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Las Especies Del Género Axonopus (Poaceae: Panicoideae: Paspaleae) En Brasil*

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Las Especies Del Género Axonopus (Poaceae: Panicoideae: Paspaleae) En Brasil* BOTÁNICA LAS ESPECIES DEL GÉNERO AXONOPUS (POACEAE: PANICOIDEAE: PASPALEAE) EN BRASIL* Diego Giraldo-Cañas** RESUMEN Giraldo-Cañas, D.: Las especies del género Axonopus (Poaceae: Panicoideae: Paspaleae) en Brasil. Rev. Acad. Colomb. Cienc. 36 (140): 317-364, 2012. ISSN 0370-3908. Axonopus P. Beauv. es un género nativo del continente americano, del que se conocen unas 71 especies, princi- palmente concentradas en el norte de Sudamérica, con algunos representantes aparentemente introducidos en el Viejo Mundo. Se presenta el estudio taxonómico sinóptico de las especies brasileñas, sus sinónimos, su distri- bución geográfica y ecológica, sus nombres vernáculos, se comentan algunas particularidades morfológicas, así como relaciones morfológicas de las especies. Se discuten las afinidades morfológicas conDigitaria , Paspalum y Spheneria. Tres nombres se reducen a la sinonimia: Axonopus apricus G. A. Black [= Axonopus marginatus (Trin.) Chase ex Hitchc.], Axonopus arcuatus (Mez) G. A. Black [= Axonopus polydactylus (Steud.) Dedecca] y Axonopus obtusifolius (Raddi) Chase var. rolfsii G. A. Black [= Axonopus furcatus (Flüggé) Hitchc.]. Por otra parte, se restablecen las especies Axonopus conduplicatus G. A. Black, Axonopus schultesii G. A. Black, las que habían sido consideradas en la sinonimia de Axonopus laxiflorus (Trin.) Chase, y Axonopus eminens (Nees) G. A. Black, la cual había sido ubicada en la sinonimia de Axonopus siccus (Nees) Kuhlm. Se designa el lectotipo de Paspalum chrysodactylon (Trin.) Döll var. glabratum Döll. Cuatro especies se excluyen de la flora brasileña. Así, el género Axonopus queda representado en Brasil por 48 especies, de las cuales 16 son endémicas. Palabras clave: Axonopus, Centrochloa, Digitaria, Ophiochloa, Paspalum, Spheneria, flora de Brasil, gramí- neas neotropicales. ABSTRACT Axonopus P. Beauv. is an American genus with ca. 71 species, mainly distributed in northern South America, with some species apparently introduced to the Old World. A synoptical treatment of the Brazilian species is provided, including synonyms, ecological and geographic distribution, common names, comments on particu- lar morphological features, and morphological relationships of the species. The morphological relationships * Contribución derivada del proyecto “Estudios morfológicos, anatómicos y taxonómicos en gramíneas neotropicales”, de la Universidad Nacional de Colombia, sede Bogotá D. C. ** Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Apartado 7495, Bogotá D. C., Colombia. Correo electrónico: [email protected] 318 REV. ACAD. COLOMB. CIENC.: VOLUMEN XXXVI, NÚMERO 140 - SEPTIEMBRE 2012 with Digitaria, Paspalum, and Spheneria are discussed. Three names are synonymized: Axonopus apricus G. A. Black [= Axonopus marginatus (Trin.) Chase ex Hitchc.], Axonopus arcuatus (Mez) G. A. Black [= Axono- pus polydactylus (Steud.) Dedecca], and Axonopus obtusifolius (Raddi) Chase var. rolfsii G. A. Black [= Axo- nopus furcatus (Flüggé) Hitchc.]. Axonopus conduplicatus G. A. Black, Axonopus schultesii G. A. Black, and Axonopus eminens (Nees) G. A. Black are re-established. The name Paspalum chrysodactylon (Trin.) Döll var. glabratum Döll is lectotypified. Four species are excluded from the Brazilian flora. Thus, a total of 48Axonopus species occur in Brazil, which 16 are endemic. Key words: Axonopus, Centrochloa, Digitaria, Ophiochloa, Paspalum, Spheneria, Flora of Brazil, Neotropical grasses. Introducción de determinaciones taxonómicas equivocadas, en las que muchos botánicos (inclusive agrostólogos) confunden no La tribu Paspaleae J. Presl comprende 40 géneros y aproxi- sólo especies de Axonopus entre sí, sino que muchas de éstas madamente 581 especies, las cuales poseen un número las ubican en otros géneros (obs. pers.), tales como Digitaria cromosómico básico de x = 10, y éstas son principalmente Haller y Paspalum L., e incluso las ubican en Chloris Sw. y nativas de las Américas (Morrone et al., 2012). A esta tri- Cynodon Rich. (estos dos últimos son géneros de la subfa- bu pertenece Axonopus P. Beauv., un género nativo de las milia Chloridoideae). A ello contribuye la gran uniformidad regiones tropicales y subtropicales de América y se distri- del género Axonopus en sus caracteres morfológicos, ya que buye desde el centro-sur de los Estados Unidos de América apenas existen diferencias interespecíficas, tanto en los ór- hasta la provincia de Buenos Aires (Argentina) y Chile (sólo ganos vegetativos como en las espiguillas (Giraldo-Cañas, registrado en la Isla de Pascua) (Giraldo-Cañas, 2008a, 2000a, 2007). 