Miscellaneous notes on the fern family Blechnaceae (Polypodiopsida) from Tropical America

Abstract

We present here new discoveries in Blechnaceae (Polypodiopsida). There are new records in Austroblechnum, Blechnum, Cranfillia, and Parablechnum for different political units and areas of the Neotropics. These (and other) records are reported here, alongside with two new necessary combinations in Parablechnum and an updated key to the Brazilian species of the genus. Also, an update of Blechnum description was made.

Key words:
new records; nomenclature; Parablechnum; pteridophytes.

Resumo

São apresentadas novas descobertas em Blechnaceae (Polypodiopsida). Há novos registros em Austroblechnum, Blechnum, Cranfillia e Parablechnum para diferentes unidades políticas e áreas do Neotrópico. Esses (e outros) registros são reportados aqui, juntamente com duas novas combinações em Parablechnum e uma chave atualizada para as espécies brasileiras do gênero. Além disso, foi feita uma atualização da descrição de Blechnum.

Palavras-chave:
novos registros; nomenclatura; Parablechnum; pteridófitas.

Introduction

The family Blechnaceae was composed, until recently, by the large genus Blechnum and eight satellite genera, such as Pteridoblechnum Hennipman, Sadleria Kaulf., and Doodia R. Br. (Tryon & Tryon 1982; Kramer et al. 1990). There were evidences, however, during last decades, that Blechnum was non-monophyletic (Cranfill & Kato 2003; Schuettpelz & Pryer 2007; Shepherd et al. 2007; Rothfels et al. 2012; Gabriel y Galán et al. 2013; Perrie et al. 2014). So, a new classification proposed by Gasper et al. (2016, 2017), adopted by PPG I (2016), recognizes 24 genera, and a new genus proposed by Molino et al. (2019) expands the number to 25. The largest genus is Parablechnum C. Presl, followed by Austroblechnum Gasper & V.A.O. Dittrich and Blechnum. In the last years, many works have been conducted about the morphology, taxonomy, nomenclature, and phylogeny of the family (e.g., Chambers & Farrant 2001; Kessler et al. 2007; Shepherd et al. 2007; Rolleri et al. 2010; Dittrich et al. 2012; Gabriel y Galán 2013; Cárdenas et al. 2019; Bauret et al. 2020; Molino et al. 2020), and many more are to be expected, as researchers in Europe, South America, and Oceania are working now with the group.

Since Gasper et al. (2016, 2017), we found out that some new combinations were necessary. So, here we provide new information about the nomenclature, taxonomy, and distribution of the family Blechnaceae in tropical America.

Material and Methods

During visits to the herbaria BM, K, P, PR, S and RB (abbreviations follow Thiers, continuously updated) we found new records for a few species. We also analyzed Blechnum species and realized that a new description of the genus is needed, since a species until now considered as Blechnum was moved to Parablechnum. For Parablechnum, new records were found in Brazil, so a new key to the Brazilian species in the genus is provided, based exclusively on Brazilian samples.

Results and Discussion

New combinations and description

1. Parablechnum striatum (Sw.) Gasper & V.A.O. Dittrich, comb. nov. - Onoclea striataSwartz (1806: 304, 422). Type: Martinica, St. Kitts. Synonyms: Lomaria ryaniiKaulfuss (1824: 155). Blechnum ryanii (Kaulf.) Hieronymus (1908: 245). Parablechnum ryanii (Kaulf.) Gasper & Salino in Gasper et al. (2016: 217).

Gasper et al. (2016) combined the name Lomaria ryanii under Parablechnum (Parablechnum ryanii). Notwithstanding, in the Fern Flora of the Lesser Antilles (Proctor 1977) Blechnum striatum (Sw.) Christensen (1906: 160), and the name on which it is based, Onoclea striata Sw., are listed as synonyms of Blechnum ryanii. However, Onoclea striataSwartz (1806) is older than Lomaria ryaniiKaulfuss (1824). Before the transfer of these names to Parablechnum, there was a Blechnum striatumBrown (1810: 152), based on a type from Australia, that impeded the transfer of Onoclea striata to Blechnum. In Parablechnum there is no such impediment, so the appropriate name for these plants must be Parablechnum striatum, not Parablechnum ryanii.

