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Ceratophyllales Y Eudicotiledóneas Basales

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Ceratophyllales Y Eudicotiledóneas Basales UNNE FACENA CERATOPHYLLALES Y EUDICOTILEDÓNEAS BASALES DIVERSIDAD VEGETAL BIOTAXONOMIA DE SPERMATOFITOS 2010 Universidad Nacional del Nordeste Facultad de Ciencias Exactas, Naturales y Agrimensura. Corrientes Argentina PROLOGO El presente trabajo es un aporte de la Asignatura Diversidad Vegetal, de la Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) que tiene por finalidad facilitar la preparación del programa a los alumnos que cursan la materia. Se intenta brindar una orientación en el estudio de la exomorfología, de la biología reproductiva, de la distribución e importancia económica de los diferentes grupos de Angiospermas. Además, se introducen aspectos filogenéticos como resultado de estudios moleculares recientes. Esta labor es el resultado de la tarea conjunta de los siguientes docentes y adscriptos de la Asignatura: 9 Dirección y corrección: Dra. Elsa L. Cabral (Prof. Titular). 9 Compaginación general: Dra. Sylvina L. Casco. 9 Fotos: Elsa L. Cabral, Walter A. Medina, Roberto M. Salas, Andrea A. Cabaña Fader. 9 Elaboración original y revisión bibliográfica de los taxones que actualmente pertenecen al Clado Eudicotiledóneas basales: Dra. Sylvina L. Casco, Prof. Natalia Ayala, Dra. Cinthia González. 9 Actualización del Clado y edición de nuevas familias de acuerdo al Sistema de Clasificación APG II (Angiosperm Phylogenetic Group II) 2009/2010: Lic. Andrea A. Cabaña Fader; Lic. Sandra G. Martin. Este trabajo no es una fuente completa de información y se sugiere al alumno la consulta bibliográfica que se añade al final de cada tema. INDICE Página 4.1. Orden Ceraphyllales 1 4.1.1 Familia Ceratophylaceae 2 4.2 Filogenia de Las Eudicotiledoneas 6 4.3. Características de los integrantes 7 4.3.1. Orden Ranunculales 9 4.3.1.1. Familia Papaveraceae 10 4.3.1.2. Familia Menispermaceae 17 4.3.1.3. Familia Berberidaceae 21 4.3.1.4. Familia Ranunculaceae 27 4.3.2. Orden Proteales 35 4.3.2.1. Familia Nelumbonaceae 36 4.3.2.2. Familia Platanaceae 39 4.3.2.3. Familia Proteaceae 43 Ceratophyllales, grupo hermano de las Eudicotiledóneas En esta guía se tratará a Ceratophyllales como grupo hermano de las Eudicotiledóneas, teniendo en cuenta los estudios realizados por varios autores (Bremer et al. 2003; Hilu et al., 2003; Qiu et al., 1999; Soltis et al., 1997, 1999, 2005; Souza et al. 2005; Svolainen et al., 2000). Bremer et al. 2003 _____________________________________________________________________________ Diversidad Vegetal. Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) Ceratophyllales 1 4.1. Ceratophyllales Este orden comprende a una única familia, Ceratophyllaceae, con un sólo género, Ceratophyllum. De acuerdo a Cronquist (1981), esta familia aparece asociada a Nymphaeaceae, sin embargo, a partir de los años 90, la posición filogenética de Ceratophyllaceae ha sido motivo de gran interés. Como resultado existen numerosos trabajos de filogenia que revelan una gran disparidad en las posiciones filogenéticas resultantes de Ceratophyllum, demostrando en algunos casos ser de muy débil soporte. Dicha posición filogenética depende del número y del tipo de genes que se utilicen (e.g. si son regiones genéticas más conservadas o no). Este grupo, de acuerdo