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Zamiaceae) En La Sierra Madre Oriental, México: Un Enfoque Para Su Conservación

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Zamiaceae) En La Sierra Madre Oriental, México: Un Enfoque Para Su Conservación UNIVERSIDAD VERACRUZANA CENTRO DE INVESTIGACIONES TROPICALES TAXONOMÍA Y EVOLUCIÓN DE CERATOZAMIA (ZAMIACEAE) EN LA SIERRA MADRE ORIENTAL, MÉXICO: UN ENFOQUE PARA SU CONSERVACIÓN TESIS QUE PARA OBTENER EL GRADO DE MAESTRÍA EN ECOLOGÍA TROPICAL PRESENTA LILÍ MARTÍNEZ DOMÍNGUEZ CO-DIRECCIÓN Dr. FERNANDO NICOLALDE-MOREJÓN Dr. DENNIS WM. STEVENSON Xalapa, Veracruz, México, Marzo 2018 II III A mis padres: Lilí Domínguez Mendiola, Agustín Martínez Casanova y en especial a Rosa Martínez Casanova IV AGRADECIMIENTOS A mis dos hermanos, Guadalupe Hernández Martínez y Agustín Martínez Domínguez. A mi familia. Un agradecimiento especial a Fernando Nicolalde Morejón por ser mi principal sparring intelectual. A los siguientes investigadores por todo su apoyo durante este proyecto: Octavio Rojas Soto, Francisco Vergara Silva, Quiyari J. Santiago Jiménez y en especial a Dennis Wm. Stevenson por su apoyo y, compartir su pasión por las cícadas y la taxonomía. A los sinodales: Francisco Vergara Silva, Citlalli López Binnqüist y Noé Velázquez Rosas por sus comentarios y sugerencias para finalizar esta tesis. A Anette Méndez Durán por su apoyo en el procesamiento técnico del material botánico colectado y su valiosa amistad. A mis amigos: Leonardo Cuevas Reyes por compartir esta historia académica en el herbario; Eduardo de la Garza Pérez, Jesus Cilias Arellano, Maribel Cantoral Castro y Jesica Celaya Tejeda por compartir esta amistad de vida. A Tania Escobar por su apoyo en el trabajo de campo en la Sierra Norte de Puebla y su amistad. A mi amigo Juan Hernández por las ilustraciones botánicas. A CONACyT por la beca de maestría (No. 291047) y el financiamiento de este trabajo de tesis a través del proyecto No. 134960 otorgado al Dr. Fernando Nicolalde Morejón. Al Instituto de Investigaciones Biológicas que a través de la gestión de Alvar González Christen financió parte de este proyecto. Al herbario CIB por el material para procesar todas las colectas botánicas. Al Laboratorio de Taxonomía Integrativa. V ÍNDICE Resumen .............................................................................................................................. 16 Introducción general .......................................................................................................... 17 Historia taxonómica ......................................................................................................... 19 Integrando evidencia para la conservación ..................................................................... 20 Breve descripción metodológica ...................................................................................... 22 Preguntas de investigación ................................................................................................. 25 Hipótesis .............................................................................................................................. 25 Objetivos generales ............................................................................................................. 26 Objetivos particulares ........................................................................................................ 26 CAPÍTULO I ...................................................................................................................... 28 Filogenia del género Ceratozamia (Zamiaceae): integrando evidencia morfológica y molecular para reconstruir escenarios evolutivos ........................................................... 28 1.1 Resumen ......................................................................................................................... 29 1.2 Introducción .................................................................................................................... 30 1.3 Materiales y Métodos ..................................................................................................... 32 1.3.1 Muestreo de material biológico ............................................................................... 32 1.3.2 Codificación de caracteres morfológicos ................................................................ 32 1.3.3 Secuencias de DNA .................................................................................................. 36 1.3.4 Extracción de DNA genómico, amplificación de PCR y secuenciación.................. 36 1.3.5 Construcción de matrices ........................................................................................ 37 1.3.6 Análisis filogenéticos ............................................................................................... 37 1.3.7 Análisis de sensibilidad ........................................................................................... 38 1.3.8 Análisis de similitud ecológica ................................................................................ 38 1.4 Resultados ....................................................................................................................... 39 1.4.1 Particiones Individuales .......................................................................................... 39 VI 1.4.2 Análisis combinados: matriz cualitativa y cuantitativa .......................................... 41 1.4.3 Análisis combinados: matriz cualitativa, cuantitativa y molecular ........................ 42 1.4.4 Aporte de caracteres ................................................................................................ 42 1.4.5 Polimorfismos .......................................................................................................... 47 1.4.6 Análisis de similitud ecológica ................................................................................ 48 1.5 Discusión ........................................................................................................................ 48 1.5.1 Aporte de caracteres en la filogenia........................................................................ 48 1.5.2 Polimorfismos y homoplasia ................................................................................... 50 1.5.3 Escenarios evolutivos .............................................................................................. 52 1.6 Referencias ..................................................................................................................... 54 CAPÍTULO II ..................................................................................................................... 61 A new species in Ceratozamia (Zamiaceae) from Sierra Norte de Puebla, Mexico ...... 61 2.1 Abstract ........................................................................................................................... 62 2.2 Introduction .................................................................................................................... 63 2.3 Ceratozamia totonacorum ............................................................................................... 64 2.4 Key to species closely related to C. totonacorum .......................................................... 76 2.5 Literature cited ................................................................................................................ 76 CAPÍTULO III ................................................................................................................... 79 Qualitative and quantitative morphological evidence for recognition of a new species within Ceratozamia (Zamiaceae) from Mexico ................................................................ 79 3.1 Abstract ........................................................................................................................... 80 3.2 Introduction .................................................................................................................... 81 3.3 Materials and Methods ................................................................................................... 82 3.4 Results ............................................................................................................................ 84 3.5 Taxonomic treatment ...................................................................................................... 91 3.6 Discussion ....................................................................................................................... 93 VII 3.7 Key to northern species of Sierra Madre Oriental .......................................................... 98 3.8 References ...................................................................................................................... 98 CAPÍTULO IV .................................................................................................................. 102 Taxonomic review of Ceratozamia (Zamiaceae) in the Sierra Madre Oriental, Mexico, with notes on the history of taxonomic concepts of species .......................................... 102 4.1 Abstract ......................................................................................................................... 103 4.2 Introduction .................................................................................................................. 104 4.3 Materials and Methods ................................................................................................. 107 4.4 Results .......................................................................................................................... 107 4.4.1 Vegetative morphology. ........................................................................................
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