Grana Spore Morphology and Ornamentation in the Genus Blechnum
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This article was downloaded by: [Passarelli, Lilian Mónica] On: 18 November 2010 Access details: Access Details: [subscription number 929785472] Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37- 41 Mortimer Street, London W1T 3JH, UK Grana Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713648917 Spore morphology and ornamentation in the genus Blechnum (Blechnaceae) Lilian M. Passarellia; Jose Maria Gabriel Y Galánb; Carmen Pradab; Cristina H. Rolleria a Laboratorio de Estudios de Anatomía Vegetal Evolutiva y Sistemática, Facultad de Ciencias Naturales y Museo de La Plata, Argentina b Departamento de Biología Vegetal I, Facultad de Ciencias Biológicas, Universidad Complutense, Madrid, Spain First published on: 18 November 2010 To cite this Article Passarelli, Lilian M. , Gabriel Y Galán, Jose Maria , Prada, Carmen and Rolleri, Cristina H.(2010) 'Spore morphology and ornamentation in the genus Blechnum (Blechnaceae)', Grana,, First published on: 18 November 2010 (iFirst) To link to this Article: DOI: 10.1080/00173134.2010.524245 URL: http://dx.doi.org/10.1080/00173134.2010.524245 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Grana, 2010; iFirst: 1–20 SporeSGRA morphology and ornamentation in the genus Blechnum (Blechnaceae) LILIANSpore morphology in Blechnum M. PASSARELLI1, JOSE MARIA GABRIEL Y GALÁN2, CARMEN PRADA2 & CRISTINA H. ROLLERI1 1Laboratorio de Estudios de Anatomía Vegetal Evolutiva y Sistemática, Facultad de Ciencias Naturales y Museo de La Plata, Argentina, 2Departamento de Biología Vegetal I, Facultad de Ciencias Biológicas, Universidad Complutense, Madrid, Spain Abstract A study of the spores of 64 neotropical and palaeotropical taxa of the genus Blechnum was performed using scanning electron microscopy (SEM). Two basic types of perispores were recognised: perispore smooth or only slightly ornamented (laevigate, with micro-granules or micro-lamellae, venulose and rugulate), and perispore ornamented (rugate, low reticulate, cristate- reticulate, orbiculate, and colliculate). SEM observations allow for distinguishing differences in the structure of perispores, but the character does not correlate with the ornamentation. The smooth exospore predominates among the studied spe- cies. Results suggest that perispore characters have important diagnostic value, since different combinations of ornamenta- tion/structure were found among the studied species. Moreover, when spore characters are considered together with other morphological traits of the sporophytes and the habit of plants, ornamentation of spores becomes a useful complementary feature at the specific level. The results are discussed in light of the recent systematic studies and emphasise that any eventual subdivision of the genus Blechnum must include characters of the sporophyte as well as spore features to ensure a more robust classification than the previously proposed. The spores also show a significant affinity between Blechnum and other genera of Blechnaceae, which was also suggested by recent phylogenetic studies. Keywords: Perispore ornamentation, perispore structure, micro-ornamentation, laevigate, rugulate, foliose, rugate, reticulate, cristate-reticulate, exospore The Blechnaceae are a cosmopolitan group of ferns Based on recent phylogenetic analyses, Kenrick Downloaded By: [Passarelli, Lilian Mónica] At: 14:47 18 November 2010 including eight (de la Sota, 1977), nine (Copeland, and Crane (1997) and Pryer et al. (2001) con- 1947; Tryon & Tryon, 1982; Proctor, 1985; Lel- cluded that leptosporangiate ferns represent part linger, 1989; Kramer et al., 1990; Tryon & Stolze, of a monophyletic group, the Monilophyta, which 1993; Moran, 1995) or twelve genera (Pichi Ser- also includes the eusporangiate ferns, the Equi- molli, 1977; Cranfill & Kato 2003). Six genera are setaceae and the Psilotaceae. This configuration confined to the Old World: Brainea, Doodia, Pteri- was supported by Schuettpelz et al. (2006) and doblechnum, Sadleria, Steenisioblechnum and Steno- Schuettpelz and Pryer (2007, 2008), who sug- chlaena; Salpichlaena is exclusively American, gested a rearrangement of the families and genera while Blechnum and Woodwardia occur both in the in the largest order of the