No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Molecular Phylogenetics of Anogramma Species and Related Genera (Pteridaceae: Taenitidoideae)
Abstract
Anogramma is a genus of eight putative species with small annual sporophytes and potentially perennating gametophytes. Phylogenetic relationships within the genus as well as its relationships with other putatively taenitidoid genera and with traditionally cheilanthoid Cosentinia vellea have been poorly resolved and are investigated here. Maximum parsimony, maximum likelihood, and Bayesian inference analyses of rbcL sequences were used to test 1) the monophyly of Anogramma, 2) support for the proposed specific distinctness of A. guatemalensis and A. caespitosa from A. leptophylla, and 3) the asserted close (sister) relationship between Anogramma and Pityrogramma, as well as to infer the phylogenetic relationships among these genera. Results reveal that Anogramma in the traditional sense is polyphyletic. Anogramma guatemalensis and A. caespitosa nest within A. leptophylla. Pityrogramma is not sister to Anogramma as a whole but only to A. chaerophylla and A. novogaliciana. Anogramma osteniana is deeply separated from its traditional congeners and the new combination Jamesonia osteniana (Dutra) Gastony is made. Cosentinia vellea should not be subsumed within Cheilanthes but instead should be considered a taenitidoid genus closely related to A. leptophylla and A. lorentzii. Neighbor joining analysis of Amplified Fragment Length Polymorphism data sets inferred relationships among the A. leptophylla accessions, embedding A. guatemalensis and A. caespitosa in respective New World and Old World clades of A. leptophylla.
... Taxonomy: A genus of two to five species included in subfamily Pteridoideae, along with Pteris (Nakazato & Gastony 2003;Zhang et al. 2015;PPG 1 2016). Anogramma, as traditionally interpreted, has been shown to be polyphyletic (Nakazato & Gastony 2003), with one species more closely related to Jamesonia, others to Pityrogramma, and two previously recognised species nested within Anogramma leptophylla. ...
... Taxonomy: A genus of two to five species included in subfamily Pteridoideae, along with Pteris (Nakazato & Gastony 2003;Zhang et al. 2015;PPG 1 2016). Anogramma, as traditionally interpreted, has been shown to be polyphyletic (Nakazato & Gastony 2003), with one species more closely related to Jamesonia, others to Pityrogramma, and two previously recognised species nested within Anogramma leptophylla. Indeed, only the widespread A. leptophylla and the South American A. lorentzii remain within the Anogramma clade, along with the European Cosentinia vellea, which can be treated as a monotypic genus or included within Anogramma. ...
... A. lorentzii is restricted to South America. Other species previously included in Anogramma either belong elsewhere or are synonymous with A. leptophylla (Nakazato & Gastony 2003). One indigenous species in New Zealand. ...
Pteridaceae is a large family that is almost cosmopolitan in distribution, but has its greatest diversity in
the tropics and in arid regions. It is represented in New Zealand by five indigenous genera (Adiantum,
Anogramma, Cheilanthes, Pellaea, Pteris) and 16 indigenous species, five of which are endemic, and
another ten that are fully naturalised or casual. In addition there is a wholly naturalised genus,
Myriopteris with a single species.
Members of Pteridaceae in New Zealand are terrestrial species with erect to long-creeping rhizomes
and 1–5-pinnate fronds. The sori are either completely unprotected and extend along the veins, or are
marginal and protected by the modified inrolled lamina margin. All species have trilete spores.
Adiantum includes six indigenous species, two of them endemic, and another two naturalised, all
characterised by their polished dark brown stipes, flabellate or oblong ultimate lamina segments, and
sori protected by reniform or oblong reflexed lamina flaps. Anogramma leptophylla is the sole species
of the genus in New Zealand, distinguished by its small delicate laminae, and unprotected sori
extending along the veins. Cheilanthes is represented by two indigenous and one casual species,
having small to medium-sized, 2–3-pinnate fronds, and elongated marginal sori protected by inrolled
pinna margins. Pellaea has two non-endemic indigenous species, with long-creeping rhizomes,
pinnate fronds, scaly stipes and rachises, and scarcely protected sori almost continuous around the
pinna margins. Pteris comprises five indigenous species, three of them endemic, and seven
naturalised or casual species, with medium to large, 1–4-pinnate fronds, and sori continuous along the
pinna edges, protected by membranous inrolled pinna margins.
Most species of Pteridaceae in New Zealand occur more frequently in northern or eastern parts of the
country. Adiantum cunninghamii and Pellaea rotundifolia are unusual in being widespread species.
Adiantum aethiopicum, A. diaphanum, A. fulvum, A. hispidulum, P. macilenta, P. saxatilis and P.
tremula have a mainly northern distribution extending only to northern parts of the South Island, and P.
carsei is confined to the North Island. Anogramma leptophylla, Cheilanthes distans, C. sieberi and
Pellaea calidirupium all occur most commonly in eastern parts of the country, especially in the South
Island. Two species have very restricted distributions – Adiantum formosum currently known only from
the vicinity of the Manawatū Gorge, and Pteris epaleata from Fiordland.
The naturalised genus Myriopteris is represented by a single species, M. lendigera, which is
distinguished by its 3-pinnate fronds, long pale orange hairs on the abaxial surfaces, and sori
produced on small rounded tertiary pinnae.
... Some species closely related with Pityrogramma have been tentatively treated as members of four satellite genera: Anogramma, Cerosora, Cosentinia, and Trismeria Fée (Copeland, 1947;Pichi Sermolli, 1985;Tryon, 1962). Based on the results from rbcL analyses, Anogramma was paraphyletic with its species being resolved as sister to Cosentinia, Jamesonia, and Pityrogramma (Nakazato and Gastony, 2003;Prado et al., 2007;Schuettpelz et al., 2007). Combining new rbcL data of three samples of Anogramma microphylla (Hook.) ...
... Based on previous studies Cochran et al., 2014;Nakazato and Gastony, 2003;Schneider et al., 2013;Schuettpelz et al., 2007;, all currently recognized genera of Pteridoideae were sampled. We included nine accessions representing four species of Actiniopteris, two accessions representing two species of Austrogramme, 34 accessions representing 30 species of Jamesonia, 29 accessions representing 10 species of Onychium, six accessions representing ca. ...
... The recognition of Anogramma is supported by its small and annual habit, its rhizome only with living fronds, and its lamina glabrous or rarely sparsely pubescent (Tryon, 1962;Tryon et al., 1990). Based on rbcL data of nine samples representing seven species of Anogramma, Nakazato and Gastony (2003) found the genus in the traditional circumscription sensu (Tryon et al., 1990) is paraphyletic in relation to Cosentinia and Pityrogramma. With data of six plastid markers of 19 accessions representing all the eight species of Anogramma, our study confirms the paraphyly of Anogramma s.l. ...
As the second most genera-rich fern family, Pteridaceae contain more than 1000 species contributing to ca. 10% of extant leptosporangiate fern diversity. The subfamily Pteridoideae is one of the five subfamilies often recognized. The circumscription of Pteridoideae has not been clear. A large number of species have not yet been included in any molecular analyses before. In this study, DNA sequences of six plastid loci of 154 accessions representing ca. 87 species in 14 genera of Pteridaceae subfam. Pteridoideae and four accessions representing two species in subfam. Parkerioideae and one species of subfam. Adiantoideae as outgroups were used to infer a phylogeny using maximum likelihood and maximum parsimony. Our analyses show that (1) Pteridoideae is monophyletic and the newly defined subfamily is composed of 14 genera including a newly described genus; (2) Pteridoideae is resolved into four strongly supported monophyletic clades: the Pteris clade, the Actiniopteris + Onychium clade, the JAPSTT clade, and the GAPCC clade, these being supported by not only molecular data but also morphological features and distribution information; (3) Onychium is confirmed as monophyletic and accessions of Onychium are resolved into two strongly supported clades, the O. cryptogrammoides clade and the O. siliculosum clade; and (4) Accessions of the traditionally defined Anogramma are resolved as paraphyletic in relation to Cerosora, Cosentinica, and Pityrogramma. Three species traditionally treated in Anogramma are in fact more closely related to Cerosora and Pityrogramma than they are to Anogramma. Gastoniella Li Bing Zhang & Liang Zhang, gen. nov. is described to accommodate these species and three new combinations are provided. Three currently known species of Gastoniella are distributed in the Ascension Island in South Atlantic Ocean, central Mexico, and tropical America, respectively. The new genus is distinct from Anogramma s.s. in having ultimate segments linear not obviously broadening toward the upper portion.
