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6Cfe01-543 2/10 6CFE01-543 2/10 Anatomía de Cupresáceas ROMÁN, E.1, GARCÍA ESTEBAN, L.1, DE PALACIOS DE PALACIOS, P.1 y GARCÍA 1 FERNÁNDEZ, F. 1 Cátedra de Tecnología de la Madera. Dpto. Ingeniería Forestal. E.T.S.I. Montes – Universidad Politécnica de Madrid. Resumen Se han descrito anatómicamente las maderas de 17 géneros monoespecíficos de la familia Cupressaceae: Taiwania, Sequoiadendron, Sequoia, Metasequoia, Cryptomeria, Glyptostrobus, Thujopsis, Fokienia, Tetraclinis, Platycladus, Microbiota, Papuacedrus, Pilgerodendron, Austrocedrus, Diselma, Fitzroya y Neocallitropsis. Para la evaluación de los clades establecidos por filogenie molecular y las agrupaciones morfológicas, se han considerado aquellos caracteres de la madera que no se encuentran influenciados por factores externos o internos. Como estudios previos manifiestan, nuestros resultados revelan un alto grado de homogeneidad entre los géneros estudiados, destacando algunas características anatómicas comunes como la presencia de extensiones de toro, verrugosidades en la cara interna de las traqueidas, la presencia de parénquima axial y el campo de cruce tipo cupressoide. La existencia de estas características, corrobora la inclusión de la familia Taxodiaceae dentro de Cupressaceae, permitiéndole formar un grupo dentro de la misma por su mayor altura de los radios leñosos y la presencia de campo de cruce tipo taxodioide. Nuestro estudio anatómico también apoya la asociación entre los géneros Fitzroya-Diselma, Fitzroya-Diselma-Pilgerodendron, Glyptostrobus-Cryptomeria, Sequoia-Sequoiadendron y Sequoia-Sequoiadendron-Metasequoia. Contrariamente, las descripciones anatómicas de los 17 géneros no presentan un resultado firme para la división entre aquellos pertenecientes al Hemisferio Norte o Sur así como otras asociaciones, debilitando los grupos. Palabras clave Madera, microscopía electrónica, microscopía óptica, filogenie. 1. Introducción La familia Cupressaceae consta de 30 géneros distribuidos por todo el mundo (FARJON, 2005), 17 de los cuales son monoespecíficos. La descripción anatómica de sus especies ha sido realizada por diversos autores (PEIRCE, 1936; 1937; PHILLIPS, 1948; GREGUSS, 1955; GAUSSEN, 1968; PANSHIN & DE ZEEUW 1980; GROMYKO, 1982; ESTEBAN et al., 2002), aunque en muchas ocasiones son contradictorias. La causa de esta falta de consenso entre los diversos autores se debe a la variabilidad que muestran algunos caracteres anatómicos según los factores ambientales y a la posición de la muestra en el árbol (JONES, 1924; BANNAN, 1941; 1944; PANSHIN & DE ZEEUW 1980; YAMAMOTO, 1992; MARTÍN et al., 2010; ESTEBAN et al., 2012). Basados en criterios morfológicos (YOUNG & WATSON 1969; ECKENWALDER, 1976; HART, 1987) o moleculares (TSUMURA et al., 1995; KUSUMI et al., 2000; GADEK et al., 2000; QUINN et al., 2002), diversos autores han realizado distintas agrupaciones entre los géneros de la familia Cupressaceae, según las afinidades entre los mismos. 3/10 2. Objetivos El objetivo de este estudio es describir anatómicamente los géneros monoespecíficos de la familia Cupressaceae, según el criterio especificado por IAWA COMMITTEE (2004), y en base al mismo, discutir las asociaciones en clades establecidas por filogenie molecular y las agrupaciones con base morfológica. 3. Metodología Las especies estudiadas, así como su procedencia y la colección de origen, se encuentran enumeradas en la Tabla 1. Las preparaciones microscópicas fueron preparadas siguiendo los métodos usuales de reblandecimiento, corte (empleándose para cortar el microtomo con grosores entre 10 – 15 µm), tinción y montaje. Las muestras fueron observadas sin teñir y teñidas con safranina y Sudan 4 para colorear la resina de rojo (JANE, 1970). La descripción anatómica se realizó siguiendo la terminología de IAWA COMMITTEE (2004). Las muestras han sido observadas utilizando microscopio óptico (Ligth Microscopy) Leica DM2500 con cámara DFC 420 y software de tratamiento de imágenes IM50 v.5 release 220, y microscopio electrónico de barrido (Scanning Electron Microscopy) SEM mod. JEOL JSM-6380. Las muestras empleadas en el microscopio electrónico fueron preparadas mediante el método descrito por HEADY & EVANS (2000). 4. Resultados En la Tabla 2 se recoge la descripción anatómica de la madera, según el criterio de IAWA COMMITTEE (2004), de los 17 géneros monoespecíficos de la familia Cupressaceae. Los caracteres que se han tenido en cuenta son aquellos que no se ven afectados por factores ambientales ni por la posición de la muestra en el árbol. Dichos caracteres son: depósitos en traqueidas (48) (Fig. 1A); extensiones de toro (58) (Fig. 1B); muescas en el borde de las punteaduras (59) (Fig. 1C); verrugosidades (60) (Fig. 1D); parénquima axial presente (72); pared transversal lisa (76) (Fig. 1E); pared transversal irregular (77) (Fig. 1F); pared transversal nodular (78) (Fig. 1G); paredes axiales nodulares (86) (Fig. 1H); paredes horizontales lisas (87) (Fig. 1I); paredes horizontales punteadas (88) (Fig. 1H); hendiduras (89) (Fig. 1J); campo de cruce tipo cupressoide (93) (Fig. 1K); campo de cruce tipo taxodioide (94) (Fig. 1L); radios de menos de 4 células de altura (102); radios entre 5 y 15 células de altura (103); radios entre 16 y 30 células de altura (104); radios de más de 30 células de altura (105). 