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Taxonomía Y Análisis Filogenético Del Agaricales) Virginia Ramírez Cruz

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Taxonomía Y Análisis Filogenético Del Agaricales) Virginia Ramírez Cruz UNIVERSIDAD DE GUADAL.AJARA Centro Universitario de Ciencias Biológicas y Agropecuarias Taxonomía y análisis filogenético del género Psilocybe sensu lato (Fungi, Agaricales) Tesis que para obtener el grado de Doctora en Ciencias en Biosistemática, Ecología y Manejo de Recursos Naturales y Agrícolas Presenta Virginia Ramírez Cruz DIRECTORA Dra. Laura Guzmán Dávalos Zapopan, Jalisco 24 de julio de 2013 UNIVERSIDAD DE GUADALAJARA Centro Universitario de Ciencias Biológicas y Agropecuarias Doctorado en Ciencias en Biosistemática, Ecología y Manejo de Recursos Naturales y Agrícolas Taxonomía y análisis filogenético del género Psilocybe sensu lato (Fungi, Agaricales) Por Virginia Ramírez Cruz Tesis presentada como requisito parcial para obtener el grado de: Doctora en Ciencias en Biosistemática, Ecología y Manejo de Recursos Naturales y Agrícolas Aprobado por: _j JÜO 5 1 2()/:}; Fecha irectora de Tesis e integrante del Jurado Ju.Lo 5. 20\3 Fecha' _,-4\n-~ ~~ J;\l..,,aloba.::o '3 Julio 20'l3 Dra. Alma Rosa Villalobos Arámbula Fecha Asesor del Comité Particular e integrante del Jurado Dr. Gastón Guzmán Huerta Fecha Asesor del Comité P~icular e integrante del Jurado {~>i,IJ· Dr. José Luis Navarrete Heredia o Ju1,-o r;, ,J,o¡ 3 Fecha y Productos Bióticos Agradecimientos Agradezco al posgrado BEMARENA por la formación académica. Al Consejo Nacional de Ciencia y Tecnología (CONACYT) por otorgarme la beca de doctorado. Al proyecto CONACYT-SEP-2003-C02-42957, Universidad de Guadalajara y a Idea Wild por el financiamiento al proyecto de investigación y estancias académicas. A los organizadores del 9º Congreso Internacional de Micología (IMC9) por el apoyo concedido para asistir a dicho congreso. A la Dra. Laura Guzmán Dávalos por la dirección de éste trabajo, sus enseñanzas, consejos y todo el apoyo brindado durante el proyecto de investigación; además, Lau gracias por ocuparte no sólo de mi formación académica. A los miembros de mi comité tutora!: Dr. Aarón Rodríguez Contreras, Dr. Gastón Guzmán y Dra. Alma Rosa Villalobos Arámbula. Gracias por sus observaciones para mejorar el proyecto y por revisar esta tesis. Agradezco al Dr. José Luis Navarrete Heredia la revisión de este documento y sus consejos para mejorarlo. A la Dra. Alma Rosa Villalobos por permitirme trabajar en su Laboratorio de Biología Celular y Molecular, por su asesoría en los experimentos y por todo su apoyo. Al Dr. Aarón Rodríguez Contreras por permitirme trabajar en el Laboratorio de Sistemática Vegetal, para la obtención de secuencias. A Georgina Vargas Amado por el valioso tiempo que invirtió metiendo mis muestras al secuenciador y por su asesoría en las técnicas moleculares para la obtención de secuencias. Al Dr. Guzmán por permitirme llevar a cabo tres estancias en la colección de hongos del Instituto de Ecología de Xalapa para consultar literatura, especímenes y por los préstamos de ejemplares. Al Dr. Brandon Matheny por permitinne hacer una estancia en su laboratotio en la Universidad de Tennessee y por la asesoría en los análisis filogenéticos. A los curadores de los herbarios BPI, ENCB, F, FH, FMCE, IB, IBUG, K, LPS, MER, PC, PDD, PRM, QUE, TENN, WU y XAL por los préstamos de ejemplares tipo. Al Dr. Machiel Noordeloos por su asesoría en la detenninación de hongos. A mi familia, quienes aceptaron que estuviéramos separados durante este tiempo: Rosa Cruz Cayetano, Pedro Ramírez Ramírez, Esaú Ramírez Cruz, Reyna Vázquez Fructuoso y nuestro más reciente regalo, Eliezer Ramírez Vázquez; agradezco todo su apoyo y cariño. A mis abuelos por sus enseñanzas de vida y por todo su apoyo en todos los aspectos de mi vida: Enrique, Esperanza, Faustino y Cannen; ellos saben lo que es estar en equilibrio con la naturaleza y conocen más que yo a los hongos. A mi tía Celsa y a Israel Ramírez por su apoyo incondicional. A Lupis, por ser mi amiga y mi familia durante parte <le este tiempo, fue muy agradable compartir todos estos momentos. A mis amigos: Abi, Chayo, Dulce Munieta, Edgar Hernández, Edith Jiménez, Flor Ramírez Guillén, Gonzalo Juárez García, Gustavo, Irene Ramírez Gómez, Ivette, Jonathan Montoya López, Juan Antonio Ortega, Laura Escobar, Lnpis, Magaly, Mariel, Priscila y Rosa Luz, los tengo presentes en mi memoria. A mis compañeros del posgrado con quienes compartí este tiempo ya sea en el CUCBA o en el campo; Arturo, Lupis, Chuy, Gina, José Luis Villa, Fidel, Pili, Diego, Esther Esquive!, Marcelo y Luis Eduardo. Agradezco también a Guadalupe Munguía Lino, Arturo Castro-Castro y Jesús González por invertir tiempo en leer algunos manuscritos, gracias por sus observaciones y comentarios. A José Luis Villa por su ayuda en el uso de algunos programas de cómputo. A