Asteraceae) En El Mediterráneo Occidental

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Asteraceae) En El Mediterráneo Occidental DEPARTAMENTO DE BOTÁNICA Estudio biosistemático del género Inula L. (Asteraceae) en el Mediterráneo occidental Memoria que para optar al grado de Doctora en Biología por la Universidad de Salamanca presenta la licenciada María Santos Vicente VºBº del Director VºBº de la Directora D. Enrique Rico Hernández Dª Mª Montserrat Martínez Ortega Salamanca, 2013 DEPARTAMENTO DE BOTÁNICA D. ENRIQUE RICO HERNÁNDEZ, Catedrático de Universidad del Departamento de Botánica de la Universidad de Salamanca y Dª Mª MONTSERRAT MARTÍNEZ ORTEGA, Profesora Titular del Departamento de Botánica de la Universidad de Salamanca, INFORMAN: En calidad de directores de la Memoria de Tesis Doctoral titulada “Estudio biosistemático del género Inula L. (Asteraceae) en el Mediterráneo occidental”, realizada por la Licenciada en Biología Dª María Santos Vicente, consideran finalizado el trabajo y autorizan la presentación, a fin de que pueda ser juzgada por el Tribunal correspondiente. Y para que así conste, firmamos el presente informe en Salamanca, a 11 de junio de 2013. D. Enrique Rico Hernández Dª Mª Montserrat Martínez Ortega Índice de contenidos CAPÍTULO 1.– Introducción ....................................................................................... 1 CAPÍTULO 2.– Beck revisited: phylogeny, karyology and character evolution in the European species of Inula L. (Asteraceae, Inuleae) and resurrection of the genus Corvisartia Mérat..................................................................................................... 19 CAPÍTULO 3.– Chromosome numbers in European species of Inula L. (Asteraceae) . 101 CAPÍTULO 4.– Caracteres micromorfológicos en taxones europeos y norteafricanos del género Inula L. (Asteraceae) y su importancia taxonómica: perspectiva filogenética 107 CAPÍTULO 5.– Nomenclatural revision of the genus Inula L. (Asteraceae) and related genera .................................................................................................................. 143 CAPÍTULO 6.– Síntesis genéricas: géneros Limbarda Adans., Corvisartia Mérat e Inula L. ...... ...................................................................................................................... 191 CAPÍTULO 7.– Conclusiones y resumen ................................................................... 259 ANEXO I.– Listado de material revisado .................................................................. 299 ANEXO II.– Especies del género Inula L. incluidas en el “Decreto de Flora Protegida de Castilla y León”: informes técnicos ......................................................................... 335 ANEXO III.– Especies del género Inula L. incluidas en “Lista Roja y Atlas y Libro Rojo de la Flora Vascular Amenazada de España”: ficha técnica ........................................... 375 ANEXO IV.– Dossier fotográfico .............................................................................. 379 Índice de contenidos (detallado) CAPÍTULO 1.– INTRODUCCIÓN .................................................................................. 1 1.1. Introducción general .................................................................................... 3 1.2. Antecedentes históricos ............................................................................... 4 1.2.1. Contexto taxonómico de Inula s.l.: la familia Asteraceae ................ 4 1.2.2. Rango taxonómico, posición filogenética y delimitación de los géneros integrantes del grupo Inula s.l. ..................................................... 6 1.2.3. Circunscripción de las especies integrantes de los géneros del grupo Inula s.l. .......................................................................................... 12 1.3. Estructura de la memoria de Memoria Doctoral ....................................... 13 1.4. Objetivos generales .................................................................................... 14 1.5. General aims ............................................................................................... 15 1.6. Bibliografía citada ....................................................................................... 16 CAPÍTULO 2.– BECK REVISITED: PHYLOGENY, KARYOLOGY AND CHARACTER EVOLUTION IN THE EUROPEAN SPECIES OF INULA L. (ASTERACEAE, INULEAE) AND RESURRECTION OF THE GENUS CORVISARTIA MÉRAT ............................................. 19 2.1. Introduction ................................................................................................ 21 2.2. Material and methods ................................................................................ 23 2.2.1. DNA extractions and ITS amplifications ......................................... 23 2.2.2. Sequence alignment and editing .................................................... 24 2.2.3. Molecular phylogenetic analysis .................................................... 24 2.2.4. Karyological analysis ....................................................................... 25 2.2.5. Character‐state evolution and taxonomic information ................. 28 2.3. Results ......................................................................................................... 29 2.3.1. Molecular phylogenetic analysis .................................................... 29 2.3.2. Karyology ........................................................................................ 32 2.3.3. Chromosomal and morphological traits evolution ......................... 36 2.4. Discussion ................................................................................................... 41 2.4.1. Paraphyly of Inula s.l. and resurrection of the genus Corvisartia .. 41 2.4.2. The “Inula clade” and infrageneric classification of Inula s.s. in Europe ....................................................................................................... 45 2.4.3. Polyploidy as a major force in the evolution of Inula ..................... 49 2.5. Acknowledgments ...................................................................................... 52 2.6. Literature cited ........................................................................................... 52 Appendix 2.1. List of taxa used in the molecular analysis ................................. 60 Appendix 2.2. Aligned ITS sequences ................................................................ 63 Appendix 2.3. Summary of published chromosome counts in Inuleae ............ 69 Appendix 2.4. Selected revised material belonging to Inula s.l. ....................... 93 Appendix 2.5. New combinations in Corvisartia ............................................... 99 CAPÍTULO 3.– CHROMOSOME NUMBERS IN EUROPEAN SPECIES OF INULA L. (ASTERACEAE) ....................................................................................................... 101 Chromosome numbers in European species of Inula L. (Asteraceae) ............. 103 Literature cited ................................................................................................ 105 CAPÍTULO 4.– CARACTERES MICROMORFOLÓGICOS EN TAXONES EUROPEOS Y NORTEAFRICANOS DEL GÉNERO INULA L. (ASTERACEAE) Y SU IMPORTANCIA TAXONÓMICA: PERSPECTIVA FILOGENÉTICA ........................................................ 107 4.1. Introducción .............................................................................................. 109 4.2. Materiales y métodos ............................................................................... 110 4.2.1. Material vegetal para el análisis filogenético ............................... 110 4.2.2. Análisis filogenético ...................................................................... 111 4.2.3. Selección de los caracteres morfológicos ..................................... 111 4.2.4. Mapeo de los caracteres morfológicos ........................................ 113 4.3. Resultados ................................................................................................ 113 4.3.1. Análisis filogenético ...................................................................... 113 4.3.2. Caracteres morfológicos: descripción y mapeo de caracteres .... 114 4.4. Discusión ................................................................................................... 125 4.4.1. Los caracteres micromorfológicos apoyan la separación del género Corvisartia ............................................................................................... 125 4.4.2. Importancia filogenética del indumento del aquenio .................. 127 4.4.3. Algunos caracteres vegetativos muestran señal filogenética ...... 128 4.4.4. Valor taxonómico de los caracteres micromorfológicos .............. 129 4.5. Agradecimientos ....................................................................................... 130 4.6. Bibliografía citada ..................................................................................... 131 Apéndice 4.1. Lista de taxones empleados en los estudios micromorfológicos mediante SEM .................................................................................................. 134 Ilustraciones ..................................................................................................... 137 CAPÍTULO 5.– NOMENCLATURAL REVISION OF THE GENUS INULA L. (ASTERACEAE) AND RELATED GENERA ......................................................................................... 143 Nomenclature of Inula L. (Asteraceae) and related genera, mainly in Europe, with several typifications and taxonomical notes ..........................................
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