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Chaenorhinum (DC.) Rchb Programa Oficial de Doctorado en Biodiversidad Interés taxonómico de los caracteres micromorfológicos en el género Chaenorhinum (DC.) Rchb. (Plantaginaceae) Doctorando: María Pilar Blasco Conesa Director: Dr. Jaime Güemes Heras, Conservador del Jardín Botánico de la Universidad de Valencia Índice 1. Introducción general………………………………………………………... 1 1.1. El género Chaenorhinum (DC.) Rchb. (Antirrhineae, Plantaginaceae)…………………………………………………………………. 2 1.2. Interés taxonómico de la micromorfología de semillas y tricomas… 13 1.3. Valor taxonómico de la morfología en la época de la hegemonía de los estudios y las filogenias moleculares......................................... 17 1.4. Bibliografía………………………………………………………………. 21 2. Metodología………………………………………………………………….. 35 2.1. Material………………………………………………………...……....... 36 2.2. Técnicas de observación…………………………………………….. 37 2.3. Terminología…………………………………………………...……..... 41 2.4. Métodos estadísticos………………………………………………...... 42 2.5. Bibliografía………………………………………………………………. 44 3. Resultados……………………………………………………………………. 48 3.1. Relaciones micromorfológicas en el grupo Chaenorhinum origanifolium: ubicación de dos táxones poco conocidos, Linaria rodriguezii Porta y Ch. origanifolium subsp. cotiellae P. Monts. & G. Monts………………………………………………………………………….. 49 3.1.1. Introducción…….………………………………………………… 49 3.1.2. Material y métodos.……………………………………...………. 52 3.1.3. Resultados………………………………………………….......... 54 3.1.4. Discusión……………………………………………………......... 65 3.1.5. Bibliografía………………………………………………….......... 83 3.1.6. Apéndice………………………………………………………….. 86 3.2. Estudio de los caracteres micromorfológicos del grupo Ch. macropodum (Boiss. & Reut.) Lange y comparación con Ch. robustum Loscos..................................................................................................... 98 3.2.1. Introducción………………………………….………………….. 98 3.2.2. Material y métodos…………………………………………….... 101 3.2.3. Resultados………………………………………...…………...... 103 3.2.4. Discusión…………………………………………………………. 112 3.2.5. Bibliografía……………………………………………..……........ 125 3.2.6. Apéndice………………………………………………………...... 129 3.3. Chaenorhinum serpentinum (Plantaginaceae), nueva especie de los afloramientos ultrabásicos del sur de la Península Ibérica………. 136 3.3.1. Introducción…………………………………………………..... 136 3.3.2. Material y métodos…………………………………………….. 138 3.3.3. Resultados……………………………………………………… 141 3.3.4. Discusión……………………………………………………….. 151 3.3.5. Clave para la identificación de Chaenorhinum serpentinum 157 3.3.6. Bibliografía……………………………………………………… 171 3.3.7. Apéndice……………………………………………………….. 177 3.4. A new gypsophilous species of Chaenorhinum (Antirrhinaceae) from the south-east of the Iberian Peninsula……………………..…….. 190 3.4.1. Introduction…………………………………………...……….. 190 3.4.2. Material and methods……………………………………….... 191 3.4.3. Results………………………………………………………….. 192 3.4.4. Discussion……………………………………………………… 198 3.4.5. Acknowledgements.............................................................. 201 3.4.6. Key for identification of Chaenorhinum gamezii…………… 201 3.4.7. References……………………………………………………… 208 3.4.8. Appendix………………………………………………...……… 212 4. Conclusiones finales………………………………………………...……... 215 5. Bibliografía……………………………………………………………………. 219 1. Introducción general Introducción 1.1. El género Chaenorhinum (DC.) Rchb. (Antirrhineae, Plantaginaceae) El género Chaenorhinum se integra actualmente en la familia Plantaginaceae, aunque históricamente se ha situado en la familia Scrophulariaceae, una familia que recientemente ha sido disgregada y cuyos géneros tradicionales han sido repartidos entre siete familias distintas del orden Lamiales (Tank et al., 2006). Durante mucho tiempo, la familia Scrophulariaceae fue un grupo indiscutible formado por, aproximadamente, 270 géneros y 5100 especies (Mabberley, 1997). Estaba caracterizado por una gran variedad morfológica y metabólica: en la forma de vida (desde hierbas anuales a árboles), en las adaptaciones nutricionales (autótrofas, hemiparásitas u holoparásitas), en los mecanismos fisiológicos (rutas metabólicas C3 y C4), en la disposición de las hojas (alternas u opuestas), en la venación de las hojas (pinnada o palmeada), en el tipo de inflorescencia (flores solitarias, inflorescencias racemosas o cimosas), en la simetría de la flor (actinomorfa o cigomorfa), en el número de lóbulos del cáliz (2, 4 ó 5), de la corola (2, 4 ó 5), de estambres (2, 4 ó 5), de estaminodios (ausentes o hasta tres), de carpelos (2 ó 3), de estigmas (1 ó 2), en el tipo de fruto (secos o carnosos; dehiscentes o indehiscentes; en cápsula septicida, esquizocarpo o baya) (Watson & Dallwitz, 2000). Sin embargo, esta familia podía diferenciarse de las familias próximas con relativa facilidad (Olmstead et al., 2001), si bien es cierto que, desde