2. Parablechnum anthracinum (R.C. Moran) Gasper & V.A.O. Dittrich, comb. nov. - Blechnum anthracinumMoran (1992: 132).

When we published the new classification of Blechnaceae some years ago (Gasper et al. 2016), there was one species of Blechnum quite odd in the genus. It is a plant with very scaly rachises, pinnae weakly revolute, stipes and rachises atropurpureous and short-stalked or sessile pinnae throughout the blade, features rare or absent in Blechnum s.s. (as circumscribed in the aforementioned works). Kessler et al. (2007) argue that B. anthracinum belongs to the Blechnum occidentale L. group, despite the blackish, creeping rhizomes and the supposed epiphytic habit. Rolleri et al. (2012) believe that B. anthracinum is closely related to a hemiepiphytic group now known as Lomaridium C. Presl. Some observations about the relationships of this species were made by Moran (1992), who highlighted the presence of strongly denticulate scales. According to him, B. anthracinum is related to B. occidentale and B. glandulosum Kaulf. exLink (1822: 462) (= B. occidentale), but he has also suggested a relationship with Parablechnum stipitellatum (Sodiro) Gasper & Salino inGasper et al. (2016: 217) (as Blechnum stipitellatum Sodiro (Christensen) (1906: 160). Recently, Silva et al. (2021) noted that the spores of this species are quite distinct from spores of Blechnum, which usually have perine with or without crests, whilst P. anthracinum has cristate spores, with filaments and perforations. Therefore, it is now clear that it was not a Blechnum species, but rather a Parablechnum with monomorphic fronds, a rare feature in Parablechnum, but seen in other species such as Parablechnum monomorphum (Moran & Øllgaard 1995: 177) Gasper & Salino in Gasper et al. (2016: 216) and Parablechnum obtusum (Moran & Smith 2005: 237) Gasper & Salino in Gasper et al. (2016: 217). Consequently, we transfer this name to Parablechnum and provide a new description of Blechnum.

3. Blechnum.

Type: Blechnum occidentaleLinnaeus (1753: 1077) (as Blechnum orientale).

Plants terrestrial or epipetric;rhizomes erect to decumbent, stoloniferous, slender to rather stout, bearing linear, linear-oblong, triangular, narrowly triangular, lanceolate or linear-lanceolate, pale or dark brown, concolorous or bicolorous, entire scales (or scales with few teeth); fronds monomorphic or subdimorphic; stipes slender, short (sometimes absent) or long, stramineous to pale brown, with scales similar to those of rhizomes proximally, and sometimes with multicellular, catenate, hyaline hairs toward apices; blades concolorous, lanceolate to deltate-lanceolate, entire (rare), pinnatisect or 1-pinnate, gradually reduced or truncate proximally, apices pinnatifid or conform; rachises glabrous or pilose, without scales; buds absent; aerophores absent; pinnae sessile, adnate or subpetiolulate, lanceolate, oblong-lanceolate or ensiform, margins entire, finely denticulate; veins rarely partially anastomosing, usually free and then 1-3 furcate, each ending near margins in an enlarged tip; sori linear, continuous or rarely interrupted along the costae, indusia slightly erose or ciliate; spores reniform, exine usually smooth, without verrucae or ridges; x = 31.

New records

1. Austroblechnum organense (Brade 1935: 2) Gasper & V.A.O. Dittrich in Gasper et al. (2016: 203).

This species is so far known from Brazil, Venezuela, and Colombia (Dittrich et al. 2017). This is the first record to Ecuador: Bolívar, road Chillanes - El Tambo, 1,700-2,300 m, 19.VII.1991, H. van der Werff, B. Gray & G. Tipas 12,476 (QCNE - photo!).

