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Core Eudicotiledóneas UNNE FACENA Core Eudicotiledóneas DIVERSIDAD VEGETAL BIOTAXONOMÍA 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, Sandra G. Martin, Enrique A. Szelag, Otto Ferber. 9 Dibujos: Daniel Cian. 9 Elaboración original y revisión bibliográfica de los taxones que actualmente pertenecen al Clado Core 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 Características generales 1 5.1 Orden Saxifragales 3 5.1.1. Familia Crassulaceae 5 5.1.2. Familia Haloragaceae 9 5.1.3. Familia Hamamelidaceae 12 5.1.4. Familis Saxifragaceae 16 5.2. Orden Santalales 20 5.2.1. Familia Misodendraceae 22 5.2.2. Familia Olacaceae 25 5.2.3. Familia Opiliaceae 28 5.2.4. Familia Loranthaceae 31 5.2.5. Familia Santalaceae 34 5.3. Caryophyllales 38 5.3.1. Familia Droseraceae 40 5.3.2. Familia Nepenthaceae 45 5.3.3. Familia Tamaricaceae 48 5.3.4. Familia Plumbaginaceae 52 5.3.5. Familia Polygonaceae 55 5.3.6. Familia Caryophyllaceae 60 5.3.7. Familia Achatocarpaceae 64 5.3.8. Familia Amaranthaceae 68 5.3.9. Familia Phytolaccaceae 75 5.3.10. Familia Nyctaginaceae 81 5.3.11. Familia Molluginaceae 88 5.3.12. Familia Basellaceae 91 5.3.13. Familia Portulacaceae 94 5.3.14. Familia Cactaceae 98 5.3.15. Familia Aizoaceae 106 5.5. Familia Dilleniaceae 110 1 5. Core-Eudicotiledóneas Características generales Los integrantes del Core-Eudicotiledóneas se caracterizan por presentar flores típicamente pentámeras, una clara diferenciación entre sépalos y pétalos, piezas del androceo en número doble al de los sépalos, polen tricolpado y carpelos connados y en igual número que las piezas del cáliz (Bremer et al., 2003; Friere Fierro, 2004). Sin embargo, la delimitación de este clado no estuvo basada en caracteres morfológicos; sino en numerosos datos obtenidos del análisis de ADN que soportan fuertemente su monofilia (Soltis et al., 2005). Este clado abarca siete subgrupos fuertemente soportados (D. Soltis y otros, 2000, 2003; APG II, 2003). Los grupos desarrollados en esta guía están resaltados con negrita y con recuadro: 1. Gunnerales (Myrothamnaceae y Gunneraceae). 2. “Berberidopsidales” (grupo que no fue reconocido como tal por el APG II; comprende a las familias: Berberidopsidaceae y Aextoxicaceae). 3. Saxifragales 4. Santalales 5. Rosideas 6. Dilleniaceae/Caryophyllales Core 7. Asterideas Eudicotiledóneas http://tolweb.org/Core_Eudicots/20714 Los estudios recientes identificaron al orden Gunnerales como grupo hermano del resto de las core-eudicotiledóneas (D. Soltis y otros., 2003; Hilu et al., 2003). Sin embargo, la relación del resto de los linajes todavía se encuentra sin resolver. El orden Gunnerales es un grupo que sorprendió a los taxónomos. La familia Gunneraceae y Myrothamnaceae difieren bastante con respecto a su morfología, granos de polen y hábito; Gunneraceae tiene representantes herbáceos mesofíticos y Myrothamnaceae presenta integrantes leñosos de zonas áridas. La plesiomorfia del orden sería la similitud en la forma del perianto. Las dos familias son pequeñas y monogenéricas. En el A.P.G. II (2003), es opcional incluir a ambas dentro de un mismo orden. Y Según Wilkinson (2000), ambas deberían mantenerse como clados separados. La familia Gunneraceae presenta 5 especies en el sur de nuestro pais y Gunnera apiculata habita en Salta y Jujuy. El orden Berberidopsidales es un clado bien soportado por los estudios moleculares y la morfología de los estomas. Incluye a la familia Berberidopsidaceae de Australia y Chile y a Aextoxicaceae, familia monotípica presente en nuestro país: Aextoxicum punctatum nativa de Chubut y Rio Negro que tendría su origen en Chile. Diversidad Vegetal- Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS- Características generales 2 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: 399–436. -Bremer, K., B. Bremer y M. Thulin. 