leptosporangiate ferns, Old and New World. Anchistea, Chieniopteris and the Polypodiales. According to this concept, the Lorinseria are considered additional well-defined Blechnaceae (including Stenochlaenaceae) would genera by Pichi Sermolli (1977) or they are consist of twelve genera (i.e. Blechnum, Brainea, included in Woodwardia by most authors. Doodia, Pteridoblechnum, Sadleria, Salpichlaena, Correspondence: Lilian M. Passarelli, LEAVES (Laboratorio de Estudios de Anatomía Vegetal Evolutiva y Sistemática), Facultad de Ciencias Natu- rales y Museo La Plata, Universidad Nacional de La Plata, 64 entre 120 y diagonal 113, B1904 DZB, La Plata, República Argentina. E-mail: [email protected] (Received 27 April 2010; accepted 9 September 2010) ISSN 0017-3134 print/ISSN 1651-2049 online © 2010 Collegium Palynologicum Scandinavicum DOI: 10.1080/00173134.2010.524245 2 L. M. Passarelli et al. Steenisioblechnum, Stenochlaena, Woodwardia, Anchi- (1998) and Mickel and Smith (2004) are needed, stea, Chieniopteris and Lorinseria). and must be integrated and updated with new sets of As currently circumscribed, Blechnum is a cosmo- diagnostic characters that will allow clear distinction politan genus comprising around 200 species, of of the species. which about 138 occur in the Southern Hemisphere Any revision attempting to reach a well defined and some 78 species in the Northern Hemisphere sub-generic treatment, provided that sub-generic (Kasmirczak, 1999) with species in America (North division would be justified, should be made on the America, Central America, the Caribbean and South basis of diagnostic characters, a more complete America), Europe to the Ural, Africa, Madagascar, cytological knowledge of the species, and a selection Asia (eastward to Japan), Melanesia, Australia, of the diagnostic traits that could determine with New Zealand and some Pacific Islands (Polynesia to reasonable certainty, the limits, affinities and Tahiti, Marquesas Islands and Rapa Iti). The genus is relationships of species. The systematics conducted in adventitious in Hawaii. Its sporophytes are terrestrial, floristic treatments appears to be based on too varia- erect or scandent, epiphytic, lithophytic or amphibi- ble characters, a problem that results both from the ous plants, adapted to tropical and sub-tropical rain- variation and adaptive phenotypic plasticity of the forests, lowland or montane forests, tepuis, along the species (some characters, such as indumentum, vary edges of forests, thickets, savanna or disturbed areas, with altitude), the different size and ecology of plants, wetlands, meadows, bogs and swamps from cold and also the cytology of species, yet poorly known. sub-Antarctic regions, at altitudes ranging from sea Among potential important features for taxonomic level to 4500 m. The rhizomes are decumbent, creep- delimitations at the species level are also features of ing, scandent or erect to arborescent, scaly, with mon- the spores. While several studies have focused on the omorphic or dimorphic fronds that bear entire, morphology and structure of the spores of pinnatifid or once-pinnate laminae and free, rarely Blechnaceae, and particularly of the genus Blechnum anastomosing venation. Cenosori are formed on a vas- (e.g. Lugardon, 1965, 1974; Morbelli, 1976, 1980; cular commisure or at the base of the indusium, with- Tryon & Lugardon, 1991; Ramos Giacosa et al., out paraphyses. Spores are monolete, with a perispore. 2006, 2009; de la Sota et al., 2006), most of the The number of taxa is not established with cer- studies are mainly descriptive and do not discuss the tainty. Rolleri and Prada (2006a) recorded 92 taxa systematic implications of the features in any detail. in the Neotropics, including species and hybrids, but However, these contributions provide important new species were described later (e.g., Moran & information on spore morphology and structure that Smith, 2005; Rojas-Alvarado, 2006, 2008a, 2008b; may eventually be included in more comprehensive Kessler et al., 2007; Ramos Giacosa, 2010) and par- taxonomic/systematic studies. Some attempts to tial revisions (e.g., Rolleri and Prada, 2006b; Gabriel relate spore types within Blechnum to systematic y Galán et al., 2008; Prada et al., 2008; Rolleri et al., grouping of species have been made recently. These 2008, 2010) have been published