2. Blechnum × caudatumCavanilles (1802: 262).

This hybrid is known from Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Suriname, Ecuador, Peru, Bolivia, and southeastern/southern Brazil. This is the first record for northern Brazil: Brazil, Amazonas, Barcelos, Vicinity of Pico Rondon, Perimetral Norte. Highway km 211, southern ridge of Pico Rondon, 700 m, 1°32’N, 62°48’W, 3.II.1984, G.T. Prance et al. 28805 (INPA , NY00814006 - photos! at <https://specieslink.net/> and <http://floradobrasil.jbrj.gov.br/reflora/herbarioVirtual/ConsultaPublicoHVUC/ConsultaPublicoHVUC.do>;).

3. Blechnum × leopoldense (Dutra 1938: 36) V.A.O. Dittrich & Salino in Dittrich et al. (2015: 217).

This hybrid is known from northwestern Argentina and southern Brazil (Sehnem 1968; Sota 1973; Dittrich et al. 2015). This is a new record to Uruguay: Montevideo, 18.VI.1876, C. Fruchard (P01535236).

4. Cranfillia caudata (Baker in Hooker & Baker 1867: 179) V.A.O. Dittrich & Gasper in Dittrich et al. (2017: 13).

This species is so far recorded, in Brazil, just for Minas Gerais state (Dittrich et al. 2017, 2020) and is considered vulnerable by Dittrich et al. (2013). Here it is cited for the states of São Paulo and Rio de Janeiro and for Serra do Mar range for the first time: São Paulo, São José do Barreiro, trilha ao lado da Pousada Chão de Estrelas, 22°42’59”S, 44°36’41”W, 5.X.2006, C.M. Mynssen et al. 1045 (RB00645603); Rio de Janeiro, Nova Friburgo, Parque Estadual dos Três Picos, Três Picos, Vale dos Deuses, trilha para a Cabeça do Dragão, 22°19’23.3”S, 42°43’12.8”W, 21.VI.2014, E.R. Damasceno & I.C.G. Campos 844 (R - photo).

5. Parablechnum atropurpureum (Smith in Steyermark & Maguire 1984: 5) Gasper & Salino in Gasper et al. (2016: 215).

This species is so far known only from Venezuela (Smith 1995). Here it is recorded for the first time in Brazil (Fig. 1a): Roraima, 1,800 m, II.1910, E. Ule 8509 (K, B200034238 - photo at <https://specieslink.net/>;).

Figure 1
a. Parablechnum atropurpureum from Roraima, Brazil. b-c. Parablechnum lechleri from Bahia, Brazil. [a. collected by E. Ule 8509 (B200034238); b-c. collected by T. Laessoe & T. Silva H 53301 (K)].

6. Parablechnum lechleri (Mettenius 1859: 17) Gasper & Salino in Gasper et al. (2016: 216).

This species is so far known for the Lesser Antilles, the Guayana Highlands and the Andes from Venezuela to Bolivia (Smith & Kessler 2018). Here it is recorded in Brazil for the first time (Fig. 1b-c): Brazil, Bahia, Abaíra, Mata do Cigano, 13°15’N, 41°55’W, 1,850 m, 22.III.1992, T. Laessoe & T. Silva H 53301 (K). This record was made concurrently by Molino (2022) while this work was under submission.

7. Parablechnum loxense (Kunth in Humboldt et al. 1815: 18) Gasper & Salino in Gasper et al. (2016: 216).

This taxon is so far known for Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, and Bolivia (Tryon & Stolze 1993; Smith & Kessler 2018). Here it is recorded in Chile for the first time: Chile, sine loco, without date, Anon. s.n., P01462420 (P, Herbarium Richard). This record was made concurrently by Molino (2022) while this work was under submission.