Florencia Ramírez-Guillén por todo su apoyo durante el proyecto de investigación y estancias académicas en el Instituto de Ecología de Xalapa; agradezco también su valiosa amistad y todos lo aportes que hizo en el trabajo de investigación. Al Sr. Juan Lara por su ayuda durante las estancias académicas en la colección de hongos del Instituto de Ecología de Xalapa. A Paula Santos por los trabajos compartidos y por su amistad. A María Herrera Fonseca por su ayuda durante este tiempo, ya sea en actividades del herbatio o en observaciones al microscopio. A los integrantes del laboratorio de Micología: Lau, Isela, Oli, Luis Villaseñor, Marcelo, Ale, Luis Eduardo, Mary, Diego y César, de igual f01ma a todos los que ya no están en el laboratorio pero que fueron parte de esta familia micológica. Agradezco a Mollie Harker su amistad e invaluable apoyo. A Marisol Sánchez García por su ayuda durante mi estancia en Knoxville, por su amistad y consejos para que este trabajo mejorara. A todos aquéllos que directa o indirectamente conttibuyeron en mi vida profesional o personal: Arturo Nava, Christine Braaten, Carmen Ramos, Claudia Orozco, Etel Gándara, Isaac, Juan Jesús Ruelas, Juan Manuel Hernández, Joshua Birkebak, Julio Sierra, Lupita Gómez, Mariana Orozco, Consuelo Barajas, Margarita Delgado, Marko Gómez, Vero González y Vladimir Muro Medina. Finalmente, agradezco a quienes proporcionaron fotos y especímenes para este trabajo; Alan Rockefeller, Alonso Cortés Pérez, Brandon Fogg, Brandon Matheny, Christine Braaten, Eduardo Fanti, Fidel Landeros, Guadalupe Munguía Lino, Marko Gómez, Mikael No-Line y Saúl Ruiz. 11 CONTENIDO Resumen........................................................................................... 1 Capítulo I. Introducción........................................................................ 2 Capítulo II. Ramírez-Cruz V, Guzmán G, Guzmán-Dávalos L. 2013. Type studies in Psilocybe sensu lato (Agaricales, Strophariaceae). Sydowia 65(2)... ... .. .......... 11 Capítulo Ill. Ramírez-Cruz V, Guzmán G, Villalobos-Arámbula AR, Rodríguez A, Matheny PB, Sánchez-García M, Guzmán-Dávalos L. 2013. Phylogenetic inference and trait evolution of the psychedelic mushroom genus Psilocybe sensu lato (Agaricales). Botany.... ...... .......... ... ...... .................. ... ...... ... ..... ... ...... 53 Capítulo IV. Ramírez-Cruz V, Guzmán G, Villalobos-Arámbula AR, Rodríguez A, Matheny PB, Guzmán-Dávalos L. 2013. lnfrageneric relationships of the hallucinogenic genus Psilocybe based on the rDNA ITS region. Mycologia......... .. 89 Capítulo V. Ramírez-Cruz V, Guzmán G, Villalobos-Arámbula AR, Matheny PB, Guzmán-Dávalos L. 2013. Phylogenetic inference of the genus Deconica based on ITS rDNA sequences with description of a new species with chrysocystidia. Mycologia........................................................................................ 11 O Capítulo VI. Silva PS, Ramírez-Cruz V, Cortés-Pérez A, Guzmán G, Guzmán­ Dávalos L, Silveira RMB 2013. Deconica neorhombispora (Fungi, Agaricales, Strophariaceae): combination and synonymy. Sydowia 65 (2). .. .. ...... .. ... ...... .. 128 Capítulo VII. Ramírez-Cruz V, Guzmán G, Guzmán-Dávalos L. 2012. New combinations in Deconica. Sydowia 64(2): 217-219.................................... .. 137 Capítulo VIII. Discusión general y conclusiones........................................... 141 lll Resumen El género Psilocybe sensu lato es mundialmente conocido porque a él pertenecen los hongos alucinógenos que contienen psilocibina, redescubiertos en la sierra mazateca en el estado de Oaxaca en México. Ahora se sabe que estos hongos están presentes en las ceremonias tradicionales de varios grupos étnicos en México, además de los mazatecos. Psilocybe presenta una amplia distribución en el mundo, con la mayor diversidad concentrada en los trópicos. Por mucho tiempo se creyó que el género incluía además a especies no alucinógenas; pero desde hace una década, con base en estudios filogenéticos, se sabe que son independientes, Deconica y Psilocybe sensu stricto. Las clasificaciones infragenéricas de ambos se basan en morfología y difieren de acuerdo al autor. Este trabajo representa hipótesis sobre las relaciones de parentesco en estos géneros con base en secuencias de ADN. Se presentan aquí los resultados del estudio de ejemplares tipo en Psilocybe s.l., con el fin de verificar la determinación de las especies. En los análisis filogenéticos se usaron secuencias de tres regiones de ADN (28S, 5.8S y rpbl) de Deconica y Psilocybe. Se confirma que son géneros independientes, sin una relación de grupos hermanos y se recupera su posición filogenética en relación con otros Agaricales. Deconica pertenece a la familia Strophariaceae y Psilocybe a Hymenogastraceae. Se mapearon tres caracteres de importancia taxonómica en la topología obtenida. De los tres, sólo los crisocistidios
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