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Aportaciones Al Conocimiento Del Agregado Cystopteris Fragilis En España

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Aportaciones Al Conocimiento Del Agregado Cystopteris Fragilis En España Flora Montiberica 77: 43-51 (VII-2020) ISSN 1138-5952 – eISSN 1988-799X APORTACIONES AL CONOCIMIENTO DEL AGREGADO CYSTOPTERIS FRAGILIS EN ESPAÑA Francisco J. PÉREZ CARRO1 & Mª Pilar FERNÁNDEZ ARECES2 1 I.E.S. Bernardino de Escalante, Av. Duque de Ahumada, 5. 39770-Laredo (Cantabria). [email protected] 2 I.E.S. José del Campo, Barrio El Rocillo s/n. 39840-Ampuero (Cantabria). [email protected] RESUMEN: Se realiza un estudio del agregado Cystopteris fragilis y de C. alpina, buscando conocer las relaciones que se pueden establecer, en razón de macro y microcaracteres, entre los distintos tipos de morfologías generales y los diferentes niveles de ploidía encontrados (4x, 6x y 8x). La combinación de tamaño del exosporio y de las células guarda, permite distinguir los cinco taxones analizados, aunque no siempre. Para ello, hemos analizado 43 individuos repartidos por España, con un predominio del área cantábrica. Palabras clave: Pteridophyta; Cystopteris; Cystopteris fragilis; Cystopteridaceae; citología; nivel de ploidía; micromorfología; España. ABSTRACT: Contributions to the knowledge of the aggregate Cystopteris fragilis in Spain. A study of the aggregate Cystopteris fragilis and C. alpina is carried out, seeking to know the relationships that can be established, due to macro and microcharacters, between the different types of general morphologies and the different levels of ploidy found (4x, 6x and 8x). The combination of the size of the exospory and the stomatal guard cells, allows us to distinguish the five taxa analyzed, although not always. To do this, we have analyzed 43 individuals spread across Spain, with a predominance of the Cantabrian area. Keywords: Pteridophyta; Cystopteris; Cystopteris fragilis; Cystopteridaceae; cytology; ploidy level; micromorphology; Spain. INTRODUCCIÓN Desde el punto de vista genético, el grupo se constituye como un complejo poliploide, sin El género Cystopteris Bernh. es incluido en la actuali- representantes diploides en Europa y con elevados dad en la familia Cystopteridaceae (Payer) Shmakov (PPG niveles de ploidía LOVIS (1977). Así, para Europa, se ha I, 2016). ROTHFELS & al. (2012) cuestionaron las asigna- señalado que C. fragilis s.str. presenta tres niveles de ciones anteriores de Cystopteris a Dryopteridaceae, ploidía (4x, 6x y 8x), de C. diaphana se conoce el nivel Woodsiaceae o Athyriaceae basándose en análisis filoge- hexaploide, de C. alpina se ha encontrado el citotipo 6x néticos. Este género incluye de 10 a 30 especies, según mientras que C. dickieana presenta niveles de ploidía 4x que autores sean consultados. Tradicionalmente, en Eu- y 6x (MANTON, 1950; LÖVE & KJELLQVIST,1972; VIDA, ropa han sido reconocidos 4 taxones, C. fragilis (L.) 1974; VIDA & MOHAY, 1980; DOSTÁL & REICHSTEIN, Bernh., C. alpina (Lam.) Desv., C. diaphana (Bory) Blasdell, 1984; QUEIRÓS, 1991; IVANOVA & PIĘKOŚ-MIRKOWA, y C. dickieana R. Sim. 2003; KAWAKAMI & al., 2010; PTÁČEK, 2018; En la península ibérica los antecedentes en el estudio HANUŠOVÁ & al., 2019, entre otros). del complejo poliploide de Cystopteris se centran en las Interesados por el conocimiento del agregado Cystop- aportaciones de ROCHA (1982); PRADA, & SALVO (1985); teris, nos propusimos para el presente trabajo, partir del PRADA (1986), QUINTANILLA & al. (1999), entre las más conocimiento macromorfológico adquirido durante estu- destacadas. Otras aportaciones determinan el nivel de dios previos PÉREZ CARRO (1990), y buscar la ploidía de diversos especímenes (LOVE & KJELLQVIST, correspondencia entre los rasgos detectados, el nivel de 1972; QUEIRÓS, 1991 y 1997; QUEIRÓS & al., 1988). ploidía y los rasgos micromorfológicos (longitud del PRADA & SALVO (l.c.) proponen para C. fragilis exosporio y de las células guarda). Han sido excluidos subsp. fragilis var. huteri Hausm. ex Milde, el rango del presente estudio C. diaphana y C. dickieana (salvo subespecífico (≡ C. fragilis subsp. huteri (Hausm. ex un único individuo). Somos conscientes de las Milde) Grin., 1952); sobre esta identidad SHMAKOV & al. dificultades de diagnosticar taxones en el complejo que (2018: 293) realiza una nueva combinación, Cystopteris nos ocupa mediante el uso solo de caracteres huteri (Hausm. ex Milde) Shmakov, basándose en razones morfológicos, como ya señalan entre otros BLASDELL micro-macromorfológicas, así como en su estrecha (1963), VIDA (1974) o ROTHFELS & al. (2013). correspondencia con sustratos dolomíticos. Nos hemos guiado de las siguientes correspondencias Por otro lado, para España, de forma adicional, se ha entre características determinantes - taxón, a la hora de descrito un quinto taxón C. fragilis subsp. pseudoregia prefijar el reconocimiento de éstos: Rivas Martínez, T.E. Díaz, Fern.Prieto, Loidi & Penas (RIVAS Cystopteris alpina: Frondes tri(tetra)pinnatisectas. MARTÍNEZ & al, 1984), taxón que más tarde es Pinnas medias con pínnulas distales al menos pinnatipar- reconocido como especie C. pseudoregia (Rivas Martínez, tidas. Estas últimas en general de ovadas a lanceoladas. T.E. Díaz, Fern. Prieto, Loidi & Penas) Rivas Mart. (RIVAS Segmentos de las pínnulas con dientes emarginados en MARTÍNEZ in RIVAS MARTÍNEZ & al, 2002). cuyo seno terminan las nerviaciones. 