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1-Pteridofitas.Pdf PRÓLOGO El presente trabajo es un aporte de las Asignaturas Diversidad Vegetal y Biotaxonomía de Brióftas y Pteridófitas, de la Facultad de Ciencias Exactas y Naturales y Agrimensura (UNNE). Tiene por finalidad facilitar la preparación del contenido correspondiente a las bolillas de Pteridófitas de ambas asignaturas. Se intenta brindar una orientación en el estudio de la exomorfología, taxonomía y distribución de los diferentes grupos de Pteridófitas. Además, se introducen aspectos filogenéticos como resultado de estudios moleculares recientes siguiendo la clasificación Pryer et al. (2004). Esta labor es el resultado de la tarea conjunta de los siguientes integrantes de las Asignaturas: ¾ Dirección y corrección: Dra. Elsa L. Cabral (Prof. Titular). ¾ Elaboración original: Dra. Elsa L. Cabral (Prof. Titular). ¾ Actualización de acuerdo al Sistema de Clasificación de Pryer et al. (2004): Lic. Walter A. Medina y Laila M. Miguel (Adscriptos). ¾ Compaginación general: Laila M. Miguel y Lic. Walter A. Medina (Adscriptos) ¾ Fotos: Elsa L. Cabral, Andrea A. Cabaña Fader, Walter A. Medina, Roberto M. Salas y Sandra G. Martín. ¾ Dibujos: Daniel Cian. 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. Guía de Consulta Diversidad Vegetal- FACENA (UNNE) PTERIDOFITAS- Prólogo INDICE Página Características Generales 2 Introducción a la nueva clasificación 5 Clave didáctica para diferenciar los grupos de Pteridófitos 8 Clave de familias para Pteridófitos de Argentina 9 Lycophyta: 1. Selaginellales 14 2.Isoetales 17 3.Lycopodiales 20 Monilophytes Eusporangiados Psilotaceae 27 Ophioglossaceae 30 Equisetaceae 34 Marattiaceae 38 Leptosporangiados Osmundaceae 39 Hymenophyllaceae 42 Gleicheniaceae 47 Schizaeaceae 51 Core leptosporangiados Helechos Heterosporados Marsiliaceae 57 Salviniaceae 61 Azollaceae 63 Helechos Arborescentes Cyatheaceae 65 Lophosoriaceae 68 Dicksoniaceae 70 Helechos Polypoideos Dennstaedtiaceae 72 Aspleniaceae 74 Thelypteridaceae 77 Blechnaceae 80 Parkeriaceae 82 Pteridaceae 84 Davalliaceae 89 Dryopteridaceae 91 Grammitidaceae 94 Lomariopsidaceae 96 Polypodiaceae 98 Vittariaceae 103 BIBLIOGRAFIA 106 Guía de Consulta Diversidad Vegetal- FACENA (UNNE) PTERIDOFITAS- Indice 2 Pteridófitos Los pteridófitos son plantas cormófitas con alternancia de generaciones y fases independientes, donde el esporofito (asexual) está más desarrollado que el gametofito (sexual). El esporofito está constituido por raíz, tallo y hojas. Presenta un tejido de conducción formado por traqueidas y células cribosas. La raíz es de origen caulinar, algunas plantas epífitas o acuáticas carecen de raíces. Los tallos con frecuencia son rizomatosos breves o bien desarrollados, constituyendo verdaderos troncos (helechos arborescentes). Presenta diversidad de tipos de estela: protostélico, solenostélico o dictiostélico a veces polistélico. Con respecto al origen y la estructura de las hojas pueden ser enaciones, micrófilos y megafilos. Los esporangios pueden tener paredes gruesas (eusporangiados) o delgadas (leptosporangiados), isospóreos o heterospóreos, terminales, axilares o sobre las hojas. Esporas triletes o monoletes. Gametofitos autótrofos o micotróficos. Gametas masculinos flagelados formados en anteridio. Gameta femenina inmóvil que se encuentra en un arquegonio. Ciclo de Vida Helecho heterosporado: Selaginella sp. Por acción de la gravedad o del viento, los microsporangios con las micrósporas caen en la zona de los macrosporangios. Cada microspora presenta su prótalo reducido, de desarrollo endospórico, con anteridios y anterozoides biciliados y las macrosporas con su prótalo endospórico con arquegonios. Con las gotas de lluvia o de rocío, los anterozoides pueden llegar a los arquegonios y a la oósfera y producirse la singamia. El cigoto se desarrolla formando un embrión con suspensor, nutrido por el material almacenado en la macróspora, que posteriormente se transforma en esporofito maduro. Microsporofilo Esporofito Microsporas Megásporas Megasporofilo Protalo endospórico Anterozooides Guía de Consulta Diversidad Vegetal- FACENA (UNNE) PTERIDOFITAS- Características Generales 3 Helecho isosporado: Nephrolepis sp. En el envés de las frondes del esporofito diploide se encuentran los esporangios conteniendo esporas haploides originadas por meiosis en el tejido esporógeno. Las esporas caen al suelo y, en condiciones apropiadas de humedad, germinan dando origen al gametofito (prótalo), inconspicuo. Este produce, a partir de células epidérmicas de su cara ventral, los gametangios: anteridios y arquegonios. Los anteridios están formados por unas pocas células y en su interior se forman los anterozoides flagelados. Los arequegonios son más complejos y constan de un cuello de varias células que