8. Parablechnum obtusum (Moran & Smith 2005: 237) Gasper & Salino in Gasper et al. (2016: 217).

This taxon was described from Venezuela and is so far known exclusively from this South American country. Here we report two collections from Colombia: Cauca, Páramo de Puracé, Central Cordillera, 3,500 m, 26.V.1944, E.P. Killip & F.C. Lehmann 38591 (US barcode 01525151, photo! at <https://collections.nmnh.si.edu/search/botany/>;). Caldas, Carretera entre Manizales y el Hotel Termales del Ruiz, 3,000-3,500 m, 8.VI.1966, E. Forero et al. 526 (F C0644727F, NY04187103, photos! at <http://inct.splink.org.br/>;). This record was made concurrently by Molino (2022) while this work was under submission.

9. Parablechnum rheophyticum (Moran in Moran & Øllgaard: 52) Gasper & Salino in Gasper et al. (2016: 217).

This taxon was described from Ecuador and was never mentioned to other countries. Here we report it to Colombia for the first time: Colombia, Nariño, Mpio. de Ricaurte. Resguardo Indígena Nulpe Medio, Orillas del Río Nulpe, 01°06’N, 78°13’W, 715 m, 6.I.1996, B.R. Ramírez & M.S. González 9438 (UC1732585 - photo! at <https://pteridoportal.org/portal/collections/harvestparams.php>;).

Specimen citation (correction)

In a previous paper published by us (Dittrich et al. 2018) we have cited Parablechnum cordatum (Desvaux 1811: 330) Gasper & Salino in Gasper et al. (2016: 216) for the first time in Guyana but we did not cite the specimen seen. Here is the correct citation of it: Guyana, Roraima, 1863-1864, C.F. Appun 1154 (K).

Updated key to the Brazilian species of Parablechnum, based on Brazilian specimens

• 1. Conical aerophores present at the base of stipes 2

• 1’. Conical aerophores never present at the base of stipes (tuberculate aerophores may be present) 3

  • 2. Bases of distal pinnae with bulbils Parablechnum proliferum

  • 2’. Bases of pinnae without bulbils Parablechnum lechleri

    • 3. Pinnae clearly revolute 4

    • 3’. Pinnae flat (or plane) 5

      • 4. Pinna ascending (generally strongly), rachises stramineous; plants from SE and S Brazil Parablechnum usterianum

      • 4’. Pinnae patent to weakly ascending, rachises atropurpureous; plants from N Brazil (Roraima state) Parablechnum atropurpureum

        • 5. Pinnae bases cordate or subcordate Parablechnum cordatum

        • 5’. Pinnae bases cuneate Parablechnum glaziovii

Acknowledgements

The senior author acknowledges the CNPq/Reflora program for financial support to visit European herbaria; and we acknowledge Alan R. Smith for calling our attention to the names related to Parablechnum ryanii; and the curators of all cited herbaria. ALG thanks CNPq for the productivity grant (311303/2020-0).

Data availability statement

In accordance with Open Science communication practices, the authors inform that all data are available within the manuscript.

