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cpDNA supports the identification of the major lineages of American Blechnum (Blechnaceae, Polypodiopsida) established by morphology

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Blechnaceae is an important leptosporangiate family (9–10 genera, about 250 species). It is monophyletic and distributed mainly in tropical America and Australasia. Among the species 80% belong to Blechnum, a genus with a very complex taxonomy and uncertain internal relationships. In terms of American diversity, the results of morphological studies have arrived at 8 informal groups. Molecular works on this genus are scarce, and there is no information for the majority of American species. The main objective of this work was to evaluate whether the groups proposed to organise the diversity of American of Blechnum are consistent with a molecular analysis. We sequenced 2 chloroplastic regions from species representing all of the groups. In our molecular analysis most of the informal groups were maintained as well supported clades. Only 2 species, B. brasiliense and B. spicant, appear to be isolated from their alleged relatives. Combining our molecular results with previous morphological knowledge, we propose the recognition of 4 lineages: 1) B. serrulatum , 2) B. spicant, and 3) core Blechnum, which represents a large clade that can be divided into core Blechnum I (arborescent species, cordatoids, and B. brasiliense) and core Blechnum II (epiphytic species and herbaceous terrestrials, both monomorphic and dimorphic groups).

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... They recovered several strongly supported clades, but the relationships amongst these clades were poorly resolved. Gabriel y Galán & al. (2013) sequenced trnL-trnF and another chloroplast locus, trnG-trnR, for 17 species representing eight groups that had been recognised by morphology amongst Central and South American species, and found these groups to be largely supported. ...
... This was done by adding our own data, which focused on the south-west Pacific hotspot of species-richness for Blechnum and Doodia (see below), to representatives of Blechnaceae diversity available from GenBank. This principally involved trnL-trnF sequences from Gabriel y Galán & al. (2013), rbcL sequences from the unpublished thesis of Nakahira (2000), and rbcL, rps4, and rps4-trnS sequences from the unpublished thesis of Cranfill (2001); see Appendix 1. We also included sequences of four species of Onocleaceae as outgroups. ...
... Our sampling of taxa and characters is much greater than in previously published phylogenetic studies of the Blechnaceae Gabriel y Galán & al., 2013). Our phylogenetic analyses of chloroplast DNA sequences resolve with strong support many of the deeper relationships within the family (cf. . ...
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The fern family Blechnaceae is cosmopolitan; however, the vast majority of species are placed in Blechnum, which occurs predominantly in the Southern Hemisphere. There are two areas that are particularly species-rich: the south-west Pacific (including Australasia), and Central and South America. Using chloroplast DNA sequences, we report the first comprehensive phylogenetic analysis of the Blechnaceae, including all genera widely recognised in recent treatments, and over half of the species. There is strong support for several major clades, which we characterise morphologically and geographically, and some of their interrelationships. Blechnum is confirmed as polyphyletic. Blechnum indicum and B. serrulatum are more closely related to Salpichlaena and Stenochlaena, and are segregated as a new genus, Telmatoblechnum. Alternative generic circumscriptions are discussed for the remainder of Blechnum. In the absence of morphological characters to diagnose the clades within core Blechnum, and for the sake of taxonomic stability, we advocate a broad circumscription for Blechnum. Brainea and Sadleria are retained as their relationships are not well resolved, but Doodia and Pteridoblechnum are clearly nested within the core of Blechnum and we provide four new names in Blechnum. Additionally, given the focus of our sampling, we discuss the biogeography of the south-west Pacific, where immigration has been more important in community assembly than what might be superficially inferred from patterns of endemism (with ca. 60% of species endemic to individual land areas).
... Blechnaceae is an important leptosporangiate fern family distributed mainly in tropical regions of America and Australasia that comprises around 220-250 taxa in an uncertain number of genera [1,2]. It is estimated that about 80% of the species within Blechnaceae belong to the genus Blechnum [3]. Blechnum is well known for a very complex taxonomy and huge uncertainty regarding species relationships within the genus [3,4]. ...
... It is estimated that about 80% of the species within Blechnaceae belong to the genus Blechnum [3]. Blechnum is well known for a very complex taxonomy and huge uncertainty regarding species relationships within the genus [3,4]. ...
