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Estructura Floral De Dos Especies De Trachycarpeae (Arecaceae)

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Estructura Floral De Dos Especies De Trachycarpeae (Arecaceae) Estructura floral de dos especies de Trachycarpeae (Arecaceae) Lorena I. Guevara1, Damelis J. Jáuregui1 & Fred W. Stauffer2 1. Instituto de Botánica Agrícola Facultad de Agronomía, Universidad Central de Venezuela, apartado postal 4579, Maracay 2101, Aragua, Venezuela; [email protected], [email protected] 2. Conservatoire et Jardin botaniques de la Ville de Genève, Université de Genève, Ch. de l’Impératrice 1, case postale 60, 1292 Chambésy/ Geneva, Switzerland; [email protected] Recibido 26-IX-2013. Corregido 15-I-2014. Aceptado 13-II-2014. Abstract: Floral structure of two species of Trachycarpeae (Arecaceae). Copernicia and Washingtonia are two genera of the Trachycarpeae for which no subtribal classification has been proposed, mainly because of the lack of resolution in phylogenetic studies. Morphology and anatomy of flowers whithin Coryphoideae have proven useful for taxa delimitation and supporting relationships among their members. A description of the morphological and anatomical structure of flowers of C. tectorum and W. filifera is presented in order to explore reproductive characters that may clarify their classification within the subfamily and to contribute with floral biology studies. Flowers of cultivated specimens of both taxa and developing fruits of C. tectorum were fixed in FAA, dissected for morphological analysis, and parafin-embedded flowers and fruits were serially sectioned for obtaining permanent slides, using conventional techniques and safranin-fast green staining. All procedures were carried out in the Laboratory of Morpho-Anatomy, Agronomy Faculty of the Universidad Central de Venezuela (UCV). Both species have hermaphroditic flowers. C. tectorum flowers have a thick and pubescent perianth, six stamens with filaments forming a tube fused to the corolla, with rounded projections and an acute apex where the anthers are inserted. W. filifera flowers have an irregularly dentate calyx, and a shortly acuminate corolla, six stamens united by their filaments to the corolla which at the same time are briefly fused to the gynoecium. Cells with druse crystals in the staminal tube are reported for C. tectorum. Only one of the carpels of the gynoecium of C. tectorum develops at fruit stage, and a layer of abundant raphide cells forming a crustaceous endocarp in mature fruits, was found. W. filifera presents the perianth mesophyll with few layers of thick walled cells and schlerenchymatic tissue, gynoecium with apically fused carpels in the ventral region of ovary, free at the base and the apex of the style, where the ventral sutures are opened. C. tectorum has a ventral hypodermis in the petals made of large and thick walled cells, gynoecium with apically fused carpels in the ovary, free and adpressed basally, style-stigma completely fused, and stylar transmission channel absent distally. Distinct stylar canals in C. tectorum, united distally in W. filifera confirm the close relationship between these species and subtribe Livistoninae. Also, some floral morpho-anatomical similarities (e.g. fleshy calyx base and a hypodermis with thickened cell walls in petals) were found between C. tectorum and Pritchardia, supporting the affinities between both genera. Rev. Biol. Trop. 62 (3): 1137-1146. Epub 2014 September 01. Key words: anatomy, Arecaceae, Copernicia tectorum, flowers, morphology, Trachycarpeae, Washingtonia filifera. La tribu pantropical Trachycarpeae (sub- dificultándose la identificación de sus especies familia Coryphoideae) contiene géneros prin- (Henderson, Galeano, & Bernal, 1995). En el cipalmente neotropicales que no han sido último análisis filogenético para las subfami- ubicados en subtribus por la carencia de reso- lias de Arecaceae, Baker et al. (2009) utiliza- lución en los estudios filogenéticos (Drans- ron gran cantidad de evidencia molecular y field et al., 2008). Dos de estos géneros, morfológica, y no lograron resolver dos de las Copernicia Mart. ex Endl. y Washingtonia H. tribus más grandes, Areceae (Arecoideae) y Wendl., poseen una taxonomía complicada, Trachycarpeae (Coryphoideae), señalando que Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 62 (3): 1137-1146, September 2014 1137 esto se puede alcanzar reuniendo más datos de en los llanos centrales venezolanos y orientales diversa índole sobre los géneros. En cuanto a colombianos; formando además en Venezuela, las posibles relaciones entre estos géneros, el comunidades en ambas riberas del Lago de estudio filogenético más reciente de Trachy- Maracaibo (Huber & Oliveira-Miranda, 2010). carpeae, confirma la cercanía en las relaciones Washingtonia filifera (Linden ex André) H. entre Pritchardia, Corpernicia y Washingtonia