References

• Bauret L, Vázquez R, Molino S, Gaudeul M, Rakotondrainibe F, Gasper AL & Rouhan G (2020) New molecular and morphological evidences favor a combination of Blechnum bakeri C. Chr. in Cranfillia Gasper & V.A.O. Dittrich (Blechnaceae, Polypodiopsida), thus extending the distribution of Cranfillia to Madagascar and East Africa. Adansonia 42: 279-289. DOI: 10.5252/adansonia2020v42a18
  » https://doi.org/10.5252/adansonia2020v42a18
• Brade AC (1935) Filices Novae Brasilianae IV. Arquivos do Instituto de Biologia Vegetal 2: 1-5.
• Brown R (1810) Roberti Brownii Prodromus florae Novae Hollandiae et Insulae Van Diemen. Veneunt a reductione Isidis, Lipsiae. 590p. DOI: 10.5962/bhl.title.3633
  » https://doi.org/10.5962/bhl.title.3633
• Cárdenas GG, Lehtonen S & Tuomisto H (2019) Taxonomy and evolutionary history of the neotropical fern genus Salpichlaena (Blechnaceae). Blumea 64: 1-22. DOI: 10.3767/blumea.2018.64.01.01
  » https://doi.org/10.3767/blumea.2018.64.01.01
• Cavanilles AC (1802) Descripción de las plantas que Antonio Josef Cavanilles demostró en las lecciones públicas del año 1801, precedida de los principios elementales de la Botánica. Imprenta Real, Madrid. 284p. Available at <https://archive.org/details/descripciondelas1802cava>. Access on 25 January 2023.
  » https://archive.org/details/descripciondelas1802cava
• Chambers TC & Farrant PA (2001) Revision of Blechnum (Blechnaceae) in Malesia. Blumea 46: 283-350. Available at <http://www.repository.naturalis.nl/record/525938>. Access on 24 March 2021.
  » http://www.repository.naturalis.nl/record/525938
• Christensen C (1906) Index filicum, sive enumeratio omnium generum specierumque Filicum et Hydropteridum ab anno 1753 ad finem anni 1905 descriptorum adjectis synonymis principalibus, area geographica etc. H. Hagerup, Copenhagen. 744p. DOI: 10.5962/bhl.title.402
  » https://doi.org/10.5962/bhl.title.402
• Cranfill RB & Kato M (2003) Phylogenetics, biogeography and classification of the woodwardioid ferns (Blechnaceae). In: Chandra S & Srivastava M (eds.) Pteridology in the New Millennium. Kluwer Academic Publishers, Dordrecht. Pp. 25-48. DOI: 10.1007/978-94-017-2811-9_4
  » https://doi.org/10.1007/978-94-017-2811-9_4
• Desvaux AN (1811) Observations sur quelques nouveaux genres de fougères et sur plusieures espéces nouvelles de la même famille. Berlinisches Magazin 5: 297-330.
• Dittrich VAO, Salino A & Almeida TE (2012) Two new species of the fern genus Blechnum with anastomosing veins from northern Brazil. Systematic Botany 37: 38-42. DOI: 10.1600/036364412x616602
  » https://doi.org/10.1600/036364412x616602
• Dittrich VAO, Valente ASM, Fernandez EP, Monteiro NP & Prieto P (2013) Blechnaceae. In: Martinelli G & Moraes MA (eds.) Livro vermelho da flora do Brasil. Andrea Jakobsson Estúdio, Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp. 313-314. <http://dspace.jbrj.gov.br/jspui/bitstream/doc/26/1/LivroVermelho.pdf>.
  » http://dspace.jbrj.gov.br/jspui/bitstream/doc/26/1/LivroVermelho.pdf
• Dittrich VAO, Salino A & Monteiro R (2015) The Blechnum occidentale (Blechnaceae, Polypodiopsida) species group in southern and southeastern Brazil. Phytotaxa 231: 201-229. DOI: 10.11646/phytotaxa.231.3.1
