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1 INTRODUCCIÓN Los Clorófitos Constituyen Uno De Los Grupos De

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1 INTRODUCCIÓN Los Clorófitos Constituyen Uno De Los Grupos De 1 INTRODUCCIÓN Los clorófitos constituyen uno de los grupos de vegetales eucariotas, que, tanto por su magnitud y belleza como por su fascinante biología han polarizado la atención de un contigente notable de investigadores. Se trata de un grupo ampliamente diversificado en las aguas continentales, el mar, los ambientes aerofíticos y la nieve. Desde el punto de vista biológico evidencian una extraordinaria diversidad de niveles de organización, innumerables estrategias adaptativas y eficaces sistemas de reproducción y multiplicación. Asímismo, los clorófitos han establecido con frecuencia relaciones de simbiosis con otros seres vivos, destacando el grupo de los hongos liquenizados o líquenes. En este caso la dependencia del hongo respecto del clorófito es máxima, ya que, sin los “productores primarios” el sistema “liquen” desaparece. A nivel ecológico, el desarrollo de los clorófitos tanto en el fitoplancton como en el fitobentos incide notablemente sobre el flujo de la materia y la energía, tanto en las aguas mesotróficas o eutróficas, como en las distróficas. Paralelamente, no hay que olvidar que los clorófitos han dado lugar a la aparición de lineas evolutivas transcendentales para la vida del planeta como son los embriófitos (briófitos y cormófitos), que se originaron a partir de una línea evolutiva derivada de las carofíceas. El ámbito geográfico que comprende esta obra abarca la península Ibérica, las islas Baleares e islas Canarias. La lista de especies se ha confeccionado a partir de la bibliografía recopilada que, prácticamente, se inicia en el siglo pasado con la obra de Colmeiro (1867). Éste registro bibliográfico no abarca el 100% de las publicaciones en que se citan clorófitos para el territorio considerado debido a la dificultad de recoger toda la bibliografía existente. Por ejemplo los trabajos antiguos de escasa difusión son difíciles de obtener. Asímismo, en las últimas décadas se ha producido un notable incremento de la actividad investigadora en limnología y ficología, generándose un número importante de trabajos difundidos en revistas nacionales e internacionales no siempre asequibles. En otros casos (tesis o tesinas) la información puede quedar prácticamente limitada a nivel local si no se publican los resultados. Por lo que se refiere a Portugal, la lista bibliográfica utilizada es insuficiente. De todas formas, el trabajo realizado es solamente un primer paso para ir recopilando, ordenando y revisando la amplia y entrópica información que se genera. Para cada taxon se menciona autor, año de descripción original y una referencia bibliográfica. Debido a la actualización taxonómica, numerosos taxones utilizados en la literatura ibérica han pasado a ser sinónimos nomenclaturales o taxonómicos. En estos casos, a continuación del taxon se citan aquellos sinónimos únicamente mencionados en la bibliografía consultada, excluyendo otros sinónimos. Paralelamente, se aporta información taxonómica en el apartado observaciones, tanto si se propone una nueva sinonimia, combinación nomenclatural o posición taxonómica de un taxon determinado. Por lo que se refiere al tratamiento taxonómico, el trabajo se ha centrado en dos niveles: 1) La ordenación sistemática de los clorófitos y 2) La actualización y revisión taxonómica de la mayor parte de taxones a nivel específico o infraespecífico. 1) La ordenación sistemática de los clorófitos es un tema realmente apasionante, sobre el que a lo largo de la historia se han venido proponiendo sistemas y más sistemas de 2 ordenación. Existen tres períodos bien diferenciados: a) La sistemática anterior a 1913- 1925, un tanto confusa; b) La obra de Pascher (“Süβwasserflora...”) realizada a principios de siglo que determina el futuro de la sistemática de clorófitos durante un largo período de tiempo y culmina con las ordenaciones de Fott y Bourrelly basadas exclusivamente en la morfología y c) El punto de inflexión introducido por Stewart et Mattox en 1975, al proponer una sistemática de clorófitos fundamentada en caracteres de la ultraestructura celular de los zoides (tipo de inserción flagelar) y la configuración mitótica a nivel de la telofase. Desde el trabajo histórico de Mattox et Stewart (l.c.) se ha ido profundizando en el conocimiento de la citología de las especies, lo cual ha contribuído a establecer una filogenia más “natural” de los grandes grupos. A partir del año 1990, con la utilización de las técnicas moleculares (especialmente del estudio de la secuencia del famoso 18S rDNA) se han obtenido datos altamente significativos, los cuales han confirmado en parte la ordenación propuesta por Stewart et Mattox (1975) así como permiten establecer una filogenia que refleja con más