Wendl. ex de Bari (palma del desierto, o palma (Bacon, Baker, & Simmons, 2012). de abanico de California) es plantada amplia- En la familia Arecaceae son conoci- mente como ornamental en zonas tropicales y dos los trabajos morfológicos y anatómicos subtropicales del mundo y se distribuye desde de desarrollo floral con fines taxonómicos el sureste de California y oeste de Arizona en (Baker, Dransfield, Harley, & Bruneau, 1999; Estados Unidos hasta el noroeste de México Kahn & Gluchy, 2002; Stauffer, Rutishauser, (Henderson et al., 1995). A pesar de la utili- & Endress, 2002; Stauffer & Endress, 2003; dad que muestran los estudios mencionados, Rudall, Abranson, Dransfield, & Baker, 2003). las investigaciones detalladas sobre la estruc- Dentro de la subfamilia Coryphoideae se ha tura morfológica y anatómica de flores en utilizado la morfo-anatomía de los órganos Coryphoideae siguen siendo escasas, por lo que reproductivos para explicar entre otras cosas, en este trabajo se presenta la descripción de la relaciones filogenéticas y aspectos de la bio- morfo-anatomía de las flores de C. tectorum y logía reproductiva. Esto se ve reflejado en W. filifera con la finalidad de: (1) aportar infor- el trabajo de Uhl, Morrow, y Moore (1969) mación para la sistemática de las Coryphoi- quienes describieron la anatomía de flores de deae, en especial para la ubicación de estos dos Rhapis excelsa (Thunb.) Henry, determinando géneros dentro de subtribus en Trachycarpeae y la placentación laminar del óvulo, relacionada (2) contribuir con información básica para los con la derivación múltiple de trazas hacia el estudios de biología floral en esta subfamilia. óvulo desde la región dorsal. En el gineceo de las Coryphoideae, Uhl y Moore (1971) realiza- MATERIALES Y MÉTODOS ron dos agrupaciones de géneros unicarpelares basados en la extensión de los haces vasculares Se extrajeron inflorescencias de especi- ventrales estigmáticos. Giddey, Spichiger, y menes cultivados en el Jardín Botánico Uni- Stauffer (2009) sugirieron la división de las versitario de Maracay, Facultad de Agronomía, especies de Rhapis en dos grupos basados Universidad Central de Venezuela (FAGRO- en caracteres de los filamentos estaminales y UCV), de Copernicia tectorum (Guevara & resaltaron la necesidad de nuevas investigacio- Lattke # 95) y Washingtonia filifera (Guevara nes sobre las flores de algunos géneros dentro et al. # 96) en diciembre de 2007; las exsicca- de Coryphoideae para lograr comparaciones tae fueron depositadas en el Herbario Víctor más detalladas. Castaño, Crèvecoeur, y Stau- Manuel Badillo de la FAGRO-UCV (MY) y ffer (2009) determinaron que los tres carpelos el Herbario Nacional de la Fundación Instituto en el gineceo de Sabal palmetto (Walter) Lodd. Botánico de Venezuela (VEN). ex Schult. estan libres en la base, apoyando las La identificación de las especies se basó en relaciones entre Sabal y Cryosophileae. Las las obras de Henderson et al. (1995) y Drans- relaciones internas de esta tribu fueron estudia- field et al. (2008). De C. tectorum se recolec- das en detalle por Castaño, Crèvecoeur, Pin- taron además, frutos en desarrollo. El material taud, y Stauffer (2011). Rudall, Ryder, y Baker fue fijado en FAA (5% formaldehído puro, 5% (2011) determinaron cuatro tipos de gineceo ácido acético puro, 90% etanol 70%). Se regis- dentro de las Coryphoideae según el número y traron mediciones de las estructuras florales grado de fusión de sus carpelos. (longitud del pedicelo y cáliz; longitud y ancho Copernicia tectorum (Kunth) Mart. de los botones florales preantéticos, filamentos, (palma llanera) es un elemento sobresaliente anteras y gineceo a nivel del ovario), utilizando 1138 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 62 (3): 1137-1146, September 2014 15 flores por especie como se ha utilizado en el fueron observadas y analizadas en un micros- estudio de la estructura y desarrollo de inflores- copio óptico Nikon Eclipse E 200, realizán- cencias en palmas (Ortega-Chávez & Stauffer, dose descripciones de los tejidos epidérmico, 2011). Para estas mediciones las flores fueron fundamental y vascular desde la base hasta el diseccionadas, separándose las partes bajo un ápice de cada estructura floral. Se obtuvieron microscopio estereoscópico. Para las partes de fotografías mediante una cámara digital Evolu- mayor tamaño, se utilizó una regla graduada y tion LC adaptada al microscopio. Estos proce- para las de menor tamaño, una micro regla con dimientos fueron realizados en el Laboratorio 1división=0.1mm (#TDI MRM100H - Micro de Morfoanatomía Vegetal FAGRO-UCV. Ruler), previamente utilizada en mediciones morfológicas (Salazar-Vallejo & Buzhinskaja, RESULTADOS 2013). Estos implementos se colocaron de manera paralela a cada una de las partes flora- C. tectorum: Flores hermafroditas, 1.5- les para tomar las medidas en milímetros.
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