ABSTRACT
Although the genus Blechnum has been largely studied, problems of species characterization, nomenclature, classification and taxonomy remain unsolved. For these reasons, in this study we intend to deepen in the characterization of the arborescent group of Blechnum with a review of eight species: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare and B. werckleanum.Type material and several collections from diverse herbaria were studied. For morphological studies we used both herbarium and fresh material collected by the authors in fieldwork between 2005 and 2010. Rhizomes, stipes, vascularization of costae, as well as several characters of the lamina, such as outline, division, venation, margin structure, epidermal patterns, stomata, presence and types of hypodermis, and mesophyll of pinnae were studied. Important aspects of the morphology of the sporophyte have been analyzed, and external characters have been correlated with internal diagnostic ones to use the most reliable external traits in keys and descriptions. Taxonomy has been updated, and corrections have been introduced to the species nomenclature and comments were included to summarize the current knowledge of the geographical distribution and ecology. The presence of B. tabulare in America is confirmed, and the taxonomy of the species is updated, since external and internal morphology indicated that other related, American and African species, such as B. spannagelii and B. madagascariense must be included under that name. Similar studies carried out on B. schomburgkii indicated that the name is a synonym of B. auratum subsp. auratum, while B. auratum subsp. columbiense is treated here as a well characterized species. Detailed study of numerous specimens allowed to characterize B. moritzianum, being the correct name to apply to the recently described B. yungense. A refreshed description of the species is presented together with a list of new, selected materials to illustrate current known geographical distribution. Expanded data on ecology, geographical distribution and affinities are given, and a key for species identification is presented.
La bibliografía sobre el género Blechnum L. es muy abundante tanto en lo que respecta a las especies neotropicales como paleotropicales. Sin embargo, aún faltan estudios anatómicos completos, una caracterización actualizada de su morfología externa e interna, taxonomía, citología y afinidades. En este estudio se revisaron ocho especies de Blechnum de hábito arborescente: B. brasiliense, B. columbiense, B. cycadifolium, B. magellanicum, B. moritzianum, B. schomburgkii, B. tabulare y B. werckleanum. Se analizaron caracteres de rizomas, estípites y costas, división de la lámina, y venación, estructura de márgenes, modelos epidérmicos, presencia y tipos de hipodermis y mesofilo de las pinnas. Se estudió por primera vez la hipodermis, un área dermatoide adaxial presente en el mesofilo de todas las especies. Se correlacionaron caracteres externos e internos con el fin de utilizar los externos más confiables en claves y descripciones. Los estudios realizados confirmaron nuevamente la presencia de B. tabulare en América, que incluyen en su sinonimia a B. spannagelii de América tropical y B. madagascariense de África. También se concluyó que el nombre B. schomburgkii es un sinónimo de B. auratum subsp. auratum, en tanto que B. auratum subsp. columbiense se trata aquí con rango de especie. Se caracterizó a B. moritzianum, el nombre que corresponde aplicar a la recientemente descrita B. yungense. Se presentó una descripción actualizada de las especies, se resolvieron varios problemas nomenclaturales pendientes; se presentaron datos amplios sobre la ecología, distribución geográfica y afinidades entre las especies, así como una clave para determinarlas.
Subject(s)
Ferns/anatomy & histology , Ferns/classification , Classification/methods , Terminology as TopicABSTRACT
Morphological stomatal traits, such as size, form and frequency, have been subject of much literature, including their relationships with environmental factors. However, little effort have focused on ferns, and very few in the genus Blechnum. Stomatal length, width and frequency (as stomatal index) of a number of specimens of fourteen Neotropical species of Blechnum were measured in adult pinnae. The aim of the work was to find biometrical relationships between stomatal traits and between stomatal traits and habit, habitat and ecosystem of the plants. Statistical analyses of data were conducted using Exploratory Data Analysis and Multivariate Statistical Methods. Stomatal length and width showed a very high correlation, suggesting an endogenous, genetic control, thus giving these traits a considerable diagnostic utility. With respect to the relationships between stomatal traits and environment, we found significant statistical relationships between altitude and stomatal index. We also addressed the interpretation of the ecological- selective significance of various assemblages of stomatal traits in a diverse conjunction of habits, habitats and ecosystems. Rev. Biol. Trop. 59 (1): 403-415. Epub 2011 March 01.
