Figure 1
Blechnum species silhouettes representing some of the morphological groups assessed in the current work (from herbaria material). A-B. loxense (gr. loxense); B-B. stipitellatum (gr. cordatum), sterile frond; C-B. tabulare (gr. tabulare), fertile and sterile fronds; D-B fragile (gr. fragile), sterile fronds; E-B. penna-marina (gr. penna-marina), individual with sterile and fertile fronds; F-B. laevigatum (gr. occidentale). Scale bar: A, B = 11 cm; C = 12.5 cm; D = 8 cm; E, F = 5 cm.
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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...
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... order to avoid a taxonomically uncertain scenario, specialists have proposed informal groups within Blechnum using morphological criteria. For the American diversity, where around 80 species are recognised (Rolleri & Prada, 2006a), a first approach involving 8 groups has been made, mostly according to characters of rhizome, frond architecture, dimorphism, and habit ( Figure 1 Mett. group;and B. serrulatum Rich. group (Tryon & Tryon, 1982;Durán, 1997;Dittrich, 2005;Rolleri & Prada, 2006b;Passarelli, 2007;Prada et al., 2007;Gabriel y Galán et al., 2008;Ramos Giacosa, 2008;Rolleri et al., 2008;Passarelli et al., 2010;Rolleri et al., 2010;Gabriel y Galán et al., 2011;Rolleri et al., 2012a;Rolleri et al., 2012b;Rolleri et al., 2013). ...
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... 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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... 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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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.
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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.
... 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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... 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. . ...
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.
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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
Blechnum L. is a genus belonging to the Blechnaceae family with 236 accepted species that grow in intertropical, subtropical, and southern temperate regions. Several species of the genus have long been used in folk medicines to treat a broad spectrum of ailments, including typhoid, urinary infections, influenza, wounds, pulmonary complaints, blisters, boils, and antihelmintic-related complications. So far, about 91 chemical compounds have been isolated from different parts of 20 Blechnum species. Among these metabolites, phenolic compounds, sterols, and fatty acids are the main constituents. Modern pharmacological investigations revealed several isolated compounds and extracts to exhibit exceptional biological properties including the antioxidant, antimicrobial, anti-inflammatory, anticancer, insecticidal, antitrematocidal and wound healing. In various tests, both quercetin-7′,3′,4′-trimethoxy and phytol metabolites showed potential antioxidant and antitrematocidal properties, while ponasterone exhibited insecticidal activity. Despite having a broad range of traditional medicinal benefits and biological properties, understanding the scientific connotations based on the available data is still challenging. This article presents a comprehensive review of the traditional uses, phytochemical compounds, and pharmacological aspects of the Blechnum species.
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.
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
