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Lankesteriana 1 LANKESTERIANA 4(1): 5-36. 2004. ESTUDIO MORFOLÓGICO DE SMILAX L. (SMILACACEAE) EN COSTA RICA, CON IMPLICACIONES SISTEMÁTICAS LILIAN FERRUFINO ACOSTA & JORGE GÓMEZ LAURITO Escuela de Biología, Universidad de Costa Rica 2060 San José, Costa Rica ABSTRACT. A morphologic revision of Costa Rican species of Smilax is presented. Traditionally, up to 14 species were accepted. In the present paper 7 species are recognized: Smilax domingensis, S. mollis, S. pana- mensis, S. spinosa, S. spissa, S. subpubescens, and S. vanilliodora. The following names are treated as synonyms: Smilax engleriana and S. kunthii of S. domingensis; S. hirsutior, S. angustiflora and S. candelariae of S. mollis, and S. chiriquensis and S. regelii var. albida of S. vanilliodora. Smilax regelii is excluded as a valid taxon and a lectotype of S. gymnopoda is designated. Dichotomous keys with vegetatives and reproduc- tive characters (flowers and fruits) are presented, on the basis of field and herbaria observations. For all the species many important characters useful for identification were included, such as rhizome, stem, size of tepals and variation in berry colour during development stages. RESUMEN. Se realizó una revisión morfológica de las especies de Smilax de Costa Rica. Tradicionalmente se han aceptado hasta 14 especies. En este trabajo se reconocen 7: Smilax domingensis, S. mollis, S. panamensis, S. spinosa, S. spissa, S. subpubescens y S. vanilliodora. Los siguientes nombres se tratan como sinónimos: Smilax engleriana y S. kunthii de S. domingensis; S. hirsutior, S. angustiflora y S. candelariae de S. mollis, y S. chiriquensis y S. regelii var. albida de S. vanilliodora. Smilax regelii se excluye como taxon válido y se designa un lectotipo de S. gymnopoda. Se elaboraron claves dicotómicas con características vegetativas y reproductivas (flores y frutos), con base en observaciones de campo y especímenes de herbario. En todas las especies se incluyeron diversos caracteres importantes para la identificación, como el rizoma, el tallo, el tamaño de los tépalos y la variación del color de las bayas a lo largo del desarrollo. PALABRAS CLAVE / KEY WORDS: Smilacaceae, Smilax, morfología, taxonomía, Costa Rica. El género Smilax consta aproximadamente de 300 semillas (Morice 1970) y un estudio monográfico de especies que habitan en zonas templadas y tropicales las especies brasileñas (Andreata 1980). Tanto estu- en ambos hemisferios (Judd et al. 2002). En dios morfológicos como análisis genéticos (ribulosa Mesoamérica se han registrado al menos 25 especies. 1,5 bifosfato carboxilasa / oxigenasa, rbcL, y las Standley (1937) comunica 14 especies en Costa Rica regiones no codificadoras de ADN del cloroplasto, y Huft (1994) registra las siguientes 13 especies: trn-L, trn-F) indican que la familia Smilacaceae es Smilax angustiflora, S. candelariae, S. chiriquensis, monofilética y pertenece al orden Liliales (Judd et al. S. domingensis, S. engleriana, S. hirsutior, S. kunthii, 1999, 2002, Rudall et al. 2000). S. mollis, S. panamensis, S. spinosa, S. spissa, S. sub- En el neotrópico se han realizado estudios ta- pubescens y S. vanilliodora. xonómicos para tratar de distinguir las especies de Durante mucho tiempo el género Smilax formó este género. Gaskin & Berry (1998) proponen una parte de la familia Liliaceae; pero en años recientes se sinonimia en Smilax de la Guayana Venezolana y el ha incluido en su propia familia, Smilacaceae uso de una combinación novedosa de caracteres, tales (Cronquist 1968, Dahlgren et al. 1985, Judd et al. como: morfología de los tallos, bases de brotes 2002). Algunos estudios recientes argumentan la se- laterales, presencia y ausencia de vainas aladas, arti- paración de Smilax de la familia Liliaceae, tales culación y longitud del pecíolo, morfología de las como: estudios de cromosomas (Vijayavalli & hojas, tipo de inflorescencia, longitud del pedicelo y Mathew 1990), composición de ácidos grasos en del receptáculo, número de flores por receptáculo y 6 LANKESTERIANA Vol. 4, Nº 1 características florales. Estos autores aceptan tres tropicales y subtropicales (Heywood 1978, Dahlgren especies y reducen nueve nombres a sinonimia. et al. 1985). Este taxon fue descrito por Ventenat (1799) con Guaglianone & Gattuso (1991) realizan una base en el género Smilax L. Este concepto fue segui- revisión morfológica y un estudio anatómico de las do por diversos autores (Endlicher 1836, Lindley estructuras vegetativas y reproductivas, así como de 1836, Grisebach 1842, Kunth 1850, De Candolle la distribución geográfica, la ecología y la fenología 1878). Otros autores (Bentham & Hooker 1880, de Smilax en Argentina, donde describen e ilustran Engler 1888, Melchior 1964) tratan Smilax en la cinco especies. Andreata (1980, 1997) también rea- familia