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Blechnum spicant var. fallax Lange is a fern taxon endemic to Iceland where it occurs in the vicinity of hot springs on geothermally heated soils. The taxon was first described by a Danish botanist Johan Martin Christian Lange in 1880 on the basis of plant material collected by Christian Grønlund in Iceland. Because its holotype was not designated in the protologue, we examined the extant original material including illustrations from Flora Danica and a single plant on sheet C10021769 (deposited in C) that was the basis for the respective plate. We select this specimen as the lectotype of Blechnum spicant var. fallax.
... Eight chloroplast markers (atpA, atpB, psbC−trnS, rbcL, rps4−trnS, trnG−R, trnL−F, and trnP−petG) were amplified. Primer design and reaction procedure were performed according to the previous studies , Nagalingum et al. 2007, Lu et al. 2011, Galan et al. 2013. Sanger DNA sequencing was performed at Tsinke (Kunming, China). ...
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The Adiantum pedatum complex is disjunctly distributed in North America and eastern Asia. In this study, we carried out a detailed morphological study based on 137 specimens representing the biogeographic diversity of this complex. The sequences of eight chloroplast markers of 35 samples were analyzed with maximum likelihood and Bayesian inference. The morphological and phylogenetic analyses support the recognition of a new species of the A. pedatum complex from Japan. We propose the new species as A. japonicum based on the examinations of specimens from Japan and the comparative analyses of the whole complex. This new species can be distinguished from A. pedatum by the erect rhizome and shows significant molecular differences from other species in this complex. A taxonomic description with detailed morphological characters of the new species is presented.
... Estas apreciaciones condujeron a una necesaria reconsideración de los taxones de categoría superior, proponiéndose un nuevo sistema para los licófitos y helechos (Smith et al. 2006). Con respecto a los rangos de familia y menores, el tiempo también ha derivado en un aumento notable de su conocimiento filogenético, lo que ha llevado asociados cambios taxonómicos bastante significativos, sobre todo, y lógicamente, en familias grandes tales como Aspleniaceae, Athyriaceae, Blechnaceae, Dryopteridaceae, Polypodiaceae, Pteridaceae y Thelypteridaceae (Schneider et al. 2004a, Christenhusz et al. 2011, Rothfels et al. 2012a, Gabriel y Galán et al. 2013, Schneider et al. 2013, Almeida et al. 2016, Gasper et al. 2016b, Liu et al. 2016, Zhang et al. 2016, por nombrar algunas de las que tienen representación ibérica. ...
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Resumen. Recientemente ha sido publicada una nueva propuesta de clasificación de las plantas vasculares sin semilla (PPG1) hasta el rango de género, basada en caracteres morfológicos y filogenias moleculares, siendo consensuada por un gran número de especialistas en pteridología. Tras un año desde su aparición ha sido ampliamente aceptada por la comunidad científica. Esta nueva propuesta de clasificación presenta una serie de importantes cambios respecto a sistemas anteriores, entre ellos el empleado para la Flora Iberica I. Este trabajo plantea una actualización a la propuesta del PPG1 de la clasificación y nomenclatura de los taxones de licófitos y helechos de la flora ibérica. Abstract. Recently, a new classification proposal for the seedless vascular plants, until the range of genus (PPG1), has come to light. This system considers both morphological characters and molecular phylogenies, and is based on consensus by a large number of specialists in pteridology. In its first year of life, it is being widely accepted by the scientific community. This taxonomic classification presents a series of novelties with respect to previous systems, including the one used for Flora Iberica. This work aims to be an update to the PPG1 proposal of the classification and nomenclature of the Iberian lycophytes and ferns.
... Our results are based on the largest data set thus far compiled for the family, with a sampling of the two main diversity centres: Neotropics and Australasia/ Oceania. With the inclusion of a large neotropical sampling (about 80% of total species), we were able to expand on previous phylogenies (Gabriel y Gal an et al., 2013;. Increased sampling (about 60% of species in the family) made it possible to address critical unresolved issues regarding generic circumscription in Blechnaceae . ...
