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Análisis Filogenético De La Subclase Marchantiidae (Marchantiophyta): Clarificando La Relación Entre Órdenes Y Familias

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Análisis Filogenético De La Subclase Marchantiidae (Marchantiophyta): Clarificando La Relación Entre Órdenes Y Familias ANÁLISIS FILOGENÉTICO DE LA SUBCLASE MARCHANTIIDAE (MARCHANTIOPHYTA): CLARIFICANDO LA RELACIÓN ENTRE ÓRDENES Y FAMILIAS Jorge Rafael Flores Tesis realizada para optar al título de Doctor en Ciencias Biológicas Universidad Nacional de Tucumán - 2017 - 0 Universidad Nacional de Tucumán Facultad de Ciencias Naturales e Instituto Miguel Lillo ANÁLISIS FILOGENÉTICO DE LA SUBCLASE MARCHANTIIDAE (MARCHANTIOPHYTA): CLARIFICANDO LA RELACIÓN ENTRE ÓRDENES Y FAMILIAS Jorge Rafael Flores Tesis realizada para optar al título de Doctor en Ciencias Biológicas Director: Dr. Guillermo M. Suárez Director Asociado: Dr. Santiago A. Catalano -2017- Comisión de Supervisión Dr. Guillermo M. Suarez Dr. Santiago A. Catalano Ph. D. Pablo A. Goloboff Dr. Juan B. Larraín AGRADECIMIENTOS Quisiera comenzar por agradecer a mis directores, Guille Suarez y Santi Catalano, quienes siempre contribuyeron, y aún contribuyen, a mi crecimiento profesional mediante el intercambio sincero de ideas y sugerencias. A Sole, compañera y amiga de ruta, con quién tuve la fortuna de compartir viajes de campo, cursos, congresos, etc. A mis amigos colegas de La Plata, Buenos Aires, Córdoba y de la Patagonia por su buena onda en cada encuentro. No puedo dejar de mencionar a mis amigos de Tucumán; especialmente a Cristian y a Gonzalo, por su compañerismo y amistad (y, desde luego, por las “cervezeadas”). También, y no menos por ser de otra especie, a la Lara (¡mi bóxer!). Siempre acompañándome en todo momento con su alegría (aunque sospecho que, en realidad, esperaba ligar comida). En el plano personal, quiero agradecer a mis viejos y mis hermanos por su esfuerzo interminable en darme todo lo necesario para alcanzar mis metas. No puedo imaginar lo distinto que hubiese sido mi vida sin su esfuerzo. A la gente de “mis pagos”, a mis amigos, a todos ellos, a los que están y ya no están. No olvido jamás lo que aprendí de ellos, la lealtad y la coherencia. Aún seguimos siendo los mismos de siempre, no hay diploma ni títulos que lo cambien; todavía somos los de la esquina del barrio. A Vale por su compañía, su cariño y las interminables charlas reflexivas. Descubrí en ella no sólo a una gran compañera sino también a una persona apasionada y sensible. Gracias por enseñarme tanto. A todos ellos por hacerme quien soy. “[…] ven pasar árboles y pájaros e ignoran a qué sombra pertenecen […] es posible que hayan extraviado la brújula y hoy vaguen preguntando, preguntando dónde carajo queda el buen amor porque vienen del odio […]” Desaparecidos, Benedetti-Viglietti A mi tía, a su impronta. ANÁLISIS FILOGENÉTICO DE LA SUBCLASE MARCHANTIIDAE (MARCHANTIOPHYTA): CLARIFICANDO LA RELACIÓN ENTRE ÓRDENES Y FAMILIAS ÍNDICE RESUMEN ...................................................................................................................................... 8 ABSTRACT ..................................................................................................................................... 9 CAPÍTULO I - INTRODUCCIÓN ..................................................................................................... 10 Resumen – .............................................................................................................................. 11 BRIÓFITOS, ASPECTOS GENERALES ......................................................................................... 11 LA DIVISIÓN MARCHANTIOPHYTA Y LA SUBCLASE MARCHANTIIDAE (HEPÁTICAS TALOSAS COMPLEJAS) ............................................................................................................................ 16 OBJETIVOS............................................................................................................................... 20 MATERIALES Y MÉTODOS ....................................................................................................... 21 CARACTERES MORFOLÓGICOS Y ANATÓMICOS DE LA SUBCLASE MARCHANTIIDAE .......... 26 CARACTERES GAMETOFÍTICOS ............................................................................................ 27 CARACTERES ESPOROFÍTICOS .............................................................................................. 56 DEFINICIÓN DE CARACTERES .................................................................................................. 67 DISCUSIÓN Y CONCLUSIÓN .................................................................................................... 86 1 REFERENCIAS – ........................................................................................................................ 88 CAPÍTULO II - FILOGENIA ............................................................................................................ 