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9-Asterides.Pdf 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 del clado Asterideas incluida dentro de las Core Eudicotiledóneas. 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, Andrea A. Cabaña Fader, D. Cardoso, Sylvina Casco, Walter A. Medina, Roberto M. Salas, Otto F. Ferber, Sandra G. Martín y L. P. Queiroz. 9 Dibujos: Daniel Cian. 9 Elaboración original y revisión bibliográfica de los grupos del Clado Asterides: Prof. Carlos Passicot y Dra. Elsa L. Cabral. 9 Actualización y edición de nuevas familias de acuerdo al Sistema de Clasificación APG II (Angiosperm Phylogenetic Group II) 2008: Lic. Roberto M. Salas 2009: Lic. Sandra G. Martín 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 Página 1.1. Introducción 1 1.3.4. Euasterídeas II o Campanulideas 159 1.2. Filogenia 1 1.3.4.1. Orden Aquifoliales 160 1.3. Características de los integrantes de las 3 1.3.4.1.1. Familia Aquifoliaceae 161 Asterides 1.3.1. Orden Cornales 5 1.3.4.2. Orden Apiales 164 1.3.1.1. Familia Loasaceae 6 1.3.4.2.1. Familia Araliaceae 165 1.3.1.2. Familia Hydrangeaceae 12 1.3.4.2.2. Familia Apiaceae 169 1.3.2. Orden Ericales 16 1.3.4.3. Orden Asterales 172 1.3.2.1. Familia Balsaminaceae 18 1.3.4.3.1. Familia Campanulaceae 174 1.3.2.2. Familia Polemoniaceae 21 1.3.4.3.2. Familia Asteraceae 178 1.3.2.3. Familia Lecythidaceae 24 1.3.4.3.3. Familia Calyceraceae 186 1.3.2.4. Familia Sapotaceae 27 1.3.4.4. Orden Dipsacales 189 1.3.2.5. Familia Ebenaceae 33 1.3.3.8.1. Familia Adoxaceae 190 1.3.2.6. Familia Primulaceae 36 1.3.3.8.2. Familia Caprifoliaceae 194 1.3.2.7. Familia Theaceae 39 1.3.3.8.3. Familia Dipsacaceae 197 1.3.2.8. Familia Actinidiaceae 43 1.3.3.8.4. Familia Valerianaceae 200 1.3.2.9. Familia Sarraceniaceae 45 1.3.2.10. Familia Ericaceae 49 1.3.3. Euasterídeas I o Lamiideas 52 1.3.3.1. Familia Boraginaceae 53 1.3.3.2. Orden Gentianales 57 1.3.3.2.1. Familia Rubiaceae 58 1.3.3.2.2. Familia Gentianaceae 66 1.3.3.2.3. Familia Loganiaceae 69 1.3.3.2.4. Familia Apocynaceae 72 1.3.3.3. Orden Lamiales 84 1.3.3.3.1. Familia Oleaceae 85 1.3.3.3.2. Família Gesneriaceae 89 1.3.3.3.3. Familia Plantaginaceae 93 1.3.3.3.4. Familia Scrophulariaceae 100 1.3.3.3.5.Familia Acanthaceae 105 1.3.3.3.6. Familia Bignoniaceae 111 1.3.3.3.7. Familia Lentibulariaceae 118 1.3.3.3.8. Familia Lamiaceae 121 1.3.3.3.9. Familia Verbenaceae 127 1.3.3.3.10. Familia Martyniaceae 133 1.3.3.3.11. Familia Pedaliaceae 137 1.3.3.4. Orden Solanales 140 1.3.3.4.1. Familia Hydroleaceae 141 1.3.3.4.2. Famila Convolvulaceae 143 1.3.3.4.3. Familia Solanaceae 150 1 1. Asterídeas 1.1. Introducción En el sistema de clasificación actual el clado de las Asterideas conforma un grupo monofilético fuertemente soportado, tanto por caracteres moleculares como por morfológicos. La circunscripción de las Asterideas, salvo excepciones de exclusión o inclusión de algunos taxones, no ha sido modificada significativamente en más de 200 años (Freire Fierro, 2004, Soltis et al., 2005). En uno de los primeros sistemas de clasificación, Jussieu (1789) las denomina Monopetalae, posteriormente en el sistema de Engler (1898) fueron agrupadas en Sympetalae, en el sistema de Cronquist (1981) conformaban la Subclase Asteridae y más recientemente en el clado de las Asterideas en AGP I (1989), APG II (2003) y APG 2008 (Stevens, 2001). El sistema de clasificación actual el Clado de las Asterideas está constituido por familias antes incluidas en sistema de Cronquist (1981) en Hammamelidae (Eucomiaceae), Dilleniidae (Theales, Lecythidales, Ericales, Ebenales, Primulales, Sarraceniaceae, Loasaceae) y en Rosidae (Hydrangeaceae, Cornales, Pittosporaceae, Apiales, Aquifoliaceae y Balsaminaceae), la mayoría de ellas con corolas dialipétalas. Las Asterideas comprenden cerca de 1/3 de las plantas con flores, con aproximadamente 80.000 especies, cerca de 4.700 géneros y 114 familias. La mayoría de lo taxones incluidos en este grupo tienen los siguientes caracteres en común: flores gamopétalas (excepto Apiales, que probamente hayan derivado de un ancestro con pétalos unidos), óvulos tenuinucelados (excepto Cornales, el grupo más basal), unitégmicos (excepto Primulaceae y pocos géneros de Cornales) y presencia de compuestos secundarios terpenioideos de tipo iridoide o bien compuestos alcaloides (Freire Fierro, 2004). Este grupo de plantas, además de los caracteres morfológicos y químicos antes mencionados, presentan tubos cribosos, generalmente con plastidios tipo s. Las hojas son mayormente simples, con bordes enteros, dentadas o lobuladas; compuestas, palmaticompuestas con los bordes aserrados; de disposición alterna, verticiladas, decusadas, arrosetadas. Las flores son vistosas bien desarrolladas, perfectas, por lo general pentámeras. Androceo es isómero, usualmente unido al tubo corolino y los estambres son alternos a los lóbulos de la corola. Gineceo gamocarpelar con 2 a 5 cárpelos, con ovario súpero o ínfero. Frutos variados: bayas, cápsulas, cariopses, drupas y pixidios (Cronquist, 1981; Judd et al. 1999, 2002). 