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Blechnaceae, a leptosporangiate fern family nested within eupolypods II, comprises 200–250 species, typically divided among seven to nine genera. Despite recent molecular studies of the family, it still lacks a modern taxonomic update based on broad sampling from the two centres of diversity—the Neotropics and Australasia/Oceania. To test generic circumscriptions, we have assembled the broadest dataset thus far, from three plastid regions (rbcL, rps4-trnS, trnL-trnF) and with taxonomic sampling focused on both major diversity centres. Our sampling includes 156 taxa and 178 newly generated sequences. We recognize three subfamilies, each corresponding to a highly supported clade across all analyses (maximum parsimony, Bayesian inference and maximum likelihood). The genera Salpichlaena, Stenochlaena and Telmatoblechnum are monophyletic, while Blechnum is polyphyletic, because Brainea, Doodia and Sadleria all nest within it. We outline and explain a plan to resolve the polyphyly of Blechnum by recognizing additional, monophyletic, segregate genera.
... Since the introduction of genetic tools, fern phylogeny has experienced so huge advances that previous works are in perspective seen as unimportant. This is quite far from truth, as phylogenetics confirm, not infrequently, some of the major results previously proposed by morphological and paleobotanical data , Gabriel y Galán et al. 2013. However, at the same time it is obvious that molecular techniques have allowed a very much better comprehension of taxa relationships, not only within families and genera, but also contributing novel findings at the higher levels (Hasebe et al. 1994, Pryer et al. 2004, Schuettpelz & Pryer 2007. ...
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In this review, we report on the most important ideas and features in the field of fern evolution, with particular interest into the main evolutionary mechanisms that shaped the evolutionary history of their diversification. An exhaustive account for the bibliographical corpus on the matter falls beyond the space and extent of this work; instead, we will try to focus the different issues by including featured examples from both historical and recent approaches to the study of the evolutionary mechanisms in ferns. Key words: apomixis, hybridization, molecular techniques, morphology, phylogeny, phylogeography, polyploidy, speciation, species concept
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RESUMEN. – En este trabajo se han estudiado caracteres de las pinnas esporógenas en 42 taxones de Blechnum L., con el fi n de poner en evidencia su valor diagnóstico en el nivel específi co o de grupos de especies afi nes. Se incluyeron especies palustres, herbáceas terrestres, cordatoides, arborescentes y epífi tas o hemiepífi tas. Se estudiaron la anatomía de las pinnas fértiles, los tipos de indusios en relación con las láminas, el receptáculo y la vena comisural, su modelo epidérmico y caracteres de esporangios. En general, los caracteres muestran una fuerte coincidencia con los grupos de afi nidad previamente establecidos por otros rasgos morfológicos y moleculares, y algunos se manifi estan como diagnósticos en el nivel específi co. Se hacen comentarios adicionales sobre los caracteres, a la luz de las diferentes apreciaciones fi logenéticas y evolutivas propuestas para el género. Nuestros resultados sugieren que estos caracteres deben ser incluidos en cualquier propuesta taxonómica que se haga para el género. Palabras-clave: anatomía, epidermis, evolución, indusio, taxonomía ABSTRACT. – Diagnostic characters of sporogenous fronds and sporangia of Blechnum L. (Blechnaceae). We focused on characters of sporogenous pinnae of 42 taxa in the genus Blechnum L. to highlight their diagnostic value in the genus, and whether they vary at species level or are shared by related species. This work included amphibious, herbaceous terrestrial, cordatoid, arborescent and epiphyte-hemiepiphyte taxa. We studied the anatomy of the fertile pinnae, types of indusia in relation with the lamina, the receptacle and commissural vein, the epidermal pattern and the characters of sporangia. In general, these characters correlated with the groups of species previously established by other morphological and molecular characters. Additional comments on the characters are made in the light of phylogenetic and evolutionary hypotheses proposed for Blechnum. Our results suggested that these characters should be considered in any future taxonomic approach of the genus. INTRODUCCIÓN Los estudios detallados sobre caracteres de las frondas