... Although the generic relationships within the Pteridaceae subfamily Pteridoideae have frequently been explored based on DNA sequences of one (Nakazato and Gastony, 2003;Li et al., 2004;S anchez-Baracaldo, 2004b;Prado et al., 2007;Schuettpelz et al., 2007;Bouma et al., 2010;Schneider et al., 2013;Cochran et al., 2014), or three or four Cochran et al., 2014) plastid genes, there have been very few studies focusing on the relationships within Pteris. Li et al. (2004) used trnL-F intergenic spacer sequences (< 400 base pairs) from 16 Chinese species to reconstruct the first phylogeny of Pteris, and concluded that Pteris was strongly supported as monophyletic and that P. vittata L. was resolved as the earliest diverging lineage. ...
... Six plastid regions (the atpA gene, the atpB gene, the rbcL gene, the rps4-trnS intergenic spacer, the trnL intron and the trnL-F intergenic spacer) were selected based on their use in earlier studies of the Pteridaceae (Nakazato and Gastony, 2003;Prado et al., 2007;Schuettpelz et al., 2007;Rothfels et al., 2008;Bouma et al., 2010;Lu et al., 2012;Sigel et al., 2011). The atpA gene was amplified with primers ESATPF412F and ESTRNR46F and the atpB gene with primers ESATB672F and ES-ATPE384R . ...
... Our data resolved the recently described genus Tryonia as sister to a clade containing Austrogramme, Syngramma and Taenitis, and these three together are sister to Pterozonium (Fig. 1a, b), a resolution consistent with findings by Cochran et al. (2014). Nakazato and Gastony (2003) provided the first molecular evidence that Anogramma, as traditionally circumscribed, is highly paraphyletic, with everything except A. leptophylla (and A. guatemalensis) moved to other genera, including Eriosorus, Pityrogramma, etc. Our results confirm the polyphyly of Anogramma. ...
A global plastid phylogeny of the brake fern genus Pteris (Pteridaceae) and related genera in the Pteridoideae Abstract The brake fern genus Pteris belongs to the Pteridaceae subfamily Pteridoideae. It contains 200–250 species distributed on all continents except Antarctica, with its highest species diversity in tropical and subtropical regions. The monophyly of Pteris has long been in question because of its great morphological diversity and because of the controversial relationships of the Australian endemic monospecific genus Platyzoma. The circumscription of the Pteridoideae has likewise been uncertain. Previous studies typically had sparse sampling of Pteris species and related genera and used limited DNA sequence data. In the present study, DNA sequences of six plastid loci of 146 accessions representing 119 species of Pteris (including the type of the genus) and 18 related genera were used to infer a phylogeny using maximum-likelihood, Bayesian-inference and maximum-parsimony methods. Our major results include: (i) the previous uncertain relationships of Platyzoma were due to long-branch attraction; (ii) Afropter-is, Neurocallis, Ochropteris and Platyzoma are all embedded within a well-supported Pteris sensu lato; (iii) the traditionally cir-cumscribed Jamesonia is paraphyletic in relation to a monophyletic Eriosorus; (iv) Pteridoideae contains 15 genera: Actiniopteris, Nephopteris (no molecular data), Onychium, Pityrogram-ma, Pteris, Pterozonium, Syngramma, Taenitis and Tryonia; and (v) 15 well-supported clades within Pteris are identified, which differ from one another on molecular, morphological and geographical grounds, and represent 15 major evolutionary lineages.
... Although the generic relationships within the Pteridaceae subfamily Pteridoideae have frequently been explored based on DNA sequences of one (Nakazato and Gastony, 2003;Li et al., 2004;S anchez-Baracaldo, 2004b;Prado et al., 2007;Schuettpelz et al., 2007;Bouma et al., 2010;Schneider et al., 2013;Cochran et al., 2014), or three or four Cochran et al., 2014) plastid genes, there have been very few studies focusing on the relationships within Pteris. Li et al. (2004) used trnL-F intergenic spacer sequences (< 400 base pairs) from 16 Chinese species to reconstruct the first phylogeny of Pteris, and concluded that Pteris was strongly supported as monophyletic and that P. vittata L. was resolved as the earliest diverging lineage. ...
... Six plastid regions (the atpA gene, the atpB gene, the rbcL gene, the rps4-trnS intergenic spacer, the trnL intron and the trnL-F intergenic spacer) were selected based on their use in earlier studies of the Pteridaceae (Nakazato and Gastony, 2003;Prado et al., 2007;Schuettpelz et al., 2007;Rothfels et al., 2008;Bouma et al., 2010;Lu et al., 2012;Sigel et al., 2011). The atpA gene was amplified with primers ESATPF412F and ESTRNR46F and the atpB gene with primers ESATB672F and ES-ATPE384R . ...
... Our data resolved the recently described genus Tryonia as sister to a clade containing Austrogramme, Syngramma and Taenitis, and these three together are sister to Pterozonium (Fig. 1a, b), a resolution consistent with findings by Cochran et al. (2014). Nakazato and Gastony (2003) provided the first molecular evidence that Anogramma, as traditionally circumscribed, is highly paraphyletic, with everything except A. leptophylla (and A. guatemalensis) moved to other genera, including Eriosorus, Pityrogramma, etc. Our results confirm the polyphyly of Anogramma. ...
The brake fern genus Pteris belongs to the Pteridaceae subfamily Pteridoideae. It contains 200–250 species distributed on all continents except Antarctica, with its highest species diversity in tropical and subtropical regions. The monophyly of Pteris has long been in question because of its great morphological diversity and because of the controversial relationships of the Australian endemic monospecific genus Platyzoma. The circumscription of the Pteridoideae has likewise been uncertain. Previous studies typically had sparse sampling of Pteris species and related genera and used limited DNA sequence data. In the present study, DNA sequences of six plastid loci of 146 accessions representing 119 species of Pteris (including the type of the genus) and 18 related genera were used to infer a phylogeny using maximum-likelihood, Bayesian-inference and maximum-parsimony methods. Our major results include: (i) the previous uncertain relationships of Platyzoma were due to long-branch attraction; (ii) Afropteris, Neurocallis, Ochropteris and Platyzoma are all embedded within a well-supported Pteris sensu lato; (iii) the traditionally circumscribed Jamesonia is paraphyletic in relation to a monophyletic Eriosorus; (iv) Pteridoideae contains 15 genera: Actiniopteris, Anogramma, Austrogramme, Cerosora, Cosentinia, Eriosorus, Jamesonia, Nephopteris (no molecular data), Onychium, Pityrogramma, Pteris, Pterozonium, Syngramma, Taenitis and Tryonia; and (v) 15 well-supported clades within Pteris are identified, which differ from one another on molecular, morphological and geographical grounds, and represent 15 major evolutionary lineages.
... Anogramma spp. are unique among ferns in having pea-like structures called tubercles, except in the case of A. osteniana Dutra (Nakazato & Gastony, 2003). The tubercles have the capacity to stay dormant during environmentally stressful periods and to generate new gametophytic lobes when favourable environmental conditions resume (Baroustis, 1976;Pangua, Pérez-Ruzafa & Pajarón, 2011). ...
... With the exception of A. ascensionis, DNA sequences used for phylogenetic analysis were downloaded from GenBank, using the same accessions as in Nakazato & Gastony (2003). Genomic DNA from A. ascensionis, previously collected at the original locality on Ascension Island, was obtained from the RBG, Kew DNA bank. ...
... Genomic DNA from A. ascensionis, previously collected at the original locality on Ascension Island, was obtained from the RBG, Kew DNA bank. The rbcL region was amplified by polymerase chain reaction (PCR) using the primer combinations described in Nakazato & Gastony (2003). PCR products were purified using DNA purification columns according to the manufacturer's protocol (QIAquick; Qiagen Ltd, Crawley, UK) and cycle sequenced using Big Dye terminator v3.1 chemistry (ABI, Warrington, Cheshire, UK) following the protocol recommended by the manufacturer. ...
Last seen in 1958, the Ascension Island endemic fern, Anogramma ascensionis, was listed as extinct on the 2003 IUCN Red List. However, a 2009 survey rediscovered four plants on Green Mountain. Spores were collected and cultured in vitro at the Royal Botanic Gardens, Kew, where a living collection of thousands of gametophytes and hundreds of sporophytes has been developed. To gain further insights into the biology of this species and the potential implications of in vitro multiplication for conservation purposes, samples were characterized from the karyological point of view. Chromosome analysis of root tips has confirmed that the species is tetraploid, and flow cytometry assessments have revealed that haploid gametophytes produce diploid sporophytes, which confirms natural fertilization. In addition, an rbcL sequence from A. ascensionis has been generated and compared with those published for other Anogramma spp., suggesting a close relationship with A. chaerophylla from Brazil. Further surveys of Green Mountain have reported the presence of 40 A. ascensionis sporophytes in total. Vegetation community analyses have suggested that the present population may be confined to suboptimal habitats. We therefore propose that, prior to the dramatic transformation of the vegetation on the island as a result of the invasion of alien species (particularly Adiantum spp.), A. ascensionis may have flourished in more humid and shaded parts of the mountain. A multidisciplinary approach involving in vitro culture, invasive species clearance and controlled translocation is discussed as the future roadmap for the conservation of this critically endangered fern. Our experiences have also highlighted lessons more broadly applicable to the conservation of extremely rare species elsewhere in the world, especially on remote island systems.