  » https://doi.org/10.11646/phytotaxa.231.3.1
• Dittrich VAO, Salino A, Monteiro R & Gasper AL (2017) The family Blechnaceae (Polypodiopsida) in Brazil: key to the genera and taxonomic treatment of Austroblechnum, Cranfillia, Lomaridium, Neoblechnum and Telmatoblechnum for southern and southeastern Brazil. Phytotaxa 303: 1-33. DOI: 10.11646/phytotaxa.303.1.1
  » https://doi.org/10.11646/phytotaxa.303.1.1
• Dittrich VAO, Salino A, Monteiro R & Gasper AL (2018) The fern genera Lomaria, Lomariocycas, and Parablechnum (Blechnaceae, Polypodiopsida) in southern and southeastern Brazil. Phytotaxa 362: 245-262. DOI: 10.11646/phytotaxa.362.3.1
  » https://doi.org/10.11646/phytotaxa.362.3.1
• Dittrich VAO, Gasper AL & Cárdenas GG (2020) Blechnaceae. In: Flora do Brasil 2020 (continuously updated) Jardim Botânico do Rio de Janeiro. Available at <http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB90784>. Access on 24 March 2021.
  » http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB90784
• Dutra J (1938) A flora pteridófita do estado do Rio Grande do Sul. Anais da Primeira Reunião Sulamericana de Botânica 2: 19-68.
• Gabriel y Galán JM, Prada C, Rolleri C, Ainouche A & Vicent M (2013) cpDNA supports the identification of the major lineages of American Blechnum (Blechnaceae, Polypodiopsida) established by morphology. Turkish Journal of Botany 37: 769-777. DOI: 10.3906/bot-1210-49
  » https://doi.org/10.3906/bot-1210-49
• Gasper AL, Dittrich VAO, Smith AR & Salino A (2016) A classification for Blechnaceae (Polypodiales: Polypodiopsida): new genera, resurrected names, and combinations. Phytotaxa 275: 191-227. DOI: 10.11646/phytotaxa.275.3.1
  » https://doi.org/10.11646/phytotaxa.275.3.1
• Gasper AL, Almeida TE, Dittrich VAO, Smith AR & Salino A (2017) Molecular phylogeny of the fern family Blechnaceae (Polypodiales) with a revised genus-level treatment. Cladistics 33: 429-446. DOI: 10.1111/cla.12173
  » https://doi.org/10.1111/cla.12173
• Hieronymus G (1908) Plantae Stübelianae. Pteridophyta. Dritter Teil. Hedwigia 47: 204-249. <https://www.biodiversitylibrary.org/page/457777>.
  » https://www.biodiversitylibrary.org/page/457777
• Hooker WJ & Baker J (1867) Synopsis Filicum; or, A synopsis of all known ferns, including the Osmundaceae, Schizaeaceae, Marattiaceae, and Ophioglossaceae (chiefly derived from the Kew herbarium). Accompanied by figures representing the essential characters of each genus. R. Hardwicke, London. 482p. Available at <http://www.europeana.eu/portal/pt/record/9200143/BibliographicResource_2000069360640.html>. Access on 25 January 2023.
  » http://www.europeana.eu/portal/pt/record/9200143/BibliographicResource_2000069360640.html
• Humboldt FWHA von, Bonpland AJA & Kunth CS (1815) Nova Genera et Species Plantarum quas in peregrinatione ad plagam æquinoctialem orbis novi collegerunt, descripserunt, partim adumbraverunt Amat. Bonpland et Alex. de Humboldt. Ex schedisautographis Amati Bonplandi in ordinem digessit Carol. Sigismund. Kunth. Accedunt Tabulæ Æri incisæ, et Alexandri de Humboldt notationes ad geographiam plantarum spectantes. Tomus Primus [Vol. 1]. Sumptibus Librariæ Græco-Latino-Germanicæ, Lutetiæ Parisiorum [Paris], [III]-LVIII, 377 pp., tt. 1-96. DOI: 10.5962/bhl.title.640