esporógenas de Blechnum y otros helechos leptosporangiados son relativamente escasos; con frecuencia se incluyen de forma descriptiva en revisiones o se aportan datos puntuales como la morfología de los indusios y las esporas, pero no existen sufi cientes estudios que analicen este aspecto del esporófi to en relación con la posible obtención de rasgos diagnósticos. Distintos autores (Bower 1914, 1928, Copeland 1947, Haider 1954, Wilson 1959) hicieron referencia a la ontogenia o a la estructura de los esporangios maduros, a la importancia taxonómica de algunos rasgos así como su variación en diversos géneros de helechos leptosporangiados, entre ellos Blechnum. Más recientemente se realizó un estudio citológico que incluyó la ontogenia del esporangio en especies del grupo B. occidentale (González et al. 2010). En trabajos y revisiones también recientes (Gabriel y Galán et al.), se aportan datos específi cos sobre esporas o morfología de los indusios que, si bien no incluyen detalles sobre otros aspectos de las frondas esporógenas, sugieren que la variación de los caracteres estructurales de éstas, así como los de esporangios y esporas, tendrían en conjunto importancia taxonómica en diferentes niveles jerárquicos. Esto es de interés además, porque en Blechnum, las colecciones de herbario de especies dimorfas suelen incluir ejemplares representados sólo por frondas fértiles o por mezcla de frondas fértiles y estériles de especies diferentes, lo que lleva a difi cultades en la determinación. Por lo que antecede, se propone aquí un estudio de la morfología de las pinnas fértiles, indusios y esporangios en 42 taxones del género Blechnum, la mayoría de ellos neotropicales, con la excepción de la especie paleotropical Blechnum ivohibense C. Chr. Tales especies fueron analizadas previamente por los autores en relación con otros caracteres del esporófi to y están representados también todos los grupos de afi nidad propuestos recientemente sobre la base de análisis moleculares (Gabriel y Galán et al. 2013), así como en los establecidos a partir de caracteres morfológicos de los esporófi tos (Tryon & Tryon 1982, Dittrich 2005). Se incluyen B. serrulatum Richard, las especies monomorfas y dimorfas arborescentes, las herbáceas terrestres dimorfas del grupo B. penna-marina Iheringia Série Botânica
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Especímenes de B. tabulare y B. magellanicum procedentes de toda su área de distribución fueron estudiados en detalle y considerados especies diferentes. Se estudiaron los siguientes caracteres: rizomas, escamas rizomáticas, estípites, división de las láminas dimórficas, contorno, textura, tamaño, margen, indumento, venación, modelo epidérmico, estomas (densidad y dimensiones), mesofilo de las pinnas estériles en sección transversal y esporas. El hábito de las plantas, el tipo de rizoma, las escamas rizomáticas y el tipo de ornamentación del perisporio son rasgos compartidos por ambos táxones, pero los restantes caracteres varían en el nivel específico y permiten distinguirlos como especies diferentes. Blechnum tabulare se distribuye en los trópicos y subtrópicos de Sudamérica, África e islas de los océanos Atlántico e Índico, mientras que B. magellanicum tiene una distribución más restringida y crece en áreas húmedas australes de la Argentina y Chile (Sudamérica). Se presenta una descripción completa de los dos táxones, se actualizan los datos sobre su distribución geográfica y ecología, y se comentan las relaciones con otras especies de hábito arborescente. Specimens of B. tabulare and B. magellanicum from their whole geographical area were studied, and taxa were treated as different species. The following characters were analyzed: rhizomes, rhizomatic scales, stipes, division of dimorphic laminae, outline, texture, size, margins, indument, venation, epidermal patterns, stomata (size and density), mesophyll of pinnae in transversal section, mucilaginiferous unicellular glands of axes and laminae, and spores. Habit of plants, type of rhizome, rhizomatic scales, mucilage glands, and type of ornamentation of the perispore are characters shared by the two species, while the other traits vary at the specific level, allowing them to be identified as two separate taxa. Blechnum tabulare is distributed in the tropics and subtropics of South America, África, and Islands of the Atlantic and Indic Oceans, while B. magellanicum is a subantarctic, more restricted species, that grows in humid areas of Argentina and Chile, South America. New descriptions of both species are given, along with comments on their affinities with other arborescent species of the genus.