Liliaceae. Engler (1888) y Melchior (1964) lizó un estudio exomorfológico y anatómico y una dividen Liliaceae en 13 subfamilias; entre éstas revisión de Smilax en Brasil, abarcando 12 especies, junto con una historia taxonómica del género. Smilacoideae, que se compone de cuatro géneros: Aunque se han considerado características diversas Rhipogonum Forst. & Forst. (variante ortográfica en para distinguir las especies, los resultados no han sido la literatura: Ripogonum), Smilax, Pseudosmilax los esperados. Hayata y Heterosmilax. Dahlgren et al. (1985) incluyen estos cuatro géneros en la familia Desde la época precolombina la raíz de cuculmeca Smilacaceae, que dividen en dos subfamilias: y zarzaparrilla, nombres populares de ciertas especies Smilacoideae, con tres géneros, y Rhipogonoideae de Smilax en Centro América, ha sido considerada en con el género Rhipogonum. la medicina popular contra problemas diuréticos, Además de Liliaceae o Smilacaceae, la familia se infecciones dermatológicas, desórdenes gastrointesti- ha incluido en Liliiflorae (Melchior 1964), Liliales nales, reumatismo, vaginitis, como anticonceptivo, (Hutchinson 1934, Cronquist 1968, Heywood 1978, para regulación menstrual, anemia, mordeduras de Goldberg 1989, Judd et al. 1999, 2002), Asparagales serpientes y artritis (Ocampo 1994, Gupta 1995). A (Dahlgren & Clifford 1982), Dioscoreales (Dahlgren pesar de que en la región mesoamericana se han rea- et al. 1985) y Smilacales (Takhtajan 1969). Dahlgren lizado investigaciones fitoquímicas y etnobotánicas, et al. (1985) comentan que la familia puede ser con- no se sabe con certeza cuáles son las especies uti- siderada junto a Petermanniaceae dentro de lizadas, lo que pone en evidencia una gran incer- Dioscoreales. Judd et al. (1999, 2002) incluyen en la tidumbre en la información disponible. familia Smilax y Rhipogonum, mientras que Thorne El objetivo principal de este trabajo fue mejorar las (1992) considera que Smilacaceae es una familia descripciones existentes y resolver problemas monogenérica. taxonómicos y de sinonimia en las especies de Costa Otra clasificación de Smilacaceae es la que incluye Rica, lo que facilitará la identificación de las especies tres géneros: Rhipogonum, Smilax y Heterosmilax, registradas en la región y zonas aledañas. Además, apoyada en gran parte por Melchior (1964), este estudio comprende un análisis morfológico de las Heywood (1978), Koyama (1983), Dahlgren et al. estructuras vegetativas y reproductivas, teniendo en (1985) y Andreata (1997). Pseudosmilax se reduce a cuenta la variabilidad morfológica de las hojas y el sinónimo de Heterosmilax (Koyama 1984). color de los frutos, que son caracteres que han sido Heterosmilax se distribuye en el sudeste de Asia. observados en la naturaleza y en especímenes de Este es un género muy próximo a Smilax, que se dife- herbario. rencia de éste por el perigonio soldado (urceolado) y tres estambres (raramente 9-12) generalmente unidos. Smilax es el género con mayor importancia Historia taxonómica de Smilacaceae y Smilax económica, distribuido en regiones templadas, tropi- cales y subtropicales en ambos hemisferios. Se carac- Familia Smilacaceae Ventenat teriza por un perigonio libre, seis estambres libres y La familia Smilacaceae está compuesta por los anteras confluentes (Koyama 1983, Dahlgren et al. géneros Smilax L. (300 spp.) y Heterosmilax Kunth 1985, Huft 1994). (11 spp.), que se distribuyen en regiones templadas, Otro factor que ha causado polémica es la posición Abril 2004 FERRUFINO & GÓMEZ-LAURITO - Smilax en Costa Rica 7 filogenética de Smilacaceae en relación con otras Endlicher (1836) sitúa Smilax en su propio orden LVI monocotiledóneas. Cronquist (1981) considera que Smilacaceae. Smilacaceae y Dioscoreaceae son las familias más Lindley (1836) incluye Smilax en el grupo IV especializadas de Liliales, mientras que Dahlgren et Retosae y en el orden CCLVI Smilacaceae. Este autor al. (1985) consideran que Smilacaceae es una de las sitúa Smilax en la clase V Distyogenes y el orden seis familias no basales de las monocotiledóneas. LXIX Smilaceae, al lado del género Rhipogonum. Ellos sostienen que Dioscoreaceae está muy rela- Grisebach (1842) realiza el tratamiento de 33 cionada con Smilacaceae y ésta última con especies de Smilax; divide el género en Petermanniaceae y Liliaceae. Tanto estudios mor- Pharmacosmilax y Pachysmilax por la base y la fológicos (Conran 1989) como de secuencias de ADN forma de la antera, y la consistencia y el patrón de la (Chase et al. 1995a, 1995b, 2000, Judd et al. 1999, nervadura de la hoja. Ubica Smilax en Smilacaceae, 2002, Rudall et al. 2000, Patterson & Givnish 2002) cerca de Herreria Ruiz & Pav. favorecen la ubicación de Smilacaceae en el orden Kunth (1850) enumera 188 especies de Smilax y Liliales.
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