  » https://doi.org/10.5962/bhl.title.640
• Kaulfuss GF (1824) Enumeratio Filicum quas in itinere circa terram legit cl. Adalbertus de Chamisso adjectis in omnia harum plantarum genera permultasque species non satis cognitas vel novas animadversionibus. C. Cnobloch, Leipzig. 300p.
• Kessler M, Smith AR & Lehnert M (2007) Ten new species and two new combinations of Blechnum (Blechnaceae, Pteridophyta) from Bolivia. American Fern Journal 97: 66-80. DOI: 10.1640/0002-8444(2007)97[66:TNSATN]2.0.CO;2
  » https://doi.org/10.1640/0002-8444(2007)97[66:TNSATN]2.0.CO;2
• Kramer KU, Chambers TC & Hennipman E (1990) Blechnaceae. In: Kramer KU & Green PS (eds.) The families and genera of vascular plants. Vol. 1. Pteridophytes and Gymnosperms. Springer-Verlag, Wien, Berlin. Pp. 60-68. DOI: 10.1007/978-3-662-02604-5_16
  » https://doi.org/10.1007/978-3-662-02604-5_16
• Link JHF (1822) Enumeratio Plantarum Horti Regii Botanici Berolinensis Altera. Pars II. Apud G. Reimer, Berolini. 478p. DOI: 10.5962/bhl.title.66
  » https://doi.org/10.5962/bhl.title.66
• Linnaeus C (1753) Species Plantarum. Vol. 2. Laurentius Salvius, Stockholm. 1200p. DOI: 10.5962/bhl.title.669
  » https://doi.org/10.5962/bhl.title.669
• Mettenius GH (1859) Filices Lechlerianae, chilenses ac peruanae 2. R.F. Hohenacker bei L. Voss, Leipzig. 38p.
• Molino S (2022) A global synopsis and molecular phylogenetic approach of the genus Parablechnum C.Presl (Blechnaceae, Polypodiopsida). PhD Thesis. Universidad Complutense de Madrid, Espanha. 445p.
• Molino S, Gabriel y Galán JM, Sessa EB & Wasowicz P (2019) A multi-character analysis of Struthiopteris leads to the rescue of Spicantopsis (Blechnaceae, Polypodiopsida). Taxon 68: 185-198. DOI: 10.1002/tax.12036
  » https://doi.org/10.1002/tax.12036
• Molino S, Prada C, Gabriel y Galán JM, Wasowicz P, Estébanez B & Vázquez R (2020) Sporangia and spores in the fern genera Spicantopsis and Struthiopteris (Blechnaceae, Polypodiopsida). The Botanical Review 86: 76-92. DOI: 10.1007/s12229-020-09217-z
  » https://doi.org/10.1007/s12229-020-09217-z
• Moran RC (1992) Five new species of ferns from the American tropics. Novon 2: 132-138. DOI: 10.2307/3391674
  » https://doi.org/10.2307/3391674
• Moran RC & Øllgaard B (1995) Six new species of ferns (Polypodiopsida) from Ecuador. Nordic Journal of Botany 15: 177-185. DOI: 10.1111/j.1756-1051.1995.tb00138.x
  » https://doi.org/10.1111/j.1756-1051.1995.tb00138.x
• Moran RC & Smith AR (2005) Blechnum obtusum (Blechnaceae), a new species from western Venezuela. Brittonia 57: 237-239. DOI: 10.1663/0007-196X(2005)057[0237:BOBANS]2.0.CO;2
  » https://doi.org/10.1663/0007-196X(2005)057[0237:BOBANS]2.0.CO;2
• Perrie LR, Wilson RK, Shepherd LD, Ohlsen DJ, Batty EL, Brownsey PJ & Bayly MJ (2014) Molecular phylogenetics and generic taxonomy of Blechnaceae ferns. Taxon 63: 745-758. DOI: 10.12705/634.13
  » https://doi.org/10.12705/634.13
• PPG I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603. DOI: 10.1111/jse.12229
  » https://doi.org/10.1111/jse.12229
• Proctor GR (1977) Pteridophyta. In: Howard RA (ed.) Flora of Lesser Antilles, Leeward and Windward Islands. Vol. 2. Harvard University, Boston. Pp. 1-414.