... Of the six subfamilies recognized in the family (Tryon et al., 1990), only the Cheilanthoideae has been extensively studied from a phylogenetic perspective, using plastid rbcL and nuclear ITS nucleotide sequences Rollo, 1995, 1998). Some members of subfamily Taenitidoideae were included in a larger phylogenetic analysis based on rbcL nucleotide sequences (Gastony and Johnson, 2001;Nakazato and Gastony 2003), and more studies are needed to understand the phylogenetic relationships amongst its members. Phylogenetic relationships within and among the other five subfamilies are yet to be resolved. ...
... Coniogramme fraxinea was also included as a more distantly related outgroup (Hasebe et al., 1995;Pryer et al., 1995;Gastony and Rollo, 1995;Gastony and Johnson, 2001). Coniogramme was initially placed in the cheilanthoids by Tryon et al. (1990) but subsequently shown to be the sister to other traditional Pteridaceae plus Vittariaceae (Hasebe et al., 1995;Gastony and Rollo, 1998;Nakazato and Gastony 2003). All three outgroups are restricted to the Old World. ...
... '' In all analyses, the species of Anogramma and Pityrogramma examined here form a monophyletic group as originally postulated by R. M. . The close relationship between Anogramma and Pityrogramma species is strongly supported by phylogenetic analyses based on rbcL sequence data (Nakazato and Gastony, 2003). They examined more species of Anogramma and Pityrogramma than here, however, finding that Anogramma sensu R. is polyphyletic, with A. osteniana more closely related to Eriosorus and Jamesonia then to other species of traditional Anogramma. ...
Thirteen genera are traditionally recognized in the subfamily Taenitidoideae, Pteridaceae. A phylogenetic study of this subfamily, based on both morphological and molecular data, was performed using an exemplar approach. Representatives of the following genera were included in the analyses: Jamesonia, Eriosorus, Pterozonium, Syngramma, Taenitis, Austrogramme, Pityrogramma, Anogramma, Actiniopteris, Onychium, and Afropteris. Specimens and DNA samples were not available for Cerosora and Nephopteris, so they were excluded. Three species were chosen as outgroups: Pteris multifida, P. quadriaurita, and Coniogramme fraxinea, all of which are restricted to the Old World. A robust phylogeny was generated based on 26 morphological characters, 578 base pairs of the plastid gene rps4 and partial data from the intergenic spacer rps4-trnS. The results reject the hypothesis of monophyly of the subfamily as presented by Tryon et al. (1990). However, the results support the monophyly of a well-supported clade consisting of Jamesonia, Eriosorus, Pterozonium, Austrogramme, Syngramma, Taenitis, Pityrogramma, and Anogramma. The New World genera Jamesonia and Eriosorus form a monophyletic group, and Pterozonium is more closely related to the Old World genera, Austrogramme, Syngramma, and Taenitis.
... All of those genera formed a clade with Pteris supported by a bootstrap value of 95%. Recently, Nakazato & Gastony (2003) used maximum likelihood analysis of a larger selection of taenitidoid genera (Taenitis, Eriosorus, Jamesonia, Anogramma, Pityrogramma, and Cosentinia) and found that these formed a strongly supported taenitidoid clade. Actiniopteris + Onychium formed a clade sister to those taenitidoids, and Pteris species formed a clade that was sister to the two preceding clades. ...
... In this study, Onychium and Actiniopteris were united with Pteris in a clade that was sister to clade formed by Pityrogramma and Anogramma, but this topology was supported by only a bootstrap value of 56% in Fig. 1. As shown in previous studies (Gastony & Johnson 1995;Nakazato & Gastony 2003) and the present study, the precise positioning of Onychium based on rbcL is still problematical. ...
... They are only distantly related and should not be classified together. This was also shown for Coniogramme and Llavea of the cryptogrammoid group by Gastony & Rollo (1995 and for Coniogramme alone by Gastony & Johnson (2001) and by Nakazato & Gastony (2003). ...
Cryptogrammoid ferns were often classified into the cheilanthoid group (of Pteridaceae sensu Tryon et al.) or Sinopteridaceae sensu Ching. In order to elucidate the phylogenetic relationships of cryptogrammoid ferns and their related taxa, we sequenced the chloroplast rbcL gene for 11 Asian species representing Cryptogramma, Onychium, Aleuritopteris, Leptolepidium, Cheilosoria, and Doryopteris. Sequences of other related groups were obtained from GenBank. Phylogenetic analysis of a data matrix including 32 taxa was conducted using the maximum-parsimony (MP) and neighbor-joining (NJ) methods. The results were as follows: (1) Cryptogramma, Coniogramme, and Llavea form a moderately supported clade and constitute a cryptogrammoid group that is distantly related to the cheilanthoid ferns. (2) Onychium is rejected from both the cheilanthoid and the cryptogrammoid group, but closely related to Actiniopteris and Pteris. (3) The cheilanthoid group is monophyletic, which is supported by a high bootstrap value. It comprises most of the genera in Sinopteridaceae sensu Ching and some genera in Hemionitidaceae sensu Pichi Sermolli. (4) In the cheilanthoid group, several genera from the Old World (including Aleuritopteris, Leptolepidiurn, Cheilosoria, and Sinopteris), which have been subsumed into Cheilanthes by some authors, are grouped together and distantly related to the American taxa of Cheilanthes. The present study supports the treatment of the cryptogrammoids as either a separate family or subfamily of a broadly circumscribed Pteridaceae, but Pichi Sermolli's circumscription of Cryptogrammaceae should be emended to include Coniogramme and to exclude Onychium.
... This taxon was described as part of the genus Anogramma, but Nakazato & Gastony (2003), based on molecular phylogenetic analyses, found that A. osteniana is sister and more closely related to Jamesonia and Eriosorus. This species forms a monophyletic group together another species of these two genera and with high support (ML = 100). ...
... So it was combined into Jamesonia. The similarity of the laminae and spores have already been indicative of the phylogenetic proximity of A. osteniana with Jamesonia and Eriosorus (Nakazato & Gastony 2003). ...
Jamesonia is a genus of Neotropical ferns that comprises about 50 species, distributed from Mexico to Uruguay. In spite of this wide distribution, this group is more diverse in Andean páramos and subpáramos. Due to the recent changes in its circumscription, with the junction of Eriosorus and segregation of Tryonia, the objective of this work was to carry out the taxonomic study of Jamesonia for Brazil, in order to elaborate its monograph to the Flora of Brazil 2020. This study was based on morphological analyses of specimens deposited in 25 Brazilian herbaria, plus online images of types, field expeditions in the southern and southeastern regions of Brazil, revision of the literature, and scanning electronic microscopy of the spores. Seven species and two hybrids were recognized: J. biardii, J. brasiliensis, J. cheilanthoides, J. flexuosa, J. insignis, J. osteniana, J. rufescens, J. brasiliensis × J. cheilanthoides, and J. ×intermedia, respectively. A distinct specimen, from the border of Brazil (Amazonas) and Venezuela, was treated as Jamesonia sp., due to the presence of only one material. Identification key, descriptions and illustrations are provided for the species and hybrids, as well as, geographical distribution data, comments, list of selected material examined for each taxon, and a full list of all exsiccatae analyzed. We also present an identification key for the genera Jamesonia and Tryonia.
... A nivel mundial, el género Anogramma Link (Pteridaceae) comprende ocho especies de helechos homospóricos, los que se distinguen por el hábito anual del esporofito y la persistencia del gametofito por medio de tubérculos (Tryon & Tryon 1982, Nakazato & Gastony 2003, Luna et al. 2016, características que otorgan a estos helechos una estrategia de vida similar a la de una briofita (Proskauer 1964, Pangua et al. 2011. ...
... Crece no sólo en los neotrópicos, sino también en la cuenca del Mediterráneo, Macaronesia, Europa Atlántica, África tropical, Sudáfrica, Asia (India), Australia y Nueva Zelandia (Bostock et al. 1998, Pangua et al. 2011). Landolt (2010) menciona a la especie como restringida a zonas templadas y tropicales, con preferencia para regiones húmedas, y generalmente con alternancia húmeda y temporadas secas (Nakazato & Gastony 2003). En este trabajo se reporta por primera vez a A. leptophylla en Chile, basado en material recolectado en la Cordillera de los Andes de la Región del Maule, y consultas a la base de datos de los herbarios de la Universidad de Concepción (Concepción), Museo Nacional de Historia Natural (Santiago), Cátalogo de plantas vasculares de Chile (Rodríguez et al. 2018) y Catálogo de las plantas vasculares del Cono Sur (Zuloaga et al. 2018 en adelante). ...