emergen de la superficie del gametofito, y de un vientre donde esta contenida la ovocélula. La ovocélula fecundada o cigota permanece contenida en el arquegonio mientras se desarrolla un embrión. Este embrión se mantiene unido al gametofito por medio de un pie, ya que el mismo le sirve de cuerpo nutricio hasta la formación de las primeras hojas. Gametofito Esporas Ovocélula Esporangio Anterozooide Esporofito Características de las Lycophytas Plantas terrestres, epipétricas, epifiticas o palustres; ejes caulinares rastreros con raíces y micrófilos, esporofilos agrupados o no en estróbilos, micrófilos con lígulas o sin lígulas. Con un esporangio libres, isospóreas y heterospóreas, triletes. Sin elateres. Los gametofitos de helechos heterosporados son unisexuales, endógenos y muy reducidos. Características de las Euphyllophytas Está representada por los helechos verdaderos, es el grupo más evolucionado, más numeroso y diversificado. Presenta un esporofito bien diferenciado en tallo, hojas (megáfilos), generalmente divididas, las raíces (excepcionalmente pueden faltar). Su tamaño es variable, desde muy pequeños de 5-10 cm alt. hasta muy altos de 10-12 (-25) m alt., en los helechos arborescentes. El rizoma es Guía de Consulta Diversidad Vegetal- FACENA (UNNE) PTERIDOFITAS- Características Generales 4 generalmente subterráneo, corto o más o menos erecto, hasta largamente rastrero, o columnar, en los helechos arborescentes. Es frecuente la presencia de hojas de tamaño considerable frondes de vernación circinada. En la mayor parte de las especies las hojas cumplen doble función, fotosíntesis y producción de esporas (trofosporofilos), pero en otras son dimorfas, diferenciadas en las hojas que sólo realizan la función fotosintética (trofofilos) y las hojas que producen esporas (esporofilos). Con respecto a la lámina sólo el 10 % de las especies presentan lámina entera, el resto tienen divididas (pinnadas, bi-tripinnadas). El tamaño varía desde unos centímetros hasta varios metros como en Dicranopteris, que mantiene su capacidad de crecimiento apical durante mucho tiempo. Los esporangios pueden aparecer sobre los esporofilos de la siguiente forma: aislados, soldados en sinangios o agrupados en soros. Son isosporados, salvo los helechos acuáticos Marsileales y Salviniales que son heterosporados. Cuando los esporangios están agrupados en soros, éstos se pueden diferenciar por el orden de desarrollo y maduración de los esporangios en tres tipos: simples, cuando todos los esporangios crecen y maduran al mismo tiempo; gradual, cuando primero maduran los esporangios del centro y mixto cuando en un mismo soro se encuentran esporangios en distintos estadíos de desarrollo. Algunos autores consideran a los soros mixtos como los más especializados. Con respecto a la posición de los soros hay una tendencia evolutiva de los helechos desde la posición terminal, pasando por la marginal hasta la netamente abaxial, lo que se conoce como deslizamiento filético. Los soros pueden aparecer desprovistos de estructura protectora (Osmunda) o estar cubiertos por el indusio, que es una membrana de origen epidérmico, de morfología variable y de valor taxonómico. La pared del esporangio está formada por dos estratos de células (eusporangiados) o por un solo estrato (leptosporangiados). Éstos últimos, excepto Osmundales, Salviniales y Marsileales, se caracterizan por la existencia de un mecanismo de dehiscencia, el anillo mecánico de dehiscencia, éste puede estar situado transversal, oblicuo o longitudinal o en forma de un casquete en los más primitivos. Presentan las paredes interiores y laterales engrosadas y las exteriores delgadas. Al madurar las esporas en el interior del esporangio, las células del anillo pierden agua y se produce la rotura violenta de la pared del esporangio por unas células especiales de apertura, el estomio. Los movimientos de apertura facilitan la dispersión de la espora. Los gametofitos de helechos isosporados son hermafroditas, laminares, epigeos y clorofilianos, a veces filamentosos o pueden ser subterráneos y heterótrofos. Los gametofitos de helechos heterosporados son unisexuales, endógenos y muy reducidos. Los anterozoides son pluriciliados. Esta clase comprende cerca de 51 familias, 400 géneros y 10.000 especies. El 10% de los taxones son cosmopolitas, la mayoría viven en un área tropical y aproximadamente el 40% son endémicas de América Central y el Caribe. Predominan las especies terrestres, unas pocas son epífitas y escasas son acuáticas. Distribución y hábitat. Crecen en una diversidad de ambientes: terrestre, saxícola, epífito, palustre y acuático. Se encuentran desde el nivel del mar hasta los 5000 m alt. Se concentran en los trópicos y su distribución se extiende hasta las regiones
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