• Rolleri CH., Prada C, Gabriel y Galán JM, Passarelli LM & Ciciarelli MM (2010) Morphology of the sporophyte and gametophyte of the swamp fern, Blechnum serrulatum (Blechnaceae, Pteridophyta). Australian Journal of Botany 58: 508-518. DOI: 10.1071/BT09238
  » https://doi.org/10.1071/BT09238
• Rolleri CH, Prada C, Passarelli LM, Gabriel y Galán JM & Ciciarelli MM (2012) Revisión de especies monomorfas y subdimorfas del género Blechnum (Blechnaceae-Pteridophyta). Botanica Complutensis 36: 51-77. DOI: 10.5209/rev_BOCM.2012.v36.39444
  » https://doi.org/10.5209/rev_BOCM.2012.v36.39444
• Rothfels CJ, Sundue MA, Kuo LY, Larsson A, Kato M, Schuettpelz E & Pryer KM (2012) A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61: 515-533. DOI: 10.1002/tax.613003
  » https://doi.org/10.1002/tax.613003
• Schuettpelz E & Pryer KM (2007) Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037-1050. DOI: 10.2307/25065903
  » https://doi.org/10.2307/25065903
• Sehnem A (1968) Blecnáceas. In: Reitz R (ed.) Flora Ilustrada Catarinense, BLEC. Herbário Barbosa Rodrigues, Itajaí. Pp. 1-90.
• Shepherd LD, Perrie LR, Parris BS & Brownsey PJ (2007) A molecular phylogeny for the New Zealand Blechnaceae ferns from analyses of chloroplast trnL-trnF DNA sequence. New Zealand Journal of Botany 45: 67-80. DOI: 10.1080/00288250709509703
  » https://doi.org/10.1080/00288250709509703
• Silva DM, Sylvestre LS, Mendonça CBF & Gonçalves-Esteves V (2021) Palynology of selected species of Blechnaceae (Polypodiopsida: Polypodiales). Palynology 45: 507-520. DOI: 10.1080/01916122.2021.1872728
  » https://doi.org/10.1080/01916122.2021.1872728
• Smith AR (1995) Blechnaceae. In: Steyermark J, Berry P, Holst B & Yatskievych K (eds.) Flora of the Venezuelan Guayana. Vol. 2. Pteridophytes, Spermatophytes - Acanthaceae-Araceae. Missouri Botanical Garden Press, St. Louis. Pp. 23-29.
• Smith AR & Kessler M (2018) Prodromus of a fern flora for Bolivia. XXXIII. Blechnaceae. Phytotaxa 334: 99-117. DOI: 10.11646/phytotaxa.334.2.1
  » https://doi.org/10.11646/phytotaxa.334.2.1
• Sota ER de la (1973) Sinopsis de las pteridófitas del Noroeste de Argentina, II. Darwiniana 18: 173-263. Available at <https://www.jstor.org/stable/23215673>.
  » https://www.jstor.org/stable/23215673
• Steyermark J & Maguire B (1984) Nuevos taxa de la Guayana venezolana. Acta Botanica Venezuelica 14: 5-52. Availabre at <https://www.jstor.org/stable/41740439>.
  » https://www.jstor.org/stable/41740439
• Swartz O (1806) Synopsis Filicum. Impensis Bibliopolii Novi Academici, Kiliae. 445p. DOI: 10.5962/bhl.title.81948
  » https://doi.org/10.5962/bhl.title.81948
• Thiers B (continuously updated) Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available at <http://sweetgum.nybg.org/science/ih/>. Access on 25 January 2023.
  » http://sweetgum.nybg.org/science/ih/
• Tryon RM & Stolze RG (1993) Pteridophyta of Peru. Part V. 18. Aspleniaceae - 21. Polypodiaceae. Fieldiana, Botany, new series 32: 1-204.
• Tryon RM & Tryon AF (1982) Ferns and Allied plants: with special reference to Tropical America. Springer-Verlag, New York. 857p.

Edited by

Area Editor: Dra. Thaís Almeida

Publication Dates

• Publication in this collection
  15 Jan 2024
• Date of issue
  2024

History

• Received
  14 July 2023
• Accepted
  17 Oct 2023