The genus Anogramma Link (Pteridaceae), by means of Anogramma leptophylla (L.) Link is reported for the first time in Chile, from a collection recently made in the Andes Mountains, Maule Region.
... The genus Anogramma Link (Pteridaceae) comprises seven species of homosporous ferns with typically annual small sporophytes and potentially perennating gametophytes (Nakazato & Gastony, 2003). These species grow often in environments with alternating wet and dry seasons (Tryon & Tryon, 1982). ...
... Anogramma is found mainly in the tropics of Mexico and Central and South America, although Anogramma leptophylla (L.) Link inhabits also in Africa, southern Europe to northern India, Australia and New Zealand (Tryon & Tryon, 1982;Nakazato & Gastony, 2003). ...
p>El cultivo in vitro de esporas y los aspectos reproductivos del helecho anual Anogramma chaerophylla (Pteridaceae). Anogramma chaerophylla pertenece a un género de helechos con esporofitos anuales y gametofitos potencialmente perennes. En el área de estudio, la Reserva Natural Punta Lara, la vegetación crece bajo estrés debido principalmente a la contaminación y la invasión de especies exóticas. Como parte de un proyecto de conservación de helechos que habitan en la provincia de Buenos Aires, los objetivos del trabajo fueron evaluar en Anogramma chaerophylla las condiciones para la germinación de esporas in-vitro y analizar las distintas etapas de su ciclo reproductivo. Las esporas fueron desinfectadas en solución 10 % de NaCIO durante distintos tiempos y sembradas luego en medio de cultivo Murashige – Skoog, sin agregado de sacarosa. Las cápsulas se incubaron con un fotoperíodo de 12 h luz/oscuridad y una temperatura de 22 (± 2) °C. A las dos semanas germinaron el 80% de las esporas. El patrón de germinación fue tipo Vittaria y el desarrollo del protalo tipo Ceratopteris.
Los gametangios aparecieron primero en la región engrosada del protalo donde posteriormente se originaron los tubérculos bisexuales. Los esporofitos se desarrollaron solo en asociación con los tubérculos. Durante su cultivo en macetas, éstos dieron origen a una segunda generación de protalos.
Nuestros resultados demuestran que la técnica de cultivo in vitro es apropiada para la propagación de Anogramma chaerophylla y como estrategia para su conservación ex-situ.</p
... The genus Anogramma Link (Pteridaceae) comprises seven species of homosporous ferns with typically annual small sporophytes and potentially perennating gametophytes (Nakazato & Gastony, 2003). These species grow often in environments with alternating wet and dry seasons (Tryon & Tryon, 1982). ...
... Anogramma is found mainly in the tropics of Mexico and Central and South America, although Anogramma leptophylla (L.) Link inhabits also in Africa, southern Europe to northern India, Australia and New Zealand (Tryon & Tryon, 1982;Nakazato & Gastony, 2003). ...
Anogramma chaerophylla belongs to a fern genus with annual sporophytes and potentially
perennating gametophytes. In the studied area, Natural Reserve Punta Lara, the plants grow stressed
mainly by pollution and the invasion of exotic species. As a part of a project on conservation of ferns
inhabiting Buenos Aires province, the objectives of this work were to evaluate in Anogramma chaerophylla
the conditions for in-vitro spore germination and to analyse the different stages of its reproductive cycle.
Spores were sterilized in an aqueous solution 10 % of NaCIO during different times and then sown in Petri
dishes with Murashige & Skoog medium, without the addition of sucrose. The dishes were kept under
laboratory conditions, at 12 h light/ darkness photoperiod and a temperature of 22 (± 2) °C. After two
weeks, the percentage of germination was 80%. The spore germination pattern corresponds to the Vittaria
type and the prothallus development was Ceratopteris type. Gametangia developed first in the bended
thalloid region of the prothallus and then bisexual tubercles originated near this zone. The sporophytes
developed only in association with the tubercles. During culture in plastic pots the sporophytes gave origin
to a second generation of prothalli. Our preliminary findings demonstrate that the in vitro culture technique
is suitable for A. chaerophylla propagation as a strategy for ex-situ conservation.
... Separate analyses were conducted for each of the five genera indigenous to New Zealand. For each genus, the placement of the New Zealand species was determined within a strongly supported subset of the Pteridaceae, by using previous studies (e.g. Nakazato and Gastony 2003; Kirkpatrick 2007; Prado et al. 2007; Zhang et al. 2007; Rothfels et al. 2008) as a guide. Relevant accessions from GenBank were selected from the aforementioned studies and via NCBI BLAST (http://blast.ncbi.nlm.nih.gov/) ...
... Our rbcL sequence for A. capillus-veneris, from a plant naturalised in New Zealand, was identical to those reported by Zhang et al. (2007) from a plant cultivated in China and by Hasebe et al. (1993) from a plant cultivated in Japan; only a single representative was included in our analyses. For Anogramma, rbcL sequence for a sample of A. leptophylla from New Zealand's South Island had been analysed previously by Nakazato and Gastony (2003). We compared our North Island sample with it, as well as with other sequences from their Clade C. For Cheilanthes, we focussed on the hemionitidoid clade of Rothfels et al. (2008). ...
Previous molecular phylogenetic studies of the species-rich Pteridaceae fern family have revealed that many of its constituent genera are not monophyletic. Within this context, we generated rbcL chloroplast DNA sequences for the 17 Pteridaceae species indigenous to New Zealand to assess how they are related to the type species of their genus. Of the five genera presently recognised in New Zealand, no taxonomic change is needed for the species of Anogramma, or, probably, of Cheilanthes and Pellaea. In contrast, most species presently attributed to Pteris, including those in New Zealand, probably do not belong there. The status of Adiantum remains unclear, although the New Zealand species are not especially closely related to the type species. The Adiantum species in New Zealand belong to a wide-ranging, principally southern hemisphere, clade that appears to be pivotal to resolving the relationships of Adiantum, although it has been little sampled. The closest relatives of the New Zealand species are in Australia and South America for Cheilanthes, Australia and South-east Asia for Pellaea, and the south-western Pacific for both of the Pteris lineages in New Zealand, whereas Anogramma leptophylla (L.) Link is subcosmopolitan.
... The genus Anogramma was circumscribed sensu Nakazato and Gastony (2003) in PPG I (2016) and this is followed here; however, this circumscription has been shown to be polyphyletic Hennequin et al. 2014). As circumscribed here, Anogramma includes~5 species, with the type species being widespread in Australia (1833) Type: Pityrogramma chrysophylla (Sw.) ...
The classification and typification of all Australian ferns and lycophytes is updated to reflect the Pteridophyte Phylogeny Group I classification and the International Code of Nomenclature for algae, fungi, and plants, presenting 8 new nomenclatural combinations as well as 85 lectotypifications. The Australian fern and lycophyte flora comprises 2 classes, 14 orders, 32 families, 134 genera and 528 species and subspecies with the addition of 8 newly recorded and 6 newly recognised species since the publication of the Flora of Australia fern volume in 1998. Overall, 208 species are endemic to Australia, with Queensland having the highest species diversity and endemism by state or territory, and Lord Howe Island having the highest concentration of species and endemics per unit area. The Australian fern and lycophyte flora shows diverse links with Africa, Asia and Oceania, with the largest overlaps being shared with Asia and Oceania. More species are endemic to Australia+Oceania than to Australia+Asia. Contrasting with the classification presented in the Flora of Australia, no genera of ferns and lycophytes are now considered to be wholly endemic to Australia.
... * Anogramma Link Persisten dudas sobre el carácter mono- filético de este género (Nakazato & Gastony 2003). ...
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.
... Notes:-Recognized within the genus by having rhizomes and usually the bases of the petioles with hairs and scales (1-1.5 × 0.1 mm); blades ovate to deltate, fairly large, to 40 cm long; segments mostly with acute lobes; and spores whitish to tan. Phylogenetic studies have shown that this species is more closely related, and sister, to Pityrogramma than to the other species of Anogramma ( Nakazato & Gastony 2003), from which it is well removed. However, a taxonomic reorganization of the genus has not yet been proposed. ...
We provide a synopsis to the Pteridaceae s.l. in Bolivia, a family comprising four subfamilies, 27 genera, and 154 species. These totals, approximating those for Dryopteridaceae and Polypodiaceae, make this one of the most genus-and species-rich families in Bolivia. New combinations are made for two species of Argyrochosma: A. flava and A. tenera. Eight species of Pteridaceae are considered endemic: Adiantum solomonii, A. squamulosum, Cheilanthes glutinosa, Eriosorus angustus, E. ascendens, E. madidiensis, Pteris boliviensis, and P. lellingeri.