a diferentes autores, ha sido ubicado como taxón hermano del gran linaje de Eudicotiledóneas (Bremer et al. 2003; Hilu et al., 2003; Qiu et al., 1999; Soltis et al., 1997, 1999, 2005; Souza et al. 2005; Svolainen et al., 2000), como grupo hermano de Monocotiledóneas (Qiu et al., 1999; Soltis et al., 1997, 2005; Svolainen et al., 2000) o bien como grupo hermano de todas las plantas con flores (Les et al., 1991; Chase et al., 1993; Qiu et al., 1993). El tipo polen inaperturado de las especies de Ceratophyllum nos plantea una pregunta desde el punto de vista filogenético, si este derivó del tipo tricolpado (típico de Eudicotiledóneas) o bien es resultado de un antiguo linaje de plantas acuáticas (adquisición temprana). Por otro lado, Ceratophyllum tiene características morfológicas muy derivadas, comunes a ciertas plantas acuáticas, lo que aparentemente refleja cierto paralelismo con algunas monocotiledóneas inaperturadas (cfr. Alismatales) (Stevens, 2001). Algunos de los primeros trabajos revelaron a partir del análisis de una secuencia de rbcL que podría ser el clado hermano de las demás Angiospermas (Les et al., 1991; Chase et al., 1993; Qiu et al., 1993). Hilu et al. (2003) y Bremer et al. (2003) sugirieron que Ceratophyllales debía ser considerado como clado hermano de las Eudicotiledóneas. Análisis un tanto más recientes de atpB y 18S rADN (Soltis et al., 1997, 2005; Svolainen et al., 2000), de secuencias de genes nucleares, mitocondriales y plastidiales (Qiu et al., 1999, en Soltis et al., 2005) indican una posición alternativa de las Ceratophyllaceae, considerándolas grupo hermano de las Monocotiledóneas. Soltis et al. (1999), propone una politomía en la participación de este taxón y de los demás grupos basales, aunque con bajo soporte. Por lo tanto, queda en evidencia que a la luz de los avances todavía es un grupo en vías de resolución. Diversidad Vegetal Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) Ceratophyllales: Ceratophylaceae 2 4.1. 1. Ceratophyllaceae 4.1.1.1.a. Características ¾ Porte: plantas acuáticas, herbáceas, sumergidas, muy ramificadas y articuladas. ¾ Hojas: sésiles, verticiladas, divididas dicotómicamente. ¾ Flores: actinomorfas, monoicas, sésiles, axilares, solitarias, verdosas, rodeadas por un involucro compuesto de 10-12 bractéolas calicoides. ¾ Perianto: sépalos 3, petaloideos, libres, oblongos. ¾ Estambres: 12-20, anteras sésiles o subsésiles, extrorsas, bitecas, apendiculadas superiormente. ¾ Gineceo: súpero, unilocular, uniovulado, estilo simple, estigma oblicuo. ¾ Fruto: núcula o aquenio con el estilo persistente y un par de espinas basales. ¾ Semilla: con testa delgada, sin perisperma ni endosperma. Ceratophyllum demersum Estilo Estilo persistente Anteras con Flores y frutos de expansión del Detalle de la flor conectivo en estaminada forma de aguijón Detalle de la flor Detalle del fruto: núcula pistilada Espinas marginales 4.1.1.1.b. Biología floral y/o Fenología La polinización de esta familia es hidrófila, ocurriendo debajo del agua. Se inicia con la formación de burbujas de aire que llenan el aerénquima de los estambres maduros, provocando la ruptura de las anteras y la liberación del polen en el agua. La dehiscencia de los estambres ocurre cuando todavía están fijos a la planta o después de la degeneración de las células basales. Los estambres flotantes ascienden y se abren para llevar el agua con polen, el cual se hunde y llega a los estambres abiertos. Florece en verano. Se multiplica por división (Kubitski, 1993). 