Los caracteres morfológicos estomáticos, tales como tamaño, forma y frecuencia, han sido objeto de abundante investigación, incluyendo su relación con los factores ambientales. Sin embargo, poco esfuerzo se ha realizado en esta materia en helechos y menos todavía en el género Blechnum. En este trabajo se midieron la longitud, anchura y frecuencia (como índice estomático) de estomas de pinnas adultas de un número de individuos en catorce especies de Blechnum neotropicales. El objetivo fue encontrar relaciones biométricas entre los caracteres estomáticos, y entre los caracteres estomáticos y el hábito, hábitat y ecosistema de las plantas. Se realizaron análisis estadísticos como Análisis Exploratorios de Datos y Métodos Estadísticos Multivariantes. La longitud y la anchura de los estomas mostraron una muy fuerte correlación, sugiriendo un control genético endógeno que otorga a estos caracteres un considerable valor diagnóstico. Con respecto a las relaciones entre los caracteres estomáticos y el ambiente, encontramos una relación estadísticamente significativa entre la altitud y el índice estomático. También se incluyen interpretaciones de la significación ecológico- selectiva de un conjunto de caracteres estomáticos en diferentes conjuntos de hábitos, hábitats y ecosistemas.
Subject(s)
Biometry/methods , Ecosystem , Ferns/anatomy & histology , Ferns/classification , Plant Stomata/anatomy & histologyABSTRACT
The fern Blechnum sprucei grows in Mesoamerica (Costa Rica) and South America, from Colombia to Bolivia, SE and centre of Brazil, Paraguay and Argentina. It is a distinctive, somewhat vulnerable, mostly orophilous species. Fresh and dry herbarium material was used for this study. Herbarium material for anatomical studies comes from CTES, BA, LP, MA, SI and UC (Holmgren et al.1990). Selected representative specimens are additionally cited after taxonomic treatment of the species. Dry material was restored with aqueous 4:1 butil cellosolve. Pinnae were cleared with aqueous 6% NaOH, then coloured with aqueous 1 % TBO (Gurr 1966). Hand made transverse sections of young and adult stipes, and costae were done in fresh and restored herbarium material. Venation and epidermal patterns were analyzed in basal, apical and medium pinnae, but only the latter were illustrated. The size and density of stomata were measured in medium pinnae from all studied samples, values shown are the average of 25 measures per sample; sizes are expressed as minimum, media and maximum length x width, in µm, and density as minimum, media and maximum number of stomata / mm². Spores were studied with SEM, mounted on metal stubs with double sided tape, covered with gold under vacuum and photographed with a Jeol /EO JSM 6360 (15 KV) SEM. Spores were also studied with light microscope, mounted in DePex (DePex mounting medium, Gurr, BDH Laboratory Supplies, Poole BH15 1TD, UK) and measured using an ocular micrometer. Measurements are based on a minimum sample of 100 spores taken from different specimens. Sizes are expressed as the longest equatorial diameter/ polar diameter, in µm. Gametophytes were studied from material collected in the subtropical forest of Tucumán Province, Argentina. Spore samples for cultures were taken from single sporophytes kept dry at room temperature since the date plants were collected. Gametophytes were grown under fluorescent light. Multispore cultures were established on mineral agar. Percentage of germination was recorded for a random sample of 50 spores from each of the two plates, every three days until there was no further increasing. Gametophytes were stained with chloral hydrate acetocarmine. The species has large sporophytes, suberect, scaly rhizomes, and dimorphic fronds with short, scaly stipes and lanceolate to elliptic sterile laminae. The rachises can grow indefinitely as radicant axis that vegetatively multi-plicate the plants. Pinnae are lanceolate, herbaceous, with crenate and papillose margins, superficially scaly and hairy, peciolulate, with free, visible veins regularly once furcated near the costa, ending in large, active hydathodes. The broadly elliptic fertile laminae bear distant pinnae, with vegetative tissue reduced to the portion that supports the indusium and the continuous coenosorus; terminal indefinite rachis, not proliferous, may be present. Rev. Biol. Trop. 56 (4): 2027-2040. Epub 2008 December 12.
El helecho Blechnum sprucei crece en Mesoamérica (Costa Rica) y Sudamérica, desde Colombia a Bolivia, SE y centro de Brasil, Paraguay y Argentina. Es una especie característica, algo vulnerable y orófila. Se estudiaron caracteres exomorfológicos de especimenes provenientes de distintos puntos de su gran área de distribución. Adicionalmente, se llevó a cabo un detallado análisis de los modelos epidérmicos jóvenes y maduros, del indumento de la lámina y los ejes, y de la organización vascular en los estipes y costas. Se han estudiado por primera vez la morfología esporal, el desarrollo de los gametófitos, que resultaron cordados y pelosos, y su expresión sexual. Presenta esporófitos grandes, suberectos, con rizomas escamosos y frondas dimórficas, con estipes cortos y escamosos y láminas estériles de lanceoladas a elípticas. El raquis puede crecer indefinidamente como un eje radicante que multiplica las plantas vegetativamente. Las pinnas son lanceoladas, herbáceas, con márgenes crenados y papilosos, y la superficie escamosa y pelosa. Son pecioluladas, con venas libres y visibles, regularmente furcadas cerca de la costa, terminando en hidatodos grandes y activos. Las láminas fértiles son anchamente elípticas y portan pinnas distales; presentan tejido vegetativo reducido a la porción que soporta el indusio y el cenosoro continuo. Pueden presentar raquis indefinidos, pero no prolíferos.