... KEY WORDS.-anatomy, gametophyte, embryo, adaptations, Anogramma Anogramma Link (Pteridaceae) is a homosporous fern genus with seven species distributed mainly in the tropics of Mexico and Central and South America and less frequently in Africa, from southern Europe to northern India, and in Australia and New Zealand (Tryon et al., 1990;Nakazato and Gastony, 2003). In Argentina, Anogramma is represented by two species: Anogramma chaerophylla (Desv.) ...
The fern genus Anogramma is characterized by the presence of annual sporophytes and tubercles that persist through dry periods. Tubercles may host embryos that develop when climatic conditions are more favorable. As information on the structure of the tubercle and sporophyte of Anogramma chaerophylla is incomplete, the objective of this work was to analyze anatomical characteristics during development and deepen knowledge of the adaptive strategies of this species. Spores were grown in-vitro and different stages of tubercle development and embryonic sporophytes were fixed in FAA, embedded in Paraplast and analyzed using light microscopy. Initially, the green gametophyte developed antheridia and archegonia in its thickened portion, where later bisexual tubercles differentiated. Embryos developed only from the archegonia located in the tubercles, with one embryo developing per tubercule. During sporophyte growth the reserves accumulated in the tubercle were consumed. Oversized cells were observed at the junction between the green gametophyte and the tubercle, suggesting a possible role in translocation of substances. The young sporophyte consisted of a short shoot and a prominent foot. The placenta comprised the foot cells and the adjacent tubercle cells. The first leaves protruded soon and developed early as photosynthetic organs. Sporophytes remained attached to the tubercles until advanced stages of development. Vegetative propagation was documented in smaller tubercles that did not develop gametangia. Because few sporophytes were observed in vivo, it is likely that natural populations are maintained through vegetative propagation of the gametophyte until favorable conditions encourage development of sporophytes.
... A. leptophylla is a cosmopolitan species restricted to temperate and tropical zones (Landolt 2010), with preference for humid regions, usually with alternating wet and dry seasons (Nakazato andGastony 2003, Studnička 2009). In European context, it is considered an oceanic-suboceanic (Meusel et al. 1965, Dostál 1984 or oceanic-submediterranean species (Fischer et al. 2008). ...
Anogramma leptophylla is one of the rarest fern species in Balkan Peninsula. In Croatia, several localities were noted prior to this study, when its presence was confirmed with a discovery of a small population on the island of Mljet (Southern Adriatic). This was, after almost 80 years, the first reliable finding of this species along Eastern Adriatic. The establishment of A. leptophylla on the western part of the island of Mljet may be attributable to certain favourable environmental conditions, but essentially to higher air and soil humidity. Its unusual bryophyte-like life strategy, with short-living annual sporophytes, facilitates its survival under Mediterranean climate, generally unfavourable for pteridophytes.
... While there is reasonable phylogenetic knowledge available today and many significant changes have been made in the classification of ferns and lycophytes, the following nonexhaustive list of studies (with significant neotropical sampling) shows that few present a thorough sampling of studied taxa: Anemiaceae À Labiak et al., 2015; Aspleniaceae À Murakami & Schaal, 1994; Murakami et al., 1999; Schneider et al., 2004a; Dyer et al., 2012; Athyriaceae À Mynssen et al., 2016; Cibotiaceae À Geiger et al., 2013; Korall et al., 2006; Culcitaceae and Lophosoriaceae À Korall et al., 2006; Cyatheaceae À Korall et al., 2007; Korall & Pryer, 2014; Wolf et al., 1999; Dennstaedtiaceae À Wolf et al., 1994; Dryopteridaceae À Little & Barrington, 2003; Skog et al., 2004; Vasco et al., 2009; Moran et al., 2010; Guillou, 2011, unpublished data; Sessa et al., 2012; L origa et al., 2014; Labiak et al., 2014; McHenry & Barrington, 2014; Moran & Labiak, 2015; Vasco et al., 2015; Liu et al., 2016; Moran & Labiak, 2016; Equisetaceae À Des Marais et al., 2003; Hymenophyllaceae À Dubuisson et al., 2003; Hennequin et al., 2003; Isoetaceae À Hoot et al., 2006 ; Lindsaeaceae À Lehtonen et al., 2010; Lomariopsidaceae À Rouhan et al., 2007; Hennequin et al., 2010; Christenhusz et al., 2013; Lycopodiaceae À Wikstr€ om & Kenrick, 2000aKenrick, , 2000b Field et al., 2016; Lygodiaceae À Madeira et al., 2008; Marattiaceae À Christenhusz et al., 2008; Murdock, 2008; Marsileaceae À Nagalingum et al., 2007; Metaxyaceae À C ardenas et al., 2016; Ophioglossaceae À Hauk et al., 2003; Osmundaceae À Yatabe et al., 1999; Metzgar et al., 2008; Polypodiaceae À Schneider et al., 2004b; Schneider et al., 2006; Salino et al., 2008; Lehnert et al., 2009; Otto et al., 2009; Labiak et al., 2010a Labiak et al., , 2010b Sundue et al., 2010 Sundue et al., , 2012 Sundue et al., , 2014 Hirai et al., 2011; Sprunt et al., 2011; Assis et al., 2016; Pteridaceae À Crane et al., 1995; Nakazato & Gastony, 2003; S anchez-Baracaldo, 2004a S anchez-Baracaldo, , 2004b Kirkpatrick, 2007; Prado et al., 2007; Schuettpelz et al., 2007; Beck et al., 2010; Link-P erez et al., 2011; Sigel et al., 2011; Li et al., 2012; Grusz & Windham, 2013; Cochran et al., 2014; Pab on-Mora & Gonz alez, 2015; Yesilyurt et al., 2015; Pryer et al., 2016; Salviniaceae À Reid et al., 2006; Metzgar et al., 2007; Nagalingum et al., 2008; Schizaeaceae À Wikstr€ om et al., 2002; Selaginellaceae À Korall et al., 1999; Korall & Kenrick, 2004; Tectariaceae À Moran et al., 2014a; Thelypteridaceae À Almeida et al., 2016. So far, most studies have been using data from first generation sequencing (Sanger Sequencing) to create phylogenetic hypotheses and there still is much work to be done. ...
For ferns and lycophytes, the Neotropics is a hotspot of diversity (3000-4500 species), and second only to Southeastern Asia in richness and endemism. This paper presents the current state of knowledge on fern and lycophyte systematics in the Neotropics, and emphasizes sampling sufficiency and current taxonomic and phylogenetic knowledge. Plant systematics plays an important role in documenting diversity and geographic distribution patterns that are needed to understand relationships and evolutionary patterns, and a vital role in species conservation. Although in recent decades this field of science has undergone a revolution because of new approaches and techniques, data presented in this work shows that large gaps remain and there is still a long path towards fully understanding fern and lycophyte systematics in Neotropics. Approaches and how to choose areas that should be targeted in order to try to fulfill these knowledge gaps are discussed.
... 50-60 genera (53 accepted here). References: Beck et al. (2010), Gastony & Johnson (2001), Gastony & Rollo (1995, 1998, Grusz et al. (2009), Kirkpatrick (2007), Nakazato & Gastony (2001), Prado et al. (2007), Rohfelds et al. (2008), Ruhfel et al. (2008), Sánchez-Barracaldo (2004), , Windham et al. (2009), Zhang et al. (2005Zhang et al. ( , 2007. ...
... Nephopteris possesses a number of the traits given by Tryon et al. (1990) to the subfamily Taenitidoideae that include the following: small, terrestrial plants with creeping, pubescent rhizomes, pinnae with free flabellate venation, sterile pinnae glabrous, fertile pinnae densely pubescent beneath; sporangia borne along the veins, arranged in soral lines, lacking an indusium and intermixed with long paraphyses, and trilete spores with prominent ridges and an equatorial flange. Although the genus has been considered closely related to Eriosorus since it was originally described (Lellinger, 1966;Tryon et al., 1990), this relationship still remains to be tested in a phylogenetic analysis, since recent studies involving Taenitidoidean genera did not include samples of Nephopteris (e.g., Nakazato & Gastony, 2003;Sánchez-Baracaldo, 2004;Schuettpelz et al., 2007;Cochran et al., 2014). According to Archangelsky & Villar de Seoane (1998), the spores of N. maxonii show affinities with those of the fossil genus Muricingulisporis, from the Baquero Formation of the Cretaceous of Argentina. ...
The monospecific fern genus Nephopteris has previously been collected in three subparamos of Colombia, one in the Western Cordillera, and two in the Eastern Cordillera. Only five collections are available, the most recent having been made in 1973. The genus was thought to be extinct because the three subparamos where it has been found have undergone strong anthropogenic disturbance during the last five decades. Recently, we rediscovered N. maxonii in the Central and Eastern Cordilleras. Here we report detailed observations of this poorly known genus, revise its original description, and provide the first in vivo photographs of the species. Of the anatomical traits here examined, the siphonostelic to dictyostelic rhizomes with hairs and the presence of paraphyses along the soral lines support the traditional placement of Nephopteris in the subfamily Taenitidoideae of the Pteridaceae. In addition, we propose that this taxon should be included as Critically Endangered (CR), following the IUCN Red List criteria, based on the small population size, the extreme geographical isolation of the known locations, the narrow elevational belt occupied by the species in cloud subparamos in Colombia, and the increasing disturbance of this severely fragmented and fragile habitat.