97 Resumen – .............................................................................................................................. 98 INTRODUCCIÓN: ESTATUS FILOGENÉTICO DE MARCHANTIIDAE Y LAS CLASIFICACIONES BASADAS EN EL PRINCIPIO DE LA SISTEMÁTICA FILOGENÉTICA ........................................... 99 ESTRATEGIAS DE VIDA Y FILOGENIA .................................................................................. 102 MATERIALES Y MÉTODOS ..................................................................................................... 104 MUESTREO TAXONÓMICO Y MORFOLÓGICO ................................................................... 104 DATOS MOLECULARES ....................................................................................................... 119 ANÁLISIS FILOGENÉTICOS, BÚSQUEDAS FORZADAS Y COMPARACIONES TOPOLÓGICAS . 120 RECONSTRUCCIÓN DE CARACTERES, SINAPOMORFÍAS Y DIAGNOSIS .............................. 122 PATRONES EVOLUTIVOS DE ESTRATEGIAS DE VIDA Y PERTENENCIA TAXONÓMICA ........ 123 RESULTADOS ......................................................................................................................... 123 ÁRBOL COMBINADO BAJO PESADO DE BLOQUES (BW) Y BÚSQUEDAS FORZADAS.......... 123 ANÁLISIS PARTICIONADOS ................................................................................................ 129 ANÁLISIS DE SENSITIVIDAD (ESTABILIDAD DE GRUPOS) ................................................... 132 CONTRIBUCIÓN RELATIVA DE DATOS ................................................................................ 133 2 MAPEO DE SINAPOMORFÍAS Y EVALUACIÓN DE DIAGNOSIS ........................................... 138 MAPEO DE CARACTERES ADAPTATIVOS ............................................................................ 147 DISCUSIÓN ............................................................................................................................ 149 MONOFILIA DE GRUPOS Y ESTABILIDAD ........................................................................... 149 CONTRIBUCIÓN DE DATOS, CONFLICTO Y CONGRUENCIA ................................................ 154 SINAPOMORFÍAS Y DIAGNOSIS ......................................................................................... 158 ESTRATEGIAS DE VIDA, SIMILITUD MORFOLÓGICA Y ESCENARIOS EVOLUTIVOS ............. 163 CONCLUSIONES ..................................................................................................................... 165 REFERENCIAS - ...................................................................................................................... 166 CAPÍTULO III – CORRELACIÓN DE CARACTERES ....................................................................... 174 RESUMEN – ........................................................................................................................... 175 INTRODUCCIÓN .................................................................................................................... 176 MATERIALES Y MÉTODOS ..................................................................................................... 180 Filogenia............................................................................................................................ 180 Correlación de caracteres discretos. ................................................................................ 180 RESULTADOS ......................................................................................................................... 184 VARIABILIDAD DE CARACTERES ADAPTATIVOS ................................................................. 184 3 CORRELACIÓN FILOGENÉTICA ........................................................................................... 184 DISCUSIÓN Y CONCLUSIÓN .................................................................................................. 187 REFERENCIAS - ...................................................................................................................... 192 CAPÍTULO IV – CLASIFICACIÓN Y CAMBIOS TAXONÓMICOS .................................................. 197 Resumen – ............................................................................................................................ 198 INTRODUCCIÓN .................................................................................................................... 198 PROPUESTAS CLASIFICATORIAS EN EL MARCO DE LA SISTEMÁTICA FILOGENÉTICA ........ 198 CLASIFICACIONES PREVIAS Y ACTUAL DE MARCHANTIIDAE ............................................. 201 CLASIFICACIÓN ...................................................................................................................... 206 CLASE: MARCHANTIOPSIDA Cronquist, Takht. & W. Zimm., Taxon 15: 132 (1966) ........... 209 SUBCLASE: BLASIIDAE He-Nygrén, Juslén, Ahonen,
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