1.2. Filogenia De acuerdo al APG II (2003), en el clado de las Asterideas, se reúnen cuatro grupos monofiléticos: a) Cornales, b) Ericales, c) Euasterideas I o Lamiideas (Icacinaceae, Boraginaceae, Garryales, Gentianales, Lamiales y Solanales) y d) Euasterideas II o Campanulideas (Escalloniaceae, Aquifoliales, Apiales, Asterales y Dipsacales) (Freire Fierro, 2004, Soltis et al., 2005). Sin embargo de los grupos antes mencionados, se señalan actualmente a las familias Icacinaceae, Boraginaceae y Escalloniaceae como integrantes de posición incierta (Stevens, 2001). En la delimitación actual se han excluido de este grupo a representantes con corola gamopétala (Cucurbitaceae, antes en Sympetalae) y se han incorporado familias con todos sus representantes dialipétalos (Cornaceae, Apiaceae, Balsaminaceae, Theaceae, Actinidiaceae). Por lo tanto la corola gamopétala no es una sinapomorfía de las Asterideas, ya que también aparece en otros grupos no emparentados (ej. Zingiberales o Cucurbitaceae). Este carácter morfológico debió aparecer tempranamente en la evolución de las plantas con flores (Bremer et al., 2004). Al mismo tiempo la mayoría de los representantes de este grupo son hierbas, aún cuando existen familias con numerosos representantes leñosos, la tendencia actual sigue siendo hacia el porte herbáceo. Las Eusaterideas I presentan típicamente hojas opuestas (Rubiaceae, Loganiaceae). Sin embargo, en el Orden Solanales son principalmente alternas. Las Euasterideas II presentan casi siempre ovario ínfero con un único ovulo por carpelo. Este último clado, Euasterideas II, es el más diverso y más Diversidad Vegetal- Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS- Asterideas-Características generales 2 ampliamente distribuido, ya que incluye entre sus integrantes a una de las familias más diversificada y con numerosas especies: Asteraceae (Freire Fierro, 2004). A continuación se presenta la ubicación filogenética de las Asterideas dentro del gran clado de las Angiospermas, extraído de Bremer et al. (2003). En el siguiente árbol filogenético se representa la posición de las Asterideas según APG, 2008 (Modificado de Steven, 2008). Diversidad Vegetal Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS- Asterideas-Características generales 3 1.3. Características de los integrantes de las Asterideas En esta guía de consulta se desarrollaran a aquellas familias más conocidas, con representantes nativos o de gran importancia económica o evolutiva, las mismas se resaltan con cursiva. Asterideas Basales 1. Cornales Loasaceae Hydrangeaceae Nyssaceae Cornaceae Curtisiaceae Hydrostachyaceae Grubbiaceae 2. Ericales Balsaminaceae Marcgraviaceae Tetrameristaceae Polemoniaceae Fouquieriaceae Lecythidaceae Sladeniaceae Pentaphylacaceae Sapotaceae Ebenaceae Maesaceae Theophrastaceae Primulaceae Myrsinaceae Mitrastemonaceae Theaceae Symplocaceae Styracaceae Diapensiaceae Actinidiaceae Roridulaceae Sarraceniaceae Clethraceae Cyrillaceae Ericaceae Euasterideas I o Lamiideas 3. Garryales Eucommiaceae Garryaceae 4. Boraginaceae 5. Gentianales Rubiaceae Gentianaceae Loganiaceae Gelsemiaceae Apocynaceae 6. Lamiales Oleaceae Gesneriaceae Plantaginacea Scrophulariaceae Calceolariaceae Paulowniaceae Orobanchaceae Acanthaceae Bignoniaceae Lentibulariaceae Lamiaceae Verbenaceae Martyniaceae Pedaliaceae 7. Solanales Montiniaceae Sphenocleaceae Hydroleaceae Convolvulaceae Solanaceae Euasterideas II o Campanulideas 8. Aquifoliales Cardiopteridaceae Stemonuraceae Phyllonomaceae Helwingiaceae Aquifoliaceae Diversidad Vegetal Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE) CORE EUDICOTILEDÓNEAS- Asterideas-Características generales 4 9. Apiales Pennantiaceae Torricelliaceae Griseliniaceae Araliaceae Pittosporaceae Myodocarpaceae Apiaceae 10.
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