... -The study used sequences from two plastid loci: rbcL and atpA. These genes were selected because both have been used successfully to reconstruct phylogenies in other fern lineages Nakazato and Gastony, 2003;Pryer et al., 2004;Schuettpelz et al., 2006;Tsutsumi and Kato, 2005;Wolf, 1995;Wolf et al., 1994). The rbcL gene evolves relatively slowly and is good for inferring phylogeny at the generic and familial level in ferns (Hasebe et al., 1995), as well as at lower taxonomic levels Haufler et al., 1996;Wolf et al., 1994). ...
Of the five major clades in the ecologically diverse Pteridaceae, the cheilanthoid ferns, comprised of approximately 20 genera and 400 species, are most obviously characterized by adaptations to xeric habitats. The cheilanthoids are known to be replete with paraphyletic genera due to the difficulties of recognizing monophyletic groups due to homoplasious adaptive morphology. The objectives of this dissertation were to complete a revision and explore the systematics of the Central and South American species of Adiantopsis Fée, a small neotropical genus in the hemionitid subclade within the cheilanthoids. Classically, Adiantopsis has been characterized by the combination of echinate spores, golden or golden-red paired carinae on the upper side of leaf axes and sometimes on stipe axes, and well-differentiated pseudoindusia. Laminar morphology is one of the more notable aspects of Adiantopsis because its species display palmate, pedate, and pinnate architectures. Using both traditional morphological analyses and molecular phylogenetics, this dissertation investigates i) the monophyly of Adiantopsis and its generic boundaries, ii) the enumeration of its species, iii) the history of character evolution in Adiantopsis, particularly the origin and evolution of the diverse laminar morphologies, and iv) uses a molecular phylogeny as a framework to investigate its biogeographic history. Phylogenetic analysis of combined plastid rbcL and atpA DNA sequences revealed that Adiantopsis is not monophyletic as traditionally circumscribed; with an expansion of generic boundaries and the transfer to Adiantopsis of three Cheilanthes species, Adiantopsis becomes monophyletic with strong support. Morphological characters to circumscribe the expanded Adiantopsis include: large, reddish, pluricellular hairs or carinae on axes; distinct, scarious pseudoindusia that cover one to occasionally two sori; and compound leaves with small, asymmetrical ultimate segments, at least some of which are stalked. Previously unrecognized diversity was revealed, particularly among palmate taxa: three new species are described. Pinnate architecture appears to be pleisiomorphic in Adiantopsis and the palmate architecture arose just once. Biogeographic analyses suggest an origin for the genus in South America, possibly the Cerrado and associated dry areas of Brazil, with a minimum of three migrations into the Caribbean.
... Since the advent of DNA sequencing, the vast majority of new combinations have resulted from the discovery that genera long thought to be natural are not monophyletic. However, in some cases new molecular data serve instead to illuminate historical oversights that have been carried over to the present (Nakazato & Gastony, 2003;Moran & al., 2008). In the past, such misplaced species were occasionally identified through careful morphological reassessment (Smith, 1976); DNA sequencing has simply enhanced our ability to find them. ...
Adiantum senae, a diminutive fern nearly restricted to southern Brazil, was described by Baker in 1885. In subsequent Floras, it was either completely overlooked or simply retained in the maidenhair genus Adiantum. However, a molecular phylogenetic analysis and a thorough morphological evaluation clearly demonstrate that this species does not belong in this genus. We find strong molecular support for the inclusion of A. senae within the cheilanthoid genus Adiantopsis, as a rather isolated lineage resolved as sister to the large, generally echinate-spored clade. The position of sporangia in A. senae—along the vein tips of the main laminar tissue rather than on the abaxial tissue of the pseudoindusia—confirms the separation of this species from Adiantum, and the presence of arachnoid-echinulate spores reinforces its resolved position within Adiantopsis. Based on the sum of this evidence, we here provide a new combination, Adiantopsis senae (Baker) Schuettp. & A.Davila. Additionally, we designate a lectotype for A. senae, as well as for a later synonym.
... Recent molecular phylogenetic analyses of cheilanthoid and related fern groups have used rbcL sequences (e.g., Gastony & Rollo, 1995). This marker was also used to infer phylogenetic relationships among species of Anogramma (Pteridaceae-Taenitidoideae) and between Anogramma and putative relatives, and to determine generic relationships and subfamilial placement of Cosentinia vellea (Nakazato & Gastony, 2003). Sánchez-Baracaldo (2004a), based on sequence data of the nuclear external transcribed spacer (ETS), plastid gene rps4 and intergenic spacer rps4-trnS, investigated the relationships among Jamesonia and Eriosorus (Pteridaceae-Taenitidoideae ), two traditionally recognized fern genera in the Neotropics that together form a monophyletic group. ...
Phylogenetic relationships among Pteridaceae were established using rbcL sequences, parsimony and pos-terior probabilities. The analyses involved 38 Pteridaceae species native in Brazil (12 of them endemic) and 81 species of Dennstaedtiaceae, Lindsaeaceae, Saccolomataceae (outgroups) and Pteridaceae. The resultant phylogeny comprehends five main clades: Platyzomatoideae-Pteridoideae-Taenitoideae; Ceratopteris-Acros-tichum; Adiantoideae-vittarioids; Cheilanthoideae; Coniogramme-Cryptogramma-Llavea. The cladograms support the most recent classification of Pteridaceae and demonstrate the paraphyly of Cheilanthoideae and the unnaturalness of Ceratopteridoideae, Platyzomatoideae, Pteridoideae, and Taenitidoideae as traditionally defined. Adiantoideae can only be recognized if combined with the vittarioids. Several genera of Pteridaceae appear to be paraphyletic (e.g., as Cheilanthes, Doryopteris, Pellaea, Pteris), and new generic affinities suggested by consistent internal clades are proposed.
... Contrary to its general appearance and its long history of treatment under Cheilanthes or Notholaena, this species is not closely related to Notholaena or any Cheilanthes group (Rothfels & al. 2008). Rather, it is closer to species of Anogramma and Pteris (Nakazato & Gastony 2003). Conversely, the superficially similar Cheilanthes acrostica is more closely related (evolutionarily) to Adiantum than it is to Cosentinia. ...
We present a list of ferns, lichens, mosses, and liverworts collected in the eastern UAE and northern Oman in January 2009. Of the seven fern species reported, Cosentinia vellea is confirmed for the UAE and Asplenium ceterach is reported for Oman for the second time. We briefly summarise the taxonomic and nomenclatural reasons for using the names Asplenium ceterach, Cheilanthes acrostica, and Cosentinia vellea for our collections, rather than the frequently used Ceterach officinarum, Cheilanthes pteridioides, and Cheilanthes vellea, respectively. Of the 12 lichen species reported, seven represent new records for Musandam, seven for Oman, and two for the Arabian Peninsula (Caloplaca inconnexa and Caloplaca pusilla). For the mosses, we report 13 species: seven new records for Musandam, nine for Oman, and three for Arabia (Anoectangium handelii, Orthotrichum cf. cupulatum, and Syntrichia sinensis). One of the liverworts is new for Oman (Riccia crozalsii). We include a brief description of the cryptogam flora of the UAE and northern Oman, the degree to which it is under-explored (five new records for Arabia in the course of extremely modest and relatively casual fieldwork by non-specialists), and avenues for future work.: Included in this article are records of the following species
... -The study used sequences from two plastid loci: rbcL and atpA. These genes were selected because both have been used successfully to reconstruct phylogenies in other fern lineages (Wolf & al., 1994;Wolf, 1995;Gastony & Rollo, 1996Nakazato & Gastony, 2003;Pryer & al., 2004;Tsutsumi & Kato, 2005;Schuettpelz & al., 2006;. The rbcL gene evolves relatively slowly and is good for inferring phylogeny at the generic and familial levels in ferns (Hasebe & al., 1995), as well as at lower taxonomic levels (Wolf & al., 1994;Haufler & al., 1996;Gastony & Rollo, 1998). ...
We present the first multi-gene phylogeny of taxa potentially allied to the fern genus Adiantopsis, one of about 20 genera in the large group of cheilanthoid ferns. Our goals were to clarify natural relationships, examine generic boundaries, and interpret character evolution within a phylogenetic framework. Phylogenetic analysis of combined plastid rbcL and atpA DNA sequences has led to an expansion of generic boundaries in Adiantopsis, a recircumscription that is clearly defensible with morphological data. Biogeographic analyses suggest an origin for the genus in South America, with a minimum of three migrations into the Caribbean. We include a synopsis of the genus and two new combinations.