2003. Introduction to Phylogeny and Systematics of Flowering Plants. Department of Systematic Botany Evolutionary Biology Centre. Uppsala University, 100 p. -Drinnan, A. N., P.R. Crane y S. B. Hoot.1994. Patterns of floral evolution in the early diversification of non-magnoliid dicotyledons (eudicots). Plant Systematics and evolution, supplement 8: 93-122. -Freire Fierro, A. 2004. Botánica Sistemática Ecuatoriana. Missouri Botanical Garden, FUNDACYT, QCNE, RLB y FUNBOTANICA. Murray Print, St. Louis. 209 p. -Hoot, S. B., S. Magallón y P. R. Crane. 1999. Phylogenetic of basal eudicots based on three molecular data sets: atpB, rbsL, and18S nuclear ribosomal DNAsequences. Annals of the Missouri Botanical Garden 86: 1-32. -Judd, W. S., C. S. Campbell, E. A. Kellogg, P.F. Stevens y M. J. Donoghue. 2002. Plant Systematics: a Phylogenetic Approach. Sinauer, Sunderland, MA. -Soltis, D.E., P.S. Soltis, P.K. Endress y M.W. Chase. 2005. Phylogeny and Evolution of Angiosperms. Sinauer Associates, Inc. Publishers, U.S.A. -Soltis, Doug, Pam Soltis, and Christine Edwards. 2005. Core Eudicots. Core Tricolpates. Version 01 January 2005.http://tolweb.org/Core_Eudicots/20714/2005.01.01 in The Tree of Life Web Project, http://tolweb.org/ -Stevens, P. F. 2009. Angiosperm Phylogeny Website. Version 9, June 2008 [and more or less continuously updated since]." will do.http://www.mobot.org/MOBOT/research/APweb/. -Zanis, M. J., P. S. Soltis, Y. L. Qiu, E. Zimmer y D. E. Soltis. 2003. Phylogenetic analyses and perianth evolution in basal angiosperms. Annals of the Missouri Botanical Garden 90: 129-150. Diversidad Vegetal- Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS- Características generales 3 1.1. Saxifragales La composición de este orden es una de las mayores sorpresas para la filogenia molecular de las angiospermas ya que las familias que incluye no habían sido asociadas en clasificaciones anteriores. Los miembros de este orden se encontraban ubicados en tres subclases distintas (Cronquist, 1981). Aunque este grupo incluye menos especies que otras Core Eudicotiledóneas, como Rosideas, Asterideas y Cariophyllales, presenta niveles similares de diversidad morfológica vegetativa y reproductiva. Sus flores varían considerablemente en su disposición, grado de fusión de las partes perianto, número de estambres, grado de unión y número de carpelos y posición del ovario. Además está compuesto de árboles, arbustos, lianas, hierbas anuales y perennes, suculentas y acuáticas. La circunscripción de este orden está basada únicamente en estudios de ADN (Bremer et al., 2003; Freire Fierro, 2004; Soltis et al., 2005; Souza y Lorenzi, 2005, Stevens et al., 2008). Incluye actualmente las siguientes 16 familias (Stevens, 2001) de las cuales se desarrollarán las cuatro que se destacan con negrita por poseer representantes nativos o de importancia económica o evolutiva: Altingiaceae, Aphanopetalaceae, Cercidiphyllaceae, Crassulaceae pág. 5 Cynomoriaceae, Daphniphyllaceae, Grossulariaceae, Haloragaceae pág. 9 Hamamelidaceae pág. 12 Iteaceae, Paeoniaceae, Penthoraceae, Peridiscaceae, Pterostemonaceae, Saxifragaceae pág. 16 Tetracarpaeaceae Diversidad Vegetal- Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS 4 SAXIFRAGALES Bibliografia -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: 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, 100 p. -Cronquist, A. 1981. An Integrated System of Classification of Flowering Plants. Ed. Columbia University Press, 1262 p. -Freire Fierro, A. 2004. Botánica Sistemática Ecuatoriana. Missouri Botanical Garden, FUNDACYT, QCNE, RLB y FUNBOTANICA. Murray Print, St. Louis. 209 p. -Soltis, D.E., P.S. Soltis, P.K. Endress y M.W. Chase. 2005. Phylogeny and Evolution of Angiosperms. Sinauer Associates, Inc. Publishers, U.S.A. -Souza, C.V. y H. Lorenzi. 2005. Botánica Sistemática. Guia Ilustrado para indentificação das famílias de Angiospermas da flora brasileira, baseado em APG
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