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Estudios esporales en especies del grupo Blechnum penna-marina (Blechnaceae-Pteridophyta). Las esporas de once taxones del grupo B. penna marina, B. asperum, B. blechnoides, B. corralense, B. fernandezianum, B. microphyllum, B. mochaenum subsp. achalense, B. mochaenum subsp. mochaenum, B. mochaenum subsp. squamipes, B. penna-marina, B. spicant y B. stoloniferum se estudiaron con microscopio de luz y electrónico de barrido. Las esporas son monoletes y elipsoidales en todos los taxones, con macro-ornamentación predominantemente muriforme. El esporodermo consiste de un exosporio psilado, rugulado o con superficie granular y un perisporio rugado, rugulado, venuloso-rugulado, fosulado-rugulado o crestado-reticulado. Una excepción son las esporas de B. fernandezianum, con perisporios psilados con orbículas. El perisporio es estratificado, con capas de diferente morfología y espesor: una capa externa continua, que soporta los elementos de la escultura, una media, alveolar, foliosa o con elementos columnares y una interna muy delgada que no siempre se distingue claramente. Algunos ejemplares de B. mochaenum subsp. mochaenum y B. penna-marina mostraron esporas más grandes en algunos ejemplares, lo que podría relacionarse con diferentes niveles de ploidía. La macro-ornamentación de las esporas y la morfología de las capas observadas en algunos taxones se encuentra también en esporas de otros grupos del género. Los estudios llevados a cabo sugieren que los rasgos esporales no definen un grupo particular sino taxones, una conclusión que abre perspectivas prometedoras para la sistemática del género. Study of the spores of species of the Blechnum penna-marina group (Blechnaceae-Pteridophyta). The spores of eleven taxa of the Blechnum penna-marina group, B.asperum, B. blechnoides, B. corralense, B. fernandezianum, B. microphyllum, B. mochaenum subsp. achalense, B. mochaenum subsp. mochaenum, B. mochaenum subsp. squamipes, B. penna-marina, B. spicant, and B. stoloniferum were studied with light, and scanning electron microscopes . Spores are monolete and ellipsoidal in all taxa, with macro-ornamentation predominantly among muriform types. Sporoderm consists of a psilate, rugulate, or more or less supperficially granulate exospore, and a rugate, rugulate, venulose-rugulate, fossulate-rugulate, cristate-reticulate perispore. An exception are B. fernandezianum spores, with psilate perispores that bear orbicules. The perispore is a stratified, rather complex wall formed by three strata of different thickness and morphology: an outer continuous stratum, which bears the elements of the sculpture, a middle stratum which is alveolate, foliose or bear pillar-like elements, and a very thin, not clearly defined inner stratum. Blechnum mochaenum subsp. mochaenum and B. penna-marina showed larger spores in some specimens, a condition that may be related with different ploidia. Macro-ornamentation and morphology of the perispore of some taxa is also found in species of other groups of Blechnum. Performed studies suggest that spore characters do not define a particular group but the taxa, which is considered a new promising perspective for the systematic of the whole genus.
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Revisión de los grupos de especies del género Blechnum (Blechnaceae-Pteridophyta): el grupo B. penna-marina. Blechnum asperum, B. penna-marina, B. spicant y B. stoloniferum (grupo B. penna-marina) se estudian aquí conjuntamente con'B. blechnoides, B. corralense, B. fernandezianum, B. lehmannii, B. microphyllum, B. mochaenum subsp. mochaenum, B. mochaenum subsp. achalense y B. mochaenum subsp. squamipes. Los esporófitos de estos taxones son pequeños a medianos, con rizomas postrados a erectos y frondas dimórficas. Los caracteres analizados fueron: tipo de rizoma, indumento de ejes y lámina, arquitectura de las frondas, todas con segmentos adnatos y estípites con tres haces vasculares, venación, morfología interna de los segmentos, tales como estructura marginal, modelos epidérmicos de la lámina, estomas, hidatodos, secciones transversales de los estípites, modelos epidérmicos de los indusios y esporas. Esos caracteres resultan'diagnósticos para distinguir los taxones. Se encontraron protuberancias intercelulares pécticas (PIP) en forma de verrugas y filamentos cortos en el mesofilo y tejidos parenquimáticos del estípite y raquis de algunas especies. En el parénquima de los estípites se acumula a menudo almidón de granos simples, elipsoidales. Las esporas son monoletes, aclorofílicas, con esporodermo formado por un exosporio psilado y un perisporio cuya ornamentación varía en los niveles específico e infraspecífico. Tres especímenes de B. penna-marina y tres de B. mochaenum subsps. mochaenum presentaron esporas, células epidérmicas y estomas más grandes que el resto, lo que sugiere diferencias de ploidía, aunque esos ejemplares no muestran variaciones morfológicas externas o internas. Se ha cambiado el estatus de las variedades de B. mochaenum como sigue: B. mochaenum var. fernandezianum es reconocido con el rango de especie, mientras que B. mochaenum var. squamipes y B. mochaenum var. achalense son elevadas al rango de subespecie. Blechnum microphyllum se trata aquí como una especie bien caracterizada y no una subespecie de B. penna-marina. Las plantas de B. penna-marina de Argentina, Brasil y Chile varían algo en el tamaño, pero los rasgos de la morfología interna son muy constantes y no se han reconocido subespecies en el área. Se presenta una descripción completa de los taxones, se actualizan aspectos nomenclaturales, sinonimia, tipos, distribución geográfica, ecología, se da una clave para su determinación y se discute la importancia de los caracteres analizados para redefinir el grupo.