... Molecular systematic studies in lycophytes and monilophytes (leptosporangiate ferns, eusporangiate ferns, Psilotaceae, and Equisetaceae; see Pryer et al., 2004) have generally relied on a subset of the sequences used in angiosperm systematics. The gene rbcL has been used extensively in studies for both higher-level and lower-level taxa (Dubuisson, 1997;Dubuisson et al., 1998Dubuisson et al., , 2003Gastony and Johnson, 2001;Geiger and Ranker, 2005;Hasebe et al., 1993Hasebe et al., , 1994Hasebe et al., , 1995HauXer and Ranker, 1995;HauXer et al., , 2003Hauk, 1995;Hauk et al., 2003;Hennequin et al., 2003;Kato and Setoguchi, 1998;Kenrick, 2002, 2004;Little and Barrington, 2003;Murakami et al., 1999;Nakazato and Gastony, 2003;Pinter et al., 2002;Pryer, 1999;Pryer et al., 2001aPryer et al., ,b, 2004Pryer et al., , 1995Ranker et al., 2003Ranker et al., , 2004Sano et al., 2000;Schneider et al., 2002Schneider et al., , 2004aSkog et al., 2004;Wolf, 1995;Wolf et al., 1999). Other genes such as atpB (Pryer et al., 2001a(Pryer et al., , 2004Ranker et al., 2003Ranker et al., , 2004Wolf, 1997) and rpS4 (Guillon, 2004;Hennequin et al., 2003;Pryer et al., 2001aPryer et al., , 2004Sanchez-Baracaldo, 2004a,b;Schneider et al., 2002Schneider et al., , 2004cSmith and CranWll, 2002) have also been employed. ...
Noncoding DNA sequences from numerous regions of the chloroplast genome have provided a significant source of characters for phylogenetic studies in seed plants. In lycophytes and monilophytes (leptosporangiate ferns, eusporangiate ferns, Psilotaceae, and Equisetaceae), on the other hand, relatively few noncoding chloroplast DNA regions have been explored. We screened 30 lycophyte and monilophyte species to determine the potential utility of PCR amplification primers for 18 noncoding chloroplast DNA regions that have previously been used in seed plant studies. Of these primer sets eight appear to be nearly universally capable of amplifying lycophyte and monilophyte DNAs, and an additional six are useful in at least some groups. To further explore the application of noncoding chloroplast DNA, we analyzed the relative phylogenetic utility of five cpDNA regions for resolving relationships in Botrychium s.l. (Ophioglossaceae). Previous studies have evaluated both the gene rbcL and the trnL(UAA)-trnF(GAA) intergenic spacer in this group. To these published data we added sequences of the trnS(GCU)-trnG(UUC) intergenic spacer + the trnG(UUC) intron region, the trnS(GGA)-rpS4 intergenic spacer+rpS4 gene, and the rpL16 intron. Both the trnS(GCU)-trnG(UUC) and rpL16 regions are highly variable in angiosperms and the trnS(GGA)-rpS4 region has been widely used in monilophyte phylogenetic studies. Phylogenetic resolution was equivalent across regions, but the strength of support for the phylogenies varied among regions. Of the five sampled regions the trnS(GCU)-trnG(UUC) spacer+trnG(UUC) intron region provided the strongest support for the inferred phylogeny.
... Molecular phylogenetic studies of the Pteridaceae, to date, have primarily focused on the relationships within a few segregates Rollo, 1995, 1998;Nakazato and Gastony, 2003;Sánchez-Baracaldo, 2004;Zhang et al., 2005). Our current understanding of overall Pteridaceae phylogeny mostly stems from these results combined with those from analyses aimed at leptosporangiate ferns as a whole, generally with only a few Pteridaceae exemplars and based on a single gene (Gastony and Johnson, 2001;Hasebe et al., 1995;Pryer et al., 1995). ...
The monophyletic Pteridaceae accounts for roughly 10% of extant fern diversity and occupies an unusually broad range of ecological niches, including terrestrial, epiphytic, xeric-adapted rupestral, and even aquatic species. In this study, we present the results of the first broad-scale and multi-gene phylogenetic analyses of these ferns, and determine the affinities of several previously unsampled genera. Our analyses of two newly assembled data sets (including 169 newly obtained sequences) resolve five major clades within the Pteridaceae: cryptogrammoids, ceratopteridoids, pteridoids, adiantoids, and cheilanthoids. Although the composition of these clades is in general agreement with earlier phylogenetic studies, it is very much at odds with the most recent subfamilial classification. Of the previously unsampled genera, two (Neurocallis and Ochropteris) are nested within the genus Pteris; two others (Monogramma and Rheopteris) are early diverging vittarioid ferns, with Monogramma resolved as polyphyletic; the last previously unsampled genus (Adiantopsis) occupies a rather derived position among cheilanthoids. Interestingly, some clades resolved within the Pteridaceae can be characterized by their ecological preferences, suggesting that the initial diversification in this family was tied to ecological innovation and specialization. These processes may well be the basis for the diversity and success of the Pteridaceae today.
Ferns are an integral part of the world's flora, appreciated for their beauty as ornamentals, problematic as invaders and endangered by human interference. They often dominate forest understories but also colonize open areas, invade waterways and survive in nutrient-poor wastelands and eroded pastures. Presented here is the first comprehensive summary of fern ecology, with worldwide examples from Siberia to the islands of Hawaii. Topics include a brief history of the ecological study of ferns, a global survey of fern biogeography, fern population dynamics, the role of ferns in ecosystem nutrient cycles, their adaptations to xeric environments and future directions in fern ecology. Fully illustrated concepts and processes provide a framework for future research and utilization of ferns for graduate students and professionals in ecology, conservation and land management.
Phylogeny has long informed pteridophyte classification. As our ability to infer evolutionary trees has improved, classifications aimed at recognizing natural groups have become increasingly predictive and stable. Here, we provide a modern, comprehensive classification for lycophytes and ferns, down to the genus level, utilizing a community-based approach. We use monophyly as the primary criterion for the recognition of taxa, but also aim to preserve existing taxa and circumscriptions that are both widely accepted and consistent with our understanding of pteridophyte phylogeny. In total, this classification treats an estimated 11 916 species in 337 genera, 51 families, 14 orders, and two classes. This classification is not intended as the final word on lycophyte and fern taxonomy, but rather a summary statement of current hypotheses, derived from the best available data and shaped by those most familiar with the plants in question. We hope that it will serve as a resource for those wanting references to the recent literature on pteridophyte phylogeny and classification, a framework for guiding future investigations, and a stimulus to further discourse
Abandoned croplands can be considered a new category of “scattered elements” of mountain landscapes. To gain a deeper understanding of the conservation status (sensu EEC Directive 92/43) of abandoned cropland in the northern Apennines, we used the concept of the social behavior type (SBT) of plant communities. SBTs refer to the behaviour and ecological attributes of species at a given observation level and allow the understanding of the plant community conservation status. We found that topographic and soil conditions drive species assemblage in pastures after crop abandonment, but that long-term abandonment does not lead per se to the recovery of the semi-natural grassland communities deemed worthy of conservation in the EEC Directive. It was mainly the lack of appropriate disturbance regimes that allowed the spread of dominant tall herbs, which, in turn, reduced site suitability for subordinate plants. Moreover, their spread fostered the presence of elements such as ruderals and fringe species. We concluded that these abandoned croplands had a good potential to develop into a Habitat as defined in the EU Directive but without appropriate management plans they would remain of low representativeness.
Introduction As a consequence of employing DNA sequence data and phylogenetic approaches, unprecedented progress has been made in recent years toward a full understanding of the fern tree of life. At the broadest level, molecular phylogenetic analyses have helped to elucidate which of the so-called “fern allies” are indeed ferns, and which are only distantly related (Nickrent et al., 2000; Pryer et al., 2001a; Wikström and Pryer, 2005; Qiu et al., 2006). Slightly more focused analyses have revealed the composition of, and relationships among, the major extant fern clades (Hasebe et al., 1995; Wolf, 1997; Pryer et al., 2004b; Schneider et al., 2004c; Schuettpelz et al., 2006; Schuettpelz and Pryer, 2007). A plethora of analyses, at an even finer scale, has uncovered some of the most detailed associations (numerous references cited below). Together, these studies have helped to answer many long-standing questions in fern systematics. In this chapter, a brief synopsis of vascular plant relationships - as currently understood - is initially provided to place ferns within a broader phylogenetic framework. This is followed by an overview of fern phylogeny, with most attention devoted to the leptosporangiate clade that accounts for the bulk of extant fern diversity. Discussion of finer scale relationships is generally avoided; instead, the reader is directed to the relevant literature, where more detailed information can be found. © Cambridge University Press 2008 and Cambridge University Press 2009.
