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Anatomía Foliar Comparada Y Relaciones Filogenéticas De 11 Especies De Laeliinae Con Énfasis En Brassavola (Orchidaceae)

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Anatomía Foliar Comparada Y Relaciones Filogenéticas De 11 Especies De Laeliinae Con Énfasis En Brassavola (Orchidaceae) Anatomía foliar comparada y relaciones filogenéticas de 11 especies de Laeliinae con énfasis en Brassavola (Orchidaceae) Eliana Noguera-Savelli1 & Damelis Jáuregui2 1. Posgrado en Ciencias Biológicas, Unidad de Recursos Naturales, Centro de Investigación Científica de Yucatán, A.C. (CICY), Colonia Chuburná de Hidalgo, Calle 43. N°130. Mérida, Yucatán, México C.P. 97200; [email protected] 2. Universidad Central de Venezuela, Facultad de Agronomía, Instituto de Botánica Agrícola, Laboratorio de Morfoanatomía Vegetal, Prolongación Av. 19 de abril, Maracay 2101, Estado Aragua, Venezuela; [email protected] Recibido 05-I-2011. Corregido 25-I-2011. Aceptado 14-II-2011. Abstract: Comparative leaf anatomy and phylogenetic relationships of 11 species of Laeliinae with emphasis on Brassavola (Orchidaceae). Brassavola inhabits a wide altitude range and habitat types from Northern Mexico to Northern Argentina. Classification schemes in plants have normally used vegetative and floral characters, but when species are very similar, as in this genus, conflicts arise in species delimitation, and alternative methods should be applied. In this study we explored the taxonomic and phylogenetic value of the anatomical structure of leaves in Brassavola; as ingroup, seven species of Brassavola were considered, and as an outgroup Guarianthe skinneri, Laelia anceps, Rhyncholaelia digbyana and Rhyncholaelia glauca were evaluated. Leaf anatomical characters were studied in freehand cross sections of the middle portion with a light microscope. Ten vegetative anatomical characters were selected and coded for the phylogenetic analysis. Phylogenetic reconstruction was carried out under maximum parsimony using the program NONA through WinClada. Overall, Brassavola species reveal a wide variety of anatomical characters, many of them associated with xeromorphic plants: thick cuticle, hypodermis and cells of the mesophyll with spiral thickenings in the secondary wall. Moreover, mesophyll is either homogeneous or heterogeneous, often with extravascular bundles of fibers near the epidermis at both terete and flat leaves. All vascular bundles are collateral, arranged in more than one row in the mesophyll. The phylogenetic analysis did not resolve internal relationships of the genus; we obtained a polytomy, indicating that the anatomical characters by themselves have little phylogenetic value in Brassavola. We concluded that few anatomical characters are phylogenetically important; however, they would provide more support to elucidate the phylogenetic relantionships in the Orchidaceae and other plant groups if they are used in conjunction with morphological and/or molecular characters. Rev. Biol. Trop. 59 (3): 1047- 1059. Epub 2011 September 01. Key words: Cladistic analysis, leaf anatomy, orchids, Brassavola, Laelia, Guarianthe, Rhyncholaelia. Laeliinae es una subtribu neotropical que distribuye desde el norte de México hasta el comprende alrededor de 40 géneros y 1913 norte de Argentina y ocupan intervalos de dis- especies (van den Berg 2005). Géneros como: tribución altitudinal desde los 0-1000 (-1700) Brassavola R. Br., Cattleya Lindl., Guarianthe msnm. Entonces se caracteriza por presentar Dressler & W.E. Higgins y Rhyncholaelia Schl- plantas epífitas con tallos teretes, de una sola tr., son de un extraordinario valor ornamental; hoja suculenta, e inflorescencias generalmente otros como Encyclia Hook., Epidendrum L. y racemosas, de una a tres (-18) flores. Las espe- Prosthechea Knowles & Westc., constituyen cies de Brassavola son comúnmente conocidas elementos florísticos comunes del Neotrópi- como “dama de noche” por sus flores blancas, co. Brassavola (Epidendroideae, Laeliinae) se muy fragantes al anochecer. La mayoría de Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (3): 1047-1059, September 2011 1047 las plantas muestran un aspecto similar en sus estructura anatómica foliar de un grupo de características vegetativas y florales, lo cual especies de Laeliinae con énfasis en Bras- ha causado conflictos en la delimitación de savola, con el propósito de explorar carac- las especies y en los sistemas de clasificación teres de valor taxonómico y además evaluar propuestos. El número y la delimitación de las si éstos son filogenéticamente informativos especies para el género es variable de acuerdo para este género. a diversos autores, Jones (1975), consideró 18 spp; Brieger et al. (1976), incluyeron 15 spp; MATERIALES Y MÉTODOS por su parte Withner (1998) señaló 16 spp. y posteriormente en 2007 reconsideró el género En clasificaciones taxonómicas previas constituido por 19 spp. para Brassavola (Schlechter 1919, Brieger A pesar de los avances en el conocimiento et al. 1976, Withner 1998, 2007) la mayoría de la estructura anatómica de muchos represen- coinciden en la existencia de cuatro secciones. tantes de las Orchidaceae, Stern & Carlsward En este estudio se consideraron siete especies (2009) señalaron que los estudios anatómicos pertenecientes a tres de las secciones pro- exhaustivos en la subtribu Laeliinae han sido puestas por estos autores: 1. Secc. Brassavola escasos. Sin embargo, Baker (1972) caracterizó Lindl.: B. appendiculata A. Rich. & H. G. detalladamente la morfología y anatomía foliar Galeotti; 2. Secc. Cuneilabia Rchb.: B. acau- de 36 géneros y 79 taxones de las Laeliinae. Por lis Lindley & Paxton, B. nodosa (L.) Lindl. y otra parte, Oliveira & Sajo (1999) estudiaron la B. subulifolia Lindl.; 3. Secc. Conchoglossa anatomía foliar de los géneros: Prosthechea y Schltr.: B. ceboletta Rchb., B. fragrans Barb. Encyclia, además de las especies Epidendrum Rodr. y B. tuberculata Hook.; basado en revi- campestre Lindl. y E. secundum Jacq., también siones taxonómicas previas en este trabajo se pertenecientes a las Laeliinae. acepta el nombre B. appendiculata. La especie En Brassavola las investigaciones se han Brassavola appendiculata se distribuye desde centrado principalmente en dos especies B. el Norte de México hasta Nicaragua y corres- nodosa (L.) Lindl. y B. cucullata (L.) R. Br. en ponde con lo que otros autores han llamado en las cuales se ha estudiado la anatomía de sus esta zona como B. cucullata. Adicionalmente, órganos vegetativos (Abreu & Peña 1982, Prid- se analizaron cuatro especies también perte- geon et al. 1983, Porembski & Barthlott 1988, necientes a la subtribu Laeliinae: Guarianthe Stern & Carlsward 2009). Stern & Carlsward skinneri (Bateman) Dressler & W. E. Higgins, (2009) con un enfoque evolutivo, realizaron Laelia anceps Lindl., Rhyncholaelia glauca un análisis cladístico basado en caracteres (Lindl.) Schltr. y R. digbyana (Lindl.) Schl- morfo-anatómicos vegetativos de 40 especies tr. La cita de los nombres de los autores fue (incluyendo B. cucullata) de la subtribu Lae- seguida de acuerdo a Brumitt & Powell (1992). liinae, con el fin de examinar si la anatomía Los especímenes del material estudiado pro- podría proporcionar evidencia sobre la monofi- vienen de plantas recolectadas en el campo y lia del grupo. De esta manera se evaluaron seis de colecciones particulares. Muestras repre- caracteres anatómicos foliares, los resultados sentativas de cada taxón fueron depositadas en obtenidos indicaron que los caracteres vegeta- el Herbario CICY, del Centro de Investigación tivos por si solos son insuficientes para resolver Científica de Yucatán, Mérida, México, con las relaciones filogenéticas entre los taxones los siguientes registros: R. glauca: D. Angulo estudiados, pero pueden ser útiles para evaluar sn; R. digbyana: G. Carnevali et al. # 4966; B. los patrones de evolución dentro de la subtribu. subulifolia: G. Carnevali # 7503; B. ceboletta: Dado que Brassavola es un grupo monofi- G. Carnevali #7529; B. fragrans: G. Carnevali lético (van den Berg et al. 2009) y que sus #7530; B. nodosa: G. Carnevali #7531; B. plantas presentan una morfología foliar muy acaulis: G. Carnevali #7236; G. skinneri:G. similar, en el presente trabajo se examinó la Carnevali sn; Laelia anceps: G. Carnevali sn; 1048 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 59 (3): 1047-1059, September 2011 B. appendiculata: W. Cetzal # 32; B. tubercu- dérmico, fundamental y vascular, en el Labo- lata: A. Pardo sn. ratorio de Anatomía Vegetal de la Unidad de El estudio se llevó a cabo con hojas madu- Recursos Naturales del Centro de Investigación ras de plantas vivas; se tomaron fragmentos de Científica de Yucatán, A.C. (CICY), Mérida, la porción media de la hoja y se fijaron en FAA Yucatán, México. (Formaldehido 5%, Ácido acético 5%, Agua Producto de la caracterización y obser- 40% y Etanol 50%) (Sass 1958). Dichos frag- vación de la estructura anatómica foliar se mentos fueron seccionados transversalmente codificaron 10 caracteres, los mismos definie- a mano alzada. Las secciones fueron teñidas ron sus estados en cada caso (Cuadro 1). Una con azul de tolouidina acuosa (0.5%), se mon- vez codificados todos los caracteres para cada taron en glicerina (50%) y se sellaron con taxa, se procedió a construir la matriz que se esmalte transparente para uñas. Para el estudio emplearía en el análisis cladístico (Cuadro 2). epidérmico, se colocaron porciones de hoja Los caracteres codificados fueron no ordena- de aproximadamente 0.5cm en hipoclorito de dos, discretos, binarios y multiestados. Las sodio comercial (3.75%) durante tres horas, se especies: G. skinneri, L. anceps, R. glauca y R. lavaron exhaustivamente con agua destilada y digbyana se seleccionaron como grupo exter- se eliminaron los restos del mesofilo con ayuda no, esto basado
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