5. Discusión La inclusión de la familia Taxodiaceae (menos Sciadopitys) dentro de Cupressaceae, se ha realizado en base a criterios moleculares (TSUMURA et al., 1995; KUSUMI et al., 2000; GADEK et al., 2000; QUINN et al., 2002) y morfológicos (YOUNG & WATSON 1969; ECKENWALDER, 1976; HART, 1987). Tal inclusión se ve reforzada por las características anatómicas observadas, como ya señalaron YOUNG & WATSON (1969), pudiéndose determinar rasgos casi generales entre todos los géneros de la familia Cupressaceae, como las 4/10 extensiones de toro (salvo en Tetraclinis (Fig. 1B), las verrugosidades en la cara interna de las traqueidas (menos en Papuacedrus y Neocallitropsis) (Fig. 1D), la presencia de parénquima axial (a excepción de Neocallitropsis) y el campo de cruce tipo cupressoide (Fig. 1K). Tabla 1. Lista de especies estudiadas con referencia a las colecciones de donde se han obtenido las muestras y su región de procedencia Número Especies Distribución geográfica Institución (Código) referencia Forest Products Laboratory (SJRw 3747) Fitzroya cupressoides 1 Chile y Argentina Maderas de Chile Universidad de (Molina) I. M. Johnston Valdivia Austrocedrus chilensis (D. Maderas de Chile Universidad de 2 Chile y Argentina Don) Pic. Serm. y Bizarri Valdivia Pilgerodendron uviferum Maderas de Chile Universidad de 3 Chile y Argentina (D. Don) Florin Valdivia Sequoia sempervirens (D. Costa Oeste de los Tervuren Xylarium, Royal Museum of 4 Don) Endl EEUU Central Africa (Tw 45912) Sequoiadendron Universidad Politécnica de Madrid, ETSI Costa Oeste de los 5 giganteum (Lindl.) J. Montes, Cátedra de Tecnología de la EEUU Buchholz madera (ETSIM X1559) Metasequoia Jodrell Laboratory, Royal Botanic 6 glyptostroboides Hu y W. China Gardens (Kew 24607 W2) C. Cheng Glyptostrobus pensilis Department of Botany, National Museum 7 (Stauton ex D. Don) K. China of Natural History, Smithsonian Koch Institution (USw 4420) Universidad Politécnica de Madrid, ETSI Cryptomeria japonica 8 Japón Montes, Cátedra de Tecnología de la (Thunb. ex L. f.) D. Don madera (ETSIM X1513) Myanmar, Taiwan, Taiwania cryptomerioides 9 Vietnam y el oeste de Universidad de Kyoto Hayata China Papuacedrus papuana (F. Nueva Guinea y las Jodrell Laboratory, Royal Botanic 10 Muell.) H. L. Li Molucas Gardens (Kew 19275 W2) Forest Products Laboratory (SJRw 11 Diselma archeri Hook. f. Tasmania 27044) Neocallitropsis pancheri Forest Products Laboratory (SJRw 12 Nueva Caledonia (Carrière) de Laub. 51802) Fokienia hodginsii (Dunn) Forest Products Laboratory (SJRw 13 China, Laos y Vietnam A. Henry y H. H. Thomas 23080) Universidad Politécnica de Madrid, ETSI Microbiota decussate 14 Rusia Montes, Cátedra de Tecnología de la Komarov madera (ETSIM X1198) Thujopsis dolabrata Jodrell Laboratory, Royal Botanic 15 (Thunb. ex L. f.) Siebold Japón Gardens (Kew 19329 W2) y Zucc. Platycladus orientalis (L.) Jodrell Laboratory, Royal Botanic 16 China, Corea y Rusia Franco Gardens (Kew 71699 W2) España, Algeria, Malta, Universidad Politécnica de Madrid, ETSI Tetraclinis articulata 17 Marruecos, Túnez y Montes, Cátedra de Tecnología de la (Vahl) Mast. Libia madera (ETSIM X1523) En el estudio de los géneros monoespecíficos del grupo Taxodiaceae (Sequoia, Sequoiadendron, Metasequoia, Glyptostrobus, Cryptomeria y Taiwania), se han observado características anatómicas que permiten diferenciarlos del resto de cupresáceas, ya que poseen 5/10 un mayor rango de altura de los radios leñosos, se observa la presencia en los radios de paredes horizontales tanto lisas como punteadas, y muchos de ellos presentan un campo de cruce tipo taxodioide, que sólo se ha observado fuera de este grupo en Papuacedrus y Microbiota (Fig. 1L). Dentro de este grupo se han descrito por caracteres morfológicos y moleculares dos clades bien resueltos: Sequoioid (Metasequoia-Sequoia- Sequoiadendron) y Taxodioid (Cryptomeria-Taxodium -Glyptostrobus) (ECKENWALDER, 1976; TSUMURA et al., 1995; GADEK et al., 2000). Anatómicamente se han observado características generales en los dos clades, pero se diferencian en una presencia generalizada de hendiduras y campo de cruce tipo taxodioide en Taxodioid, y una altura de los radios mayor en Sequoioid. Dentro del clade Sequoioid, los géneros Sequoia-Sequoiadendron son muy semejantes,
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