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Revisión Taxonómica De Los Géneros Phaseolus Y Vigna (Leguminosae-Papilionoideae) En Cuba

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Revisión Taxonómica De Los Géneros Phaseolus Y Vigna (Leguminosae-Papilionoideae) En Cuba View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Anales del Jardín Botánico de Madrid (Real Jardín Botánico - CSIC) Anales del Jardín Botánico de Madrid 61(2): 135-154 www.rjb.csic.es Revisión taxonómica de los géneros Phaseolus y Vigna (Leguminosae-Papilionoideae) en Cuba por Ángela Beyra & Grisel Reyes Artiles Centro de Investigaciones de Medio Ambiente de Camagüey (CIMAC). Ministerio de Ciencia, Tecnología y Medio Ambiente. Cisneros 105 / Pobre y Ángel, Camagüey, C.P. 70100, Cuba. [email protected] Resumen Abstract Se hace una revisión taxonómica de las especies de los géneros We present a taxonomic study of the Cuban species of Phaseo- Phaseolus L. y Vigna Savi presentes en Cuba. Se incluyen claves lus L. and Vigna Savi. It includes keys, descriptions, illustrations dicotómicas, descripciones, ilustraciones, datos químicos, cro- of selected taxa, chromosome numbers, palinology, phenology, mosomáticos, palinológicos, etnobotánicos, así como la distri- ecology, distribution maps, notes on chemical composition and bución y ecología de las 11 especies de ambos géneros. Se ana- ethnobotany for each of the eleven species known from Cuba. liza y se hace una estimación del valor de los caracteres que di- We also present a discussion on the value of the characters used ferencian a cada uno de los géneros y especies. in the taxonomy of the complex. Palabras clave: Cuba, Leguminosae, Papilionoideae, Phaseo- Key words: Cuba, Leguminosae, Papilionoideae, Phaseoleae, leae, Phaseolinae, Phaseolus, taxonomía, Vigna. Phaseolinae, Phaseolus, taxonomy, Vigna. Introducción un rostro más allá del estigma, lo que permitió un me- jor agrupamiento natural de los táxones de Vigna del La delimitación de los géneros Phaseolus y Vigna ha viejo mundo, y la transferencia de toda la sección asiá- sido tradicionalmente un problema difícil de solucio- tica de Phaseolus, sect. Ceratotropis (Piper) Verdc., a nar, especialmente cuando se considera también a los este género. Más tarde esta ordenación fue apoyada géneros más proximos (Verdcourt, 1970, 1980; Maré- por datos palinológicos. chal & al., 1981; Lackey, 1981, 1983; Burkill, 1995). Verdcourt (1970) hizo un primer intento serio de Además, la taxonomía del género Vigna dista mucho resolver la confusión con sus aportaciones morfológi- de ser satisfactoria (Delgado-Salinas & Torres-Colín, cas, bioquímicas, citogenéticas y palinológicas, cuyos 2001). resultados le permitieron tranferir tres secciones del En un pricipio la diferencia entre ambos géneros se género Phaseolus (sect. Sigmoidotropis Piper, sect. basaba en la morfología de la quilla, simétrica en Vig- Cochliasanthus Trev. y sect. Lasiospron Benth.), al gé- na y enrollada o curva en Phaseolus, carácter que pos- nero Vigna, y puso en duda la validez de los dos carac- teriormente se vio que no era diagnóstico. El género teres que se utilizaban para separar ambos géneros. Phaseolus incluía inicialmente un gran número de es- Posteriormente Maréchal & al. (1978), en su revisión pecies, y las secciones en las que se había dividido fue- de Vigna, consideraron especialmente importantes las ron más tarde elevadas a rango de género (Strophosty- diferencias en la morfología del polen. les Elliot, Dysolobium Prain y Macroptilium Urb.). Maréchal & al. (1981) reunieron observaciones adi- Fawcett & Rendle (1920) transfirieron Phaseolus pe- cionales sobre un buen número de táxones de Phaseo- duncularis y P. antillanus al género Vigna. Además, lus y Vigna, tales como morfología de las plántulas, Wilczek (1954) y Hepper (1958) establecieron una número cromosomático, caracteres organográficos y nueva circunscripción para Vigna basada en dos ca- palinológicos. Este volumen de información les per- racteres, estípulas alargadas y estilo prolongado por mitió realizar un estudio taxonométrico en el comple- 136 Anales del Jardín Botánico de Madrid 61(2) 2004 jo Phaseolus-Vigna y géneros próximos, en el que con- En el presente estudio se hace una evaluación de to- cluyeron que el género Phaseolus representa un grupo das las especies de Phaseolus y Vigna presentes en la separado de Vigna por varios caracteres diferenciales. flora cubana, según los criterios B y D de la UICN Propusieron un género Phaseolus restringido y ho- (2001). mogéneo basado en la presencia de pelos uncinados, brácteas persistentes y falta de nectarios extraflorales Tratamiento taxonómico (Baudet & Maréchal, 1976). En la última década se han publicado acerca de las Ambos géneros se encuadran en la subtribu Pha- Papillionoideae una serie de estudios filogenéticos ba- seolinae Bronn, que se caracteriza y diferencia de otras sados en técnicas moleculares (Bruneau & Doyle, subtribus por tener las semillas, en general, un hilo 1993; Wojciechowski & al., 1993; Sanderson & Lis- ± alargado, normalmente cubierto con los restos del ton, 1995; Kollipara & al., 1997; Falquet & al., 1997; tejido medular esponjoso funicular (Lackey, 1981). Potokina & al., 1999; Beyra & Lavin, 1999; Delgado- Además presenta adaptaciones a la entomofilia en su Salinas & al., 1999; Goel & al., 2002), que han ayuda- morfología floral. do a aclarar relaciones tanto interespecíficas como in- Número cromosomático: x = 11, con reducciones tergenéricas. Delgado-Salinas & al. (1999), con datos ocasionales a x = 10 (Lackey, 1981). de la region ITS y morfológicos, trataron sobre las re- Distribución: En el mundo incluye 23 géneros y laciones filogenéticas entre las distintas especies de aproximadamente 342 especies, la mayoría en África, Phaseolus, y Goel & al. (2002), utilizando las regiones Madagascar y Asia. Sólo seis géneros son nativos del ITS1 e ITS2 de 29 especies de Vigna y 9 especies de continente americano. Phaseolus, reconstruyen su filogenia, así como las rela- Composición química: La canavanina falta en las se- ciones con géneros próximos. millas de la subtribu Phaseolinae (Lackey, 1981). Por Delgado-Salinas & al. (1993), Vaillancourt & al. su parte, Johns (1994) determinó el contenido alelo- (1993) y Llaca & al. (1994), mediante el uso de DNA químico de semillas ricas en nitrógeno de varios géne- cloroplástico, sostienen que Phaseolus y Vigna forman ros de Phaseolinae: Lablab, Macrotyloma, Phaseolus un clado monofilético junto a Macroptilium y Stro- (5 especies), Psophocarpus y Vigna (7 especies). Agui- phostyles. Este grupo fue analizado posteriormente lar & al. (1994) estudiaron 7 táxones de Vigna y uno por Tomooka & al. (2003), que propusieron una filo- de Phaseolus útiles en los procesos de fijación de genia apoyada por los datos morfológicos y molecula- nitrógeno, y describieron la morfología del nódulo en res, además de tratar la ecogeografía de cada especie y cada una de las especies. su conservación, tanto ex situ como in situ. Usos: Una de las leguminosas más cultivadas en el Actualmente, los géneros Phaseolus y Vigna se in- mundo son los “frijoles”, especies pertenecientes a los cluyen en la tribu Phaseoleae, subtribu Phaseolinae géneros Phaseolus y Vigna, de las que se cultivan apro- (Lewis & Polhill, 1998; Delgado-Salinas & al., 1999; ximadamente treinta (Smartt, 1990). Son muy impor- Chappil, 2001; Delgado-Salinas, 2001; Delgado-Sali- tantes como alimento y forraje, y también para dar co- nas & Torres-Colín, 2001; Goel & al., 2002), que in- bertura al suelo y como fuente de abono verde. Los fri- cluye en Cuba 5 géneros y 16 especies de distribución joles y granos asiáticos, que pertenecen a Vigna subg. neotropical y 2 géneros distribuidos principalmente Ceratotropis, se consideran un componente esencial en por los paleotrópicos. la dieta de una gran parte de la población del conti- Material y métodos nente asiático, y un grupo menos conocido taxonómi- camente en comparación con los frijoles del género La presente contribución constituye una revisión Phaseolus o la soja del género Glycine (Tomooka & al., taxonómica preliminar de los géneros Phaseolus y Vig- 2003). na en el archipiélago cubano, y está basada sobre todo en el estudio de las recolecciones depositadas en los 1. Phaseolus L., Sp. Pl.: 723. 1753 herbarios cubanos y la información bibliográfica dis- Tipo: Phaseolus vulgaris L. (lectótipo, designado por ponible. Britton & Brown, 1913). La nomenclatura sobre formaciones vegetales y ti- pos de suelos es la publicada en el Nuevo Atlas Na- Hierbas anuales, plurianuales o perennes, trepado- cional de Cuba (Instituto de Geografía ACC e Institu- ras, postradas o erectas, con pelos rectos glandulares y to Cubano de Geodesia y Cartografía, 1989), en los uncinados. Raíces a veces tuberosas, frecuentemente Mapas de Vegetación Actual y Suelo, respectivamen- lignificadas, raramente fibrosas. Hojas trifolioladas te, a escala 1:1000 000. –pinnaticompuestas–; estípulas de triangulares a lan- Á. Beyra & G. Reyes: Revisión taxonómica de los géneros Phaseolus y Vigna en Cuba 137 ceoladas, raramente orbiculares, agudas, no prolon- las regiones ITS llevado a cabo por Delgado-Salinas gadas por debajo del punto de inserción, generalmen- & al. (1999) sugieren que Phaseolus es monofilético te retrorsas, a menudo pubescentes; pecíolos general- y limitado al continente americano, especialmente a mente más largos que el raquis, ambos canaliculados; las regiones tropicales y templado-cálidas de Norte- folíolos enteros o lobados, de redondeados a trunca- américa. Este clado quedó definido morfológica- dos en la base, de agudos a acuminados y mucronados mente, según los citados autores, por tener pelos un- en el ápice, el terminal ligeramente más largo que los cinados en las hojas, quilla
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