precisión las líneas evolutivas de los clorófitos (Melkonian et Surek, 1995; Friedl, 1997). No obstante, la idea que se extrae despues de consultar una extensa bibliografía sobre el tema es que nos encontramos al inicio de una nueva sistemática de clorófitos, en la cual los niveles de organización han perdido todo su protagonismo y se concede una importancia máxima a los aspectos señalados anteriormente. Por lo tanto, estamos asistiendo al desmantelamiento casi absoluto de los sistemas de clasificación considerados “tradicionales” y vemos como va emergiendo un nuevo sistema, en principio más relacionado con la evolución de los fila. Ante esta situación, se ha optado por una ordenación que refleje el monofiletismo de los grupos taxonómicos, teniendo en cuenta los conocimientos actuales, basados principalmente en la ultraestructura celular. Únicamente en algunos casos nos hemos inclinado por aceptar grupos basados en la información molecular como es el caso de las Trebouxiophyceae Friedl, Scenedesmaceae Oltmanns y Ankistrodesmaceae Korshikov. A nivel de clase el sistema es relativamente consistente o estable, con la excepción de las Prasinophyceae Christensen ex Silva, grupo que sigue siendo heterogéneo y polifilético. Probablemente, alguno de los órdenes actuales pasará a ser tratado a nivel de clase en un futuro próximo (Melkonian, com. pers.). Por otra parte, en las carofíceas sensu Mattox et Stewart, existen evidencias, tanto citológicas como moleculares, que indican un evidente monofiletismo o tronco común con los embriófitos. Éste conjunto es heterogéneo pero monofilético y algunos autores lo denominan “Streptophyta”. En las últimas décadas se han reunido bajo el nombre “Charophyceae” s.l. diversos órdenes cuyos representantes tienen fragmoplasto y una inserción flagelar subapical dotada de un haz de microtúbulos con una estructura en multicapa. Los resultados del análisis molecular confirman que las carofíceas s.l. constituyen un grupo monofilético pero también indican que se diferenciaron tres(?) líneas evolutivas. Por estas razones se ha preferido reutilizar los nombres “Charophyceae” s.s. para las Charales, “Conjugatophyceae” para las Zygnemales y Desmidiales y “Coleochaetophyceae” para las Klebsormidiales y Coleochaetales. De todas formas, es necesaria una solución más “estable” para las “Charophyceae” s.l., ya que si se considera que el filum o división “Streptophyta” es un grupo monofilético, una de las tres clases mencionadas anteriormente debería albergar también a los embriófitos!. En este caso las clases actuales de briófitos y cormófitos se verían reducidas al status de órden. 3 2) A nivel de los taxones específicos o infraespecíficos, el tratamiento ha sido de compromiso, ya que cada grupo taxonómico presenta un tratamiento independiente. Para ello se han utilizado las monografías clásicas que se mencionan en la bibliografía, así como la información recogida en los trabajos más recientes, especialmente numerosos en el campo de las formas coccales. Paralelamente a ésta labor de búsqueda y actualización nomenclatural, en algunos casos se ha realizado también una revisión taxonómica a partir de la descripción original y de los iconotipos. Cada uno de los taxones estudiados ha presentado una casuística determinada, que se ha tipificado de la forma siguiente: = sinónimo taxonómico. ?= sinónimo taxonómico dudoso. ≡ sinónimo nomenclatural. (H)= homónimo. * posición sistemática provisional o tentativa (sedis mutabilis). ING: Indice de Nombres Genéricos (Indice Nominum Genericorum). División CHLOROPHYTA Pascher, 1914 Clase PRASINOPHYCEAE T. Christensen ex Silva, 1980 4 Orden PSEUDOSCOURFIELDIALES Melkonian, 1990 Familia NEPHROSELMIDACEAE Skuja ex Silva, 1980 Género NEPHROSELMIS Stein, 1878 Nephroselmis olivacea Stein, 1878 Nephroselmis olivacea Stein; ETTL, 1983: 135, fig. 113 =Nephroselmis angulata (Korshikov) Skuja, 1948 ESPAÑA: Tarragona: Comín 1984. Nephroselmis minuta (N. Carter) Butcher, 1959 Nephroselmis minuta (N. Carter) Butcher; BUTCHER, 1959: 37, fig. 8 ESPAÑA: Madrid: Margalef et al. 1977. Tarragona: Comín 1984. Orden CHLORODENDRALES Fritsch, 1918 em. Moestrup et Throndsen, 1988 Familia CHLORODENDRACEAE Oltmanns, 1904 Género TETRASELMIS Stein, 1878 Tetraselmis arnoldii (Proshkina-Lavrenko) Norris et al., 1980 Tetraselmis arnoldii (Proshkina-Lavrenko) Norris et al.; ETTL, 1983: 146, fig. 121 ESPAÑA: Tarragona: Cambra 1993. Tetraselmis bolosiana (Margalef ex Margalef) Norris et al., 1980 Tetraselmis bolosiana (Margalef ex Margalef) Norris et al. ≡Platymonas bolosiana Margalef ex Margalef, 1946; MARGALEF, 1946a: 96 ESPAÑA: Barcelona: Catalán 1982; Margalef 1944, 1946a. 5 OBSERVACIONES: Segun ETTL (1983), la especie está insuficientemente descrita, aunque el material tipo fue revisado por Pascher (Margalef, 1944). Tetraselmis cordiformis (H.J. Carter) Stein, 1878 Tetraselmis cordiformis
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