43 Aportaciones al conocimiento del agregado Cystopteris fragilis en España Cystopteris fragilis subsp. fragilis: Frondes bi-tripin- búsqueda del nivel de significación ANOVA de Welch y natisectas con pecíolo castaño oscuro en grado diverso. para el contraste post hoc su prueba asociada, Games Segmentos de las pínnulas con dientes agudos, rara vez Howell. Cumpliéndose la homocedasticidad pero no truncados, en cuyo seno terminan las nerviaciones. claramente la normalidad, se consideró como preferente Cystopteris fragilis subsp. huteri (Hausm. ex Milde) el análisis mediante la prueba de Kruskal Wallis, Prado & Salvo: Frondes bi-tripinnatisectas con pecíolo asociada a las pruebas de Nemenyi, Dunn y Mann castaño oscuro en grado diverso. Lámina cubierta de Whitney exact. pelos unicelulares glandulares. Segmentos de las Para aproximarnos a los criterios que satisfacen las pínnulas con dientes agudos, rara vez truncados, en cuyo pruebas de, por un lado, ANOVA de Welch, y por otro, seno terminan las nerviaciones. en relación a la homocedasticidad, Kruskal Wallis, se Cystopteris pseudoregia: Frondes tripinnatisectas que procedió al ajuste de los datos mediante transformación muestran en las pinnas medias, pínnulas distales como de raíz cuadrada. En último término también se valoró mucho pinnatífidas. Estas últimas en general linear lan- ANOVA de un factor, con contraste por pares de Tukey ceoladas, sobre todo en el ápice de la fronde. Segmentos HSD. Todas ellas tuvieron la corrección de Bonferroni. de las pínnulas con algunos dientes subemarginados en Para el análisis citológico, el material fue fijado en cuyo seno terminan las nerviaciones. Farmer (alcohol absoluto: ácido acético glacial, 3:1) Algunas de las características señaladas se pueden durante 48 horas, entre -5 y 10°C, y posteriormente, observar en la figura 1. siguiendo las pautas indicadas por otros entre autores (MANTON: 1950), fue teñido con carmín acético al 2%. MATERIAL Y MÉTODOS Previamente a su observación, se procedió́ al El material utilizado para la realización del estudio, aplastamiento de los esporangios inmaduros teñidos, en tanto morfológico como citológico, se recoge en el apén- ácido acético (solución al 45%). dice 1 y se encuentra depositado en el herbario personal RESULTADOS Y DISCUSIÓN de los autores. Para realizar las medidas y el análisis citológico se usó Los datos de las localidades de los individuos analiza- un microscopio Leica DM 750 y cámara Leica ICC50 W dos y los resultados citogenéticos y micromorfológicos asociada al software para iPad, Leica AirLab App. se muestran en el apéndice 1. Por lo que se refiere al estudio de la longitud de las células guarda de las estomas, seguimos, básicamente las CITOGENÉTICOS pautas señaladas por BENNERT & al. (1989:313). Para Para el establecimiento del número cromosómico ello, procedimos a decolorar las pinnas medias de concreto en determinados individuos se han tenido material prensado, en solución de hipoclorito sódico dificultades, como ya ha sido también indicado por durante 24 horas, para, posteriormente, proceder a su MANTON (1950: 112) y JESSEN (1991: 31), entre otros. El deshidratación, en soluciones alcohólicas (30-96%). conteo impreciso, sin embargo, nos ha permitido Finalmente, el material fue incluido en DPX, con el fin establecer el nivel de ploidía y relacionar “a priori” una de proceder a su estudio. El tamaño se midió con un macromorfología, siguiendo los criterios marcados por aumento de 400 x. PRADA (1986); DOSTÁL & REICHSTEIN (1984); PÉREZ En el caso de la longitud del exosporio, se procedió a la CARRO (1990) con los tres niveles (4x, 6x y 8x) inclusión del contenido de esporangios maduros en DPX sin señalados para el complejo VIDA & MOHAY (1980); tratamiento alguno. La toma de medidas se realizó con un DOSTÁL & REICHSTEIN (1984). Los datos obtenidos se aumento de 1000 x, siguiendo las pautas de, entre otros muestran en el apéndice mencionado anteriormente. autores, (PTÁČEK, 2018:25). En el presente estudio no se han La figura 2 muestra los resultados del análisis de incluido individuos hibridógenos, habiéndose excluido los células madres de esporas de algunos de los especímenes. mismos, por observación de un contenido esporangial con Resultados con relación a C. dickieana un elevado número de esporas abortadas. Solo se ha podido estudiar un único individuo
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