2010), así como en las islas del Caribe, con algunos repre- sentantes aparentemente introducidos en África, Australia, En este contexto, en Brasil ha habido varias aproximaciones Asia tropical y Europa [Portugal (Giraldo-Cañas, 2008a) y para caracterizar la composición y la taxonomía del género Galicia (España) (Romero Buján, 2008)]. De este género se Axonopus, ya bien sea a nivel nacional o regional (véanse conocen cerca de 71 especies, las cuales están mayormente Dedecca, 1956; Black, 1963; Smith et al., 1982; da Costa concentradas en el norte de Sudamérica (Brasil, Venezuela y Monteiro & Bronzi Pace, 1984; Renvoize, 1984a, 1988; do Colombia) (Black, 1963; Giraldo-Cañas, 2008b). Sus es- Carmo Bastos, 1991; Filgueiras, 1995, 2010; Dubs, 1998; pecies crecen, principalmente, por debajo de los 1000 m de Valls et al., 2001; Zuloaga et al., 2003; Sousa da Rocha altitud, y constituyen importantes elementos en las sabanas & Secco, 2004; Quattrocchi, 2006; Dias-Melo et al., 2009; naturales, los campos, los cerrados, así como en los aflora- de Oliveira et al., 2009; Salariato et al., 2011; Filgueiras mientos rocosos de los escudos precámbricos sudamericanos & Rodrigues, 2012), pero los criterios taxonómicos son (Escudo Guayanés y Escudo Brasileño) (Giraldo-Cañas, disímiles y contradictorios, los que en muchos casos están 2007, 2008b, 2010). basados en variaciones mínimas y continuas de las dimen- siones y la densidad del indumento de hojas y espiguillas, las Axonopus incluye hierbas perennes, raras veces anuales, ces- cuales se ajustan perfectamente a variaciones locales –situa- pitosas, bajas a muy robustas, a veces rastreras, estoloníferas ción que se refleja principalmente en la aceptación de nume- o rizomatosas, de hojas tiernas a muy duras y se distingue rosos binomios o en la curiosa consideración de sinónimos de otros miembros de la tribu Paspaleae por presentar espi- sin el necesario estudio detallado del material tipo– (Tabla guillas solitarias en posición inversa, es decir, con la lema 1) (Giraldo-Cañas, 2010). Lo anterior, unido a la urgencia superior opuesta al eje del raquis y por carecer de gluma in- de desarrollar estudios taxonómicos en gramíneas neotropi- ferior. Así, las características más contundentes para delimi- cales y especialmente brasileñas –dada su alta diversidad–, tar el género Axonopus son: inflorescencias racemosas con justifican el adelantar un estudio taxonómico con el fin de espiguillas agrupadas unilateralmente, espiguillas solitarias esclarecer y delimitar las especies brasileñas de Axonopus. adaxiales, ausencia de gluma inferior y de pálea inferior y reducción del antecio basal a la lema inferior estéril (Black, Aquí presento una revisión sinóptica de las especies brasi- 1963; Giraldo-Cañas, 2007, 2008b, 2010). leñas de Axonopus, con el objetivo de contribuir con el co- nocimiento de la flora de Brasil, considerada como la más La taxonomía de Axonopus es complicada y, de hecho, resul- diversa del planeta (McNeely et al., 1990; Giulietti et al., ta difícil determinar claramente algunos ejemplares, situa- 2005; Fiaschi & Pirani, 2009, Forzza & Leitman, 2010; ción que se ve en los herbarios, dada la apreciable cantidad Giulietti, 2010; Martinelli, 2012) y especialmente de su flo- GIRALDO-CAÑAS D. - LAS ESPECIES DEL GÉNERO AXONOPUS (POACEAE: PANICOIDEAE: PASPALEAE) EN BRASIL 319 ra agrostológica, la cual asciende a 1414 especies (468 en- de nuevos sinónimos genéricos la basé en López & Mor- démicas) y 210 géneros (21 endémicos) (Longhi-Wagner, rone (2012), toda vez que dichos autores incluyeron reci- 2012), lo que la ubica como la más rica de todos los países entemente los géneros Centrochloa Swallen y Ophiochloa americanos y la segunda a nivel mundial, después de la de Filg., Davidse & Zuloaga en Axonopus, y la ubicación de China, país en el que las gramíneas están representadas por Axonopus en la tribu Paspaleae y la subtribu Paspalinae Gri- 1795 especies (810 endémicas) y 226 géneros (siete endémi- seb. la basé en Morrone et al. (2012). Cabe destacar que la cos) (Wu et al., 2006). Por otra parte, Smith & Wasshau- descripción genérica aquí presentada la construí con base en sen (1977, 1981), Burman (1985), Castro Souza (2010) y todas las especies del género y no sólo sobre la base de las Longhi-Wagner (2012) destacaron la necesidad de aportar especies brasileñas. nuevos trabajos taxonómicos para las gramíneas brasileñas, toda vez que éstos son muy escasos, máxime en los géneros La terminología de las estructuras morfológicas planas y ricamente diversificados en Brasil como lo es Axonopus, el tridimensionales la basé en la terminología clásica agros- cual ocupa el cuarto lugar en riqueza de especies (48) en la tológica, excepto en lo que tiene que ver con los términos flora agrostológica brasileña, después de Paspalum L. (202 caña (= culmo), entrenudo (= internodio), estolonífero (= especies), Panicum L. (57) y Eragrostis Wolf (52). cundidor), nervios
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