The monospecific fern genus Nephopteris is restricted to four páramos in Colombia. Until now, its phylogenetic relationships have not been clarified, although morphological traits suggest that it belongs to the taenitidoid clade of the subfamily Pteridoideae in the Pteridaceae. We isolated four chloroplast (atpA, chlL, rbcL, and rps4) loci of N. maxonii and performed maximum parsimony and maximum likelihood phylogenetic analyses, including a preliminary analysis based on rbcL of 424 accessions of leptosporangiate ferns. Then, we carried out separate and combined reconstructions for all four loci, focusing the sampling in the subfamily Pteridoideae (Pteridaceae). Our results show that Nephopteris belongs to the taenitidoid clade of the subfamily Pteridoideae and is nested in Jamesonia. Thus, the new combination Jamesonia maxonii is provided. Its dimorphic sterile and fertile leaves appear to be unique within the taenitidoids. Given that this species shares the same microhabitat with other Jamesonia species, it is unlikely that leaf shape evolution within the entire clade is a result of adaptive processes to páramo conditions.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. ABSTRACT.—A new species, Pityrogramma opalescens, is described; it is known only from Cerro del Torrá, an isolated mountain peak in the Cordillera Occidental of Colombia, which is a region of high diversity and endemism. It is most similar to Pityrogramma lehmannii, and the two species differ from all other Pityrogramma species by having creeping rhizomes, rhizome scales with turgid cells, an elongate, proximally 1-pinnate lamina, with sessile segments supplied by multiple veins that emerge from the rachis rather than from a single main vein, and farina on the abaxial side of the lamina that is borne at the apex of short hairs. It differs from P. lehmannii by more numerous segments that are narrower (3.5–4.0 mm wide vs. 10.0–17.0 mm wide), with more acutely rounded apices, and dull-yellow rather than white farina abaxially.
In an effort to obtain a solid and balanced approximation of global fern phylogeny to serve as a tool for addressing large-scale evolutionary questions, we assembled and analyzed the most inclusive molecular dataset for leptosporangiate ferns to date. Three plastid genes (rbcL, atpB, atpA), totaling more than 4,000 bp, were sequenced for each of 400 leptosporangiate fern species (selected using a proportional sampling approach) and five outgroups. Maximum likelihood analysis of these data yielded an especially robust phylogeny: 80% of the nodes were supported by a maximum likelihood bootstrap percentage ≥ 70. The scope of our analysis provides unprecedented insight into overall fern relationships, not only delivering additional support for the deepest leptosporangiate divergences, but also uncovering the composition of more recently emerging clades and their relationships to one another.
Anogramma leptophylla is one of few homosporous ferns with ephemeral sporophytes and perennial gametophytic tubercules. This rare aspect of the life cycle has considerable bearing on the reproductive features of the gametophyte. Four populations in the Iberian Peninsula were selected to provide spores for laboratory cultures and to sample natural populations of gametophytes for comparative studies. Sexual expression and the role of the tubercule in the reproductive biology of this fern were studied. No significant differences were found between percentages of sexual versus asexual prothalli in natural populations or laboratory cultures, or between percentages of unisexual and bisexual prothalli. These results may indicate that sex in A. leptophylla is genetically determined instead of environmentally controlled as in most ferns. The presence of a gametophytic tubercule with nutrient reserves seems to be most favourable for the development of archegonia, and for the establishment of young sporophytes. This tubercule, which can resist stressful environmental conditions, allows for the development of new gametophytic outgrowths and of dormant embryos as soon as environmental conditions improve. This bryophyte-like behaviour gives this fern some advantages in colonisation and has probably contributed to its wide geographic range.
The presence of the fern species Anogramma leptophylla was detected in the Zempléni Mountains (NE Hungary) in 1991. The species was known neither from the country nor from the whole Carpatho-Pannonian Region (also known as Carpathian Basin) previously. Its habitat is situated on a roadside bank, cut into an unstable rhyolite surface, above the valley of the Creek Kemence near the village of Pálháza. The fern is a cosmopolitan taxon restricted to humid environments and is considered to be an oceanic-suboceanic (Atlantic) element in Europe. The occurrence in Hungary is located more than 1000 km from the closest populations, thus, this is one of the most remote inland occurrences of this (sub)oceanic species. This striking presence of the fern may be due to the peculiar microclimatological conditions of the habitat, which are described here in order to give an exact explanation for this outstanding occurrence. The chromosomes were also counted in some individuals of the Hungarian population and were found to be n = 26II for each sample.
Recent phylogenetic studies have revealed that the traditional concept of pteridophytes which includes lycopods and ferns should be revised and a new classification of the extant monilophytes has just been published by Smith et al. China is very rich in plant diversity, with representatives of most important major groups of the lycophytes and monilophytes. Here we present the recent progress in phylogenetic analyses of lycophytes and monilophytes with a focus on relationships among the Chinese taxa. We conducted a phylogenetic analysis using sequence data of the chloroplast rbcL gene of 184 species (179 genera) representing 62 of a total of 63 families recognized by R. C. Ching. The relationships of all major lineages recovered in the rbcL phylogeny generally agree with those reconstructed in the studies that focused on these clades individually and had more extensive ingroup taxon sampling and/or character sampling. A tentative systematic arrangement focused on the Chinese lycophytes and monilophytes at the family level is presented.
We present a revised classification for extant ferns, with emphasis on ordinal and familial ranks, and a synopsis of included genera. Our classification reflects recently published phylogenetic hypotheses based on both morphological and molecular data. Within our new classification, we recognize four monophyletic classes, 11 monophyletic orders, and 37 families, 32 of which are strongly supported as monophyletic. One new family, Cibotiaceae Korall, is described. The phylogenetic affinities of a few genera in the order Polypodiales are unclear and their familial placements are therefore tentative. Alphabetical lists of accepted genera (including common synonyms), families, orders, and taxa of higher rank are provided.
Introduction and historical summary Over the past 70 years, many fern classifications, nearly all based on morphology, most explicitly or implicitly phylogenetic, have been proposed. The most complete and commonly used classifications, some intended primarily as herbarium (filing) schemes, are summarized in Table 16.1, and include: Christensen (1938), Copeland (1947), Holttum (1947, 1949), Nayar (1970), Bierhorst (1971), Crabbe et al. (1975), Pichi Sermolli (1977), Ching (1978), Tryon and Tryon (1982), Kramer (in Kubitzki, 1990), Hennipman (1996), and Stevenson and Loconte (1996). Other classifications or trees implying relationships, some with a regional focus, include Bower (1926), Ching (1940), Dickason (1946), Wagner (1969), Tagawa and Iwatsuki (1972), Holttum (1973), and M.ckel (1974). Tryon (1952) and Pichi Sermolli (1973) reviewed and reproduced many of these and still earlier classifications, and Pichi Sermolli (1970, 1981, 1982, 1986) also summarized information on family names of ferns. Smith (1996) provided a summary and discussion of recent classifications. With the advent of cladistic methods and molecular sequencing techniques, there has been an increased interest in classifications reflecting evolutionary relationships. Phylogenetic studies robustly support a basal dichotomy within vascular plants, separating the lycophytes (less than 1% of extant vascular plants) from the euphyllophytes (Figure 16.1; Raubeson and Jansen, 1992, Kenrick and Crane, 1997; Pryer et al., 2001a, 2004a, 2004b; Qiu et al., 2006). Living euphyllophytes, in turn, comprise two major clades: spermatophytes (seed plants), which are in excess of 260000 species (Thorne, 2002; Scotland and Wortley, 2003), and ferns (sensu Pryer et al. 2004b), with about 9000 species, including horsetails, whisk ferns, and all eusporangiate and leptosporangiate ferns. © Cambridge University Press 2008 and Cambridge University Press 2009.
Nine species of "Cheilantoid ferns" are known to grow in Macaronesia and the Mediterranean basin. Two of them (lacking a pseudo-indusium and having the basic chromosome number X = 29), both aggregate species which we prefer to retain in Notholaena, are not included in this study. The other seven species (with distinct pseudo-indusium and the basic chromosome number X = 30), which we accept as members of the genus Cheilanthes Sw. sensu stricto, were subjected to detailed genome analysis of their natural and experimentally produced hybrids and shown to represent an aggregate of four very distinct ancestral diploids and three allotetraploids. The latter must have once been formed by chromosome doubling in the three diploid hybrids of C. maderensis Lowe with the other three diploid species. Theoretically three more allotetraploids would be possible but their formation has obviously been prevented by the geographical separation of the three respective diploids. The most widely distributed of the tetraploids, i.e. C. pteridioides (Reich.) C.Chr. has also been resynthesized from its ancestors (still sympatric) under experimental conditions. The intermediate morphology of the allotetraploids (as compared with their diploid ancestors) is obviously the reason why their status and existence has so long escaped recognition in Europe. These seven species form a natural group and, in our opinion, should not be divided into sections.