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A critical checklist of the Mesoamerican and South American species and hybrids of Blechnum is presented. For each entry the place of original publication, types, authors, and pertinent synonyms are included. Most of the synonyms are cited with their nomenclatural types. Information on geographical distribution and altitudinal range, ecology, as well as morphological and nomenclatural comments pertinent to the treated taxa are also included. Hybrids that have been cytologically and nomeclaturally studied are treated independently, while not formally described hybrids are cited under their putative parents. Presentamos un catálogo comentado de las especies del género Blechnum L. para Mesoamérica y Sudamérica. Para la mayoría de los nombres se proporcionan los datos sobre indicaciones locotípicas, tomados del protólogo, y sobre los tipos, así como los sinónimos. Siempre que ha sido posible, los sinónimos se citan también con la reseña de sus tipos o indicaciones locotípicas. Se dan referencias sobre distribución geográfica, ecología y rango altitudinal de los táxones y se aportan observaciones que pudieran contribuir al mejor conocimiento de los mismos. Se incluyen los híbridos cuya citología y nombre se conocen, mientras que aquellos que no han sido formalmente descritos se citan por la fórmula híbrida de los nombres de los presuntos parentales.
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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 microm, 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 microm. 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 multiplicate 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 coast, 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.
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Six primers for the amplification of three non-coding regions of chloroplast DNA via the polymerase chain reaction (PCR) have been designed. In order to find out whether these primers were universal, we used them in an attempt to amplify DNA from various plant species. The primers worked for most species tested including algae, bryophytes, pteridophytes, gymnosperms and angiosperms. The fact that they amplify chloroplast DNA non-coding regions over a wide taxonomic range means that these primers may be used to study the population biology (in supplying markers) and evolution (inter- and probably intraspecific phylogenies) of plants.
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Seven Centaurea L. s.l (Asteraceae) taxa endemic to Turkey were examined in terms of anatomical and molecular aspects to contribute to their taxonomic positions. Numerical analysis of the 13 anatomical traits showed that average row number of collenchyma and the arrangements of the vascular bundles in the stem, leaf type, and abaxial surface of leaf epidermal tissue were important to determine the investigated taxa. It was also found that the 7 investigated taxa were grouped into 2 distinct clusters based on anatomical traits and combined nrDNAITS/cpDNA data.
Chapter
The woodwardioid ferns comprise fourteen species distributed among three genera of the Blechnaceae: Anchistea C. Presl, Lorinseria C. Presl, and Woodwardia Sm. The former two genera are monotypic and are confined to eastern North America, while the twelve species of the latter genus are disjunctly distributed throughout the warm temperate and subtropical regions of the northern hemisphere in a classic Arcto-Tertiary distribution pattern. Phylogenetic analyses based on characters derived from morphology and three molecular markers (rbcL,rps4, and rps4-trnS spacer) demonstrate that this group is monophyletic and is sister to the remainder of the Blechnaceae. Within the group, Woodwardia and Anchistea are sister taxa. A comparison of the phylogenetic results with known fossil history suggests that the group first arose sometime in the late Cretaceous in North America at high latitudes. Diversification of the major lineages of the group was complete by the Paleocene, with the lineages subsequently spreading southward into North America and westward into Asia, ultimately reaching Europe by the Miocene. The recent evolutionary history of the group has been characterized by extensive extinction and subsequent vicariance. We evaluate and discuss various hypotheses of relationship and character evolution proposed in the literature, and present a new classification for the group.