4.1.1.1.c. Distribución y Habitat Esta familia está ampliamente distribuida. Viven sumergidas o flotando en lagos, lagunas o ríos de corriente muy lenta, en zonas templadas y tropicales. Ecológicamente tienen importancia por ser lugar de puesta y refugio de numerosas especies de peces, pero se las considera una molestia al interferir el flujo de agua en canales de riego, la pesca, la natación, la navegación y otros recreos acuáticos, además de servir de hábitat idóneo a las poblaciones de larvas de mosquitos (con riesgo palúdico) y cobijar los caracoles acuáticos transmisores de la esquistosomiasis. En grandes cantidades han bloqueado el funcionamiento de centrales hidroeléctricas. Diversidad Vegetal Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) Ceratophyllales: Ceratophylaceae 3 http://www.mobot.org/MOBOT/Re search/APweb/welcome.html 4.1.1.1.d. Especies de la familia Ceratophyllaceae De acuerdo a Zuloaga y Morrone (1999) y Zuloaga, Morrone y Belgrano (2008), esta familia posee un sólo género con dos especies, que se encuentran en Argentina: Ceratophyllum demersum L. y C. muricatum subsp. australe (Griseb.). Distribución Nombre Vulgar Especies nativas Ceratophyllum demersum L Buenos aires, Chaco, Córdoba, Corrientes, Cola de zorro (Fig. 1) Entre Rios, Río Negro y Santa Fe. C. muricatum subsp. Chaco, Entre Rios, Salta, Santa Fe y australe (Griseb.) Tucumán. 4.1.1.1.e. Importancia Se utilizan como ornamentales en estanques y acuarios y como protección para peces (Lahitte et al., 1997). 4.1.1.1.f. Ilustraciones Fig. 1: Ceratophyllum demersum a. Detalle de la planta Judd et al., 1999 c. Detalle del fruto inmaduro b. Detalle de la flor d. Detalle del fruto maduro, aquenio http://botany.cs.tamu.edu/FLORA/ Diversidad Vegetal Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) Ceratophyllales: Ceratophylaceae 4 Fig. 1: Ceratophyllum muricatum subsp. asustrale b. Detalle de las hojas divididas a. Detalle de una rama c. Detalle del fruto Fotos: R. Salas y A. Cabaña Fader 1.2.4.1.g. Bibliografía -APG II. The Angiosperm Phylogenetic Group. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141 (4): 399–436. -Bremer, K., B. Bremer y M. Thulin. 2003. Introduction to Phylogeny and Systematics of Flowering Plants. Deparment of Systematic Botany Evolutionary Biology Centre. Uppsala University. USA. -Burkart, A. 1987. Flora Ilustrada de Entre Ríos (Argentina). III: Dicotiledóneas Arquiclamídeas: A. Salicales a Rosales (incluso Leguminosas). Colección Científica del I.N.T.A. VI. Buenos Aires, Argentina. 763 p. -Chase, M.W., D.E. Soltis, R.G. Olmstead, D. Morgan, D.H. Les, B.D. Mishler, M.R. Duvall, R.A. Price, H.G. Hills, Y.-L. Qiu, K.A. Kron, J.H. Rettig, E. Conti, J.D. Palmer, J.R. Manhart, K.J. Sytsma, H.J. Michaels, W.J. Kress, K.G. Karol, W.D. Clark, M. Hedrén, B.S. Gaut, R.K. Jansen, Kim K.-J., C.F. Wimpee, J.F. Smith, G.R. Furnier, S.H. Strauss, Q.-Y. Xiang, G.M. Plunkett, P.S. Soltis, S.M. Swensen, S.E. Williams, P.A. Gadek, C.J. Quinn, L.E. Eguiarte, E. Golenberg, G.H. Learn Jr., S.W. Graham, S.C.H. Barrett, S. Dayanandan y V.A. Albert. 1993. «Phylogenetics of seed plants: An analysis of nucleotide sequences from the plastid gene rbcL.». Ann. Missouri Bot.
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