antiguo, su delimitación tradicional estaba basada más en la ausencia de los caracteres de las familias próximas que en la presencia de caracteres comunes entre los géneros asignados a la familia (Wettstein, 1891). Aun así, era relativamente sencillo caracterizar e identificar a una “Scrophulariaceae típica”. Las características que típicamente las definían eran: flores con simetría bilateral, frecuentemente tubulares; ovarios con placentación axial; primordios seminales numerosos; frutos en cápsula septicida o poricida; y semillas con endospermo (Olmstead et al., 2001). Pero, la ausencia de caracteres que definieran de forma única a la familia, planteaba la posibilidad de que las Scrophulariaceae no fueran un grupo sistemático monofilético (Olmstead & Reeves, 1995). Dos estudios de sistemática molecular independientes (Olmstead & Reeves, 1995; dePamphilis 2 Introducción et al., 1997), identificaron la naturaleza polifilética de la tradicional familia Scrophulariaceae, corroborando así las especulaciones previas sobre la naturalidad del grupo. Los estudios posteriores que han analizado la sistemática molecular en el orden Lamiales han sostenido la polifilia de la familia (Olmstead et al., 2000, 2001; Oxelman et al., 2005; Rahmanzadeh et al., 2005) y han supuesto la desintegración de la familia Scrophulariaceae (Tank et al., 2006). El resultado de las primeras reorganizaciones, basadas en los análisis filogenéticos de secuencias de DNA plastidial de los genes rbcL, ndhF y rps2, sugería que los géneros de la clásica familia podrían quedar integrados en, al menos, 5 linajes de plantas parásitas y no parásitas. Además, la ubicación de algunos de sus géneros en clados de otras familias, como Plantaginaceae, Orobanchaceae y Globulariaceae, indicaban la necesidad de una profunda reorganización de la familia (Olmstead et al., 2001). De este modo, se propuso la desintegración de las Scrophulariaceae clásicas y sus géneros quedaron repartidos en 5 familias: Scrophulariaceae s.s., Orobanchaceae, Calceolariaceae, Stilbaceae y Plantaginaceae (Olmstead & Reeves, 1995; Olmstead et al., 2001). La última revisión (Tank et al., 2006) de Scrophulariaceae s.l. ordena los géneros y especies en siete familias distintas: Scrophulariaceae s.s., formada por unas 1000 especies ordenadas en 87 géneros, distribuida principalmente por el Hemisferio Sur, aunque dos de sus géneros más numerosos (Scrophularia L. y Verbascum L.) se encuentran predominantemente en el Hemisferio Norte; Orobanchaceae, una familia de plantas hemiparásitas u holoparásitas formada por unas 1600 especies ordenadas en 80 géneros, con una amplia distribución y presente en África, sudoeste de Asia, Europa, América y Australia; Calceolariaceae, una pequeña familia con sólo cuatro géneros y unas 280 especies, presente únicamente en Sudamérica y Nueva Zelanda; Stilbaceae, distribuida por las regiones subtropicales del sur de África, con 36 especies de 11 géneros diferentes; Phrymaceae, una familia de distribución cosmopolita formada por unas 106 especies de 10 géneros diferentes, con dos centros de diversificación, uno en el norte de América y otro en Australia; Linderniaceae, formada por unas 150 especies en 15 géneros, se presenta en África, América y Asia aunque con 3 Introducción distribución principalmente neotropical; y por último, Plantaginaceae, formada por unas 1200 especies ordenadas en unos 110 géneros, sin una localización geográfica concreta, ya que muchos de sus géneros o especies tienen una distribución cosmopolita (Tank et al., 2006). En este último grupo, Plantaginaceae, quedaron reunidos muchos de los géneros más característicos de las Scrophulariaceae clásicas: Antirrhinum L., Linaria Mill., Digitalis L., Veronica L., y también Chaenorhinum (DC.) Rchb. Con ellos se unieron géneros, morfológicamente muy distintos, situados previamente en otras familias como Plantago L., Callitriche L., Hippuris L. o Hydrotriche Zucc. Estos géneros tienen características florales muy distintas de las Scrophulariaceae típicas. Esta diversidad floral se ha interpretado como una adaptación a distintos tipos de polinización (Donoghue et al., 1998; Reeves et al., 1998). La aplicación del Código de Nomenclatura Botánica llevó a la utilización para esta nueva agrupación del nombre más antiguo entre los de las familias: Plantaginaceae. Sin embargo, por motivos ajenos a la nomenclatura, principalmente por la dificultad conceptual de aceptar que las antiguas Scrophulariaceae estuvieran bajo un nombre (Plantaginaceae) formado tradicionalmente por especies con una morfología floral muy característica y muy distinta de aquellas, se hicieron diversos intentos por eludir las normas del Código y recurrir a la conservación de nombres no prioritarios, como por ejemplo Antirrhinaceae (Reveal et al., 1999)
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