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In the present paper, we provide a revised, comprehensive description of the sporophyte and gametophyte of the swamp fern, Blechnum serrulatum Rich., from neo- and paleotropical localities. External and internal characters of the sporophyte were analysed, including axes, laminae, pinnae, indusia and spores. Intercellular pectic connections of the parenchyma of the rhizomes are reported for the first time. In stipes, cell walls of the aerenchyma tissue contain filamentous protuberances that are composed primarily of cellulose but contain also fatty substances. The morphology of the gametophyte, from spore germination to gametangia formation, is discussed. The taxonomic significance of the characters is considered, especially in regard to the relationship between B. serrulatum and the closely related B. indicum.
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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.
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The Blechnaceae is one of the most speciose fern families in New Zealand, with two genera represented: Blechnum and Doodia. We se‐quenced the chloroplast trnL‐trnF locus for all of the Blechnaceae species indigenous to New Zealand, plus several non‐indigenous species. Although deeper relationships were not well resolved by phylogenetic analyses of these DNA sequences, several groupings of species were consistently recovered. Some of these relationships have been previously suspected on the basis of morphological similarity and/or hybridisation (e.g., the B. procerum group), and are consistent with variation in base chromosome numbers, but others were unexpected (e.g., the relationship of B. fluviatile and B. vulcanicum). The species of Doodia sampled here were found to be monophyletic, and were nested within a paraphyletic Blechnum. Infraspecific variation in the trnL‐trnF locus was detected within six New Zealand species, and may prove useful for future phylogeographic and taxonomic studies.
Article
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.
jModelTest 2: more models, new heuristics and parallel computing
  • D Darriba
  • G Taboada
  • R Doallo
  • D Posada
Darriba D, Taboada G, Doallo R & Posada D (2012). jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9: 772-772.
Blechnum subgénero Blechnum en Sur América, con especial referencia a las especies de Colombia
  • M Murillo
Murillo M (1968). Blechnum subgénero Blechnum en Sur América, con especial referencia a las especies de Colombia. Nova Hedwigia 16: 329-366.
A molecular phylogeny of Blechnaceae inferred from three plastid gene sequences
  • R Cranfill
  • Y Nakahira
  • M Kato
Cranfill R, Nakahira Y & Kato M (2001). A molecular phylogeny of Blechnaceae inferred from three plastid gene sequences. Fifth International Flora Malesiana Symposium Abstract.
Estudios morfológicos, taxonómicos y biosistemáticos en el género Blechnum (Blechnaceae-Pteridophyta)
  • M Durán
Durán M (1997). Estudios morfológicos, taxonómicos y biosistemáticos en el género Blechnum (Blechnaceae-Pteridophyta). Universidad Nacional de Córdoba, Córdoba, Argentina.
  • R M Tryon
  • R G Stolze
Tryon RM & Stolze RG (1993). Pteridophyta of Peru. Part. V: Aspleniaceae-Polypodiaceae. Fieldiana Botany 32: 54-70.
Phylogenetic Studies in the Polypodiales (Pteridophyta) with an Emphasis on the Family Blechnaceae
  • R Cranfill
Cranfill R (2001). Phylogenetic Studies in the Polypodiales (Pteridophyta) with an Emphasis on the Family Blechnaceae. Berkeley, CA: University of California.
Revisión sistemática, análisis cladístico y biogeográfico de la sección Lomariocycas
  • Ramos Giacosa
Ramos Giacosa JP (2008) Revisión sistemática, análisis cladístico y biogeográfico de la sección Lomariocycas (J. Sm.) C. V. Morton del género Blechnum L. (Blechnaceae, Pteridophyta) en
Blechnaceae The Families and Genera of Vascular Plants: I. Pteridophytes and Gymnosperms
  • K Kramer
  • T Chambers
  • E Hennipman
Kramer K, Chambers T & Hennipman E (1990). Blechnaceae. In: Kramer K & Green P (eds.) The Families and Genera of Vascular Plants: I. Pteridophytes and Gymnosperms. Berlin: Springer.
Revision of the genus Salpichlaena J. Sm
  • G E Giudice
  • M L Luna
  • C Carrion
  • La De
  • E R Sota
Giudice GE, Luna ML, Carrion C & de La Sota ER (2008). Revision of the genus Salpichlaena J. Sm. American Fern Journal 98: 49-60.