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The Struthiopteris spicant (Blechnaceae, Polypodiopsida) complex in Western Europe, with proposals for taxonomic and nomenclatural changes

A Correction to this article was published on 27 April 2019

This article has been updated


Struthiopteris spicant is a medium-sized, dimorphic fern that occurs mainly in Western Europe and Western North America. The species is quite variable, with several forms differing in frond sizes and degree of dimorphism. In addition to the typical plant, in Europe two other varieties are recognized: the Iberian S. s. var. homophyllum includes plants up to 20 cm, monomorphic or subdimorphic, with fragmented cenosori; the Icelandic S. s. var. fallax comprises very small plants up to 5 cm, monomorphic, with isolated sori. Outstanding questions remain about hybridization among the different forms, their taxonomic status, and the relations between different populations. The present work aims to study additional morphological and anatomical features of the S. spicant complex in Western Europe using a statistical approach, to resolve the taxonomic position of the different forms. We observed traits ranging from the rhizome scales to the spores and used one-way ANOVA to test for significant differences between all varieties and PCA to determine whether plants assigned to different varieties can be classified statistically into distinct groups. We detected significant differences in several of the analysed traits among the varieties in the complex. Qualitative differences in pinna anatomy and epidermal cells also help segregation of the varieties considered. Our results allowed us to propose a new specific status for the Icelandic endemic var. fallax and a new endemic Spanish variety, S. s. var. pradae.

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Change history

  • 27 April 2019

    Due to an unfortunate error, the citation of the holotype specimen for the new taxon <Emphasis Type="Italic">Struthiopteris spicant</Emphasis> var. <Emphasis Type="Italic">pradae</Emphasis> is incorrectly stated as MACB 10654, when it should be MACB 110654.


  1. Breitwieser I, Brownsey PJ, Heenan PB, Nelson WA, Wilton AD (eds) (2010–2018) Taxon profile—Blechnum filiforme (based on Allan 1961). Flora of New Zealand Online. Avaliable at: Accessed 1 Aug 2018

  2. Chiou W-L, Shieh W-C, Devol CE (1994) Struthiopteris. In: Huang T-C (ed) Flora of Taiwan, vol I. Editorial committee of the Flora of Taiwan, Taipei, pp 271–273

    Google Scholar 

  3. Christ H (1904) Les fougères de la Galicie Espagnole. Bull Acad Int Géogr Bot 13:76–81

    Google Scholar 

  4. 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. Turk J Bot 37:769–777.

    CAS  Article  Google Scholar 

  5. 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.

    Article  Google Scholar 

  6. 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.

    Article  Google Scholar 

  7. Horjales M, Redondo N, Pérez Prego JM (1990) Nota citotaxonómica sobre pteridoflora del noroeste de la Península Ibérica. Anales Jard Bot Madrid 48:82–84

    Google Scholar 

  8. Hovenkamp PH (1986) A monograph of the fern genus Pyrrosia. Leiden Bot Ser 9:1–280

    Google Scholar 

  9. Iwatsuki K (ed) (1992) Ferns and fern allies of Japan. Heibonsha Publishers, Tokyo

    Google Scholar 

  10. Korpelainen H, Pietiläinen M (2008) Effort to reconstruct past population history in the fern Blechnum spicant. J Pl Res 121:293–298.

    Article  PubMed  Google Scholar 

  11. Kramer K, Chambers T, Hennipman E (1990) Blechnaceae. In: Kramer K, Green P (eds) The families and genera of vascular plants: I. Pteridophytes and Gymnosperms. Springer, Berlin, pp 60–67

    Google Scholar 

  12. Lange J (1880) Icones plantarum Florae Danicae Fasciculus L. C Philibert, Copenhagen

  13. Lawalrée A (1964) Blechnum L. In: Tutin TG, Heywoood VH, Burges NA, Valentine DH, Walters SM, Webb DA (eds) Flora Europaea, vol 1. Lycopodiaceae to Platanaceae. Cambridge University Press, Cambridge, pp 22–23

    Google Scholar 

  14. Löve A, Löve D (1966) The variation of Blechnum spicant. Bot Tidsskr 62:186–196

    Google Scholar 

  15. Löve A, Löve D (1968) Cytotaxonomy of Blechnum spicant. Collect Bot 7:665–676

    Google Scholar 

  16. Lugardon B (1965) Strutcture des parois de la spore de Blechnum spicant (L.) Roth. Pollen Spores 7:409–428

    Google Scholar 

  17. Lugardon B (1974) La structure fine de l’exospore et de la périspore des Filicinées isosporées, II. Filicales. Comment. Pollen Spores 16:161–226

    Google Scholar 

  18. Merino B (1898) Contribución a la Flora de Galicia, Suplemento I. Tipografia Regional, Tuy

  19. Molino S, de la Fuente P, Seral A, Gabriel y Galan JM (2018) Nuevas localidades de tres helechos para la Península Ibérica. Bot Complut 43:2–6

    Google Scholar 

  20. Nakato N (1987) Chromosome numbers of three endemic species of the fern genus Blechnum in Japan. J Jap Bot 62:129–133

    Google Scholar 

  21. Nauman CE (1993) Blechnum. In: FNA Editorial Committee (ed) Flora of North America North of Mexico, vol. 2. Oxford University Press, New York, Oxford, pp 223–227

    Google Scholar 

  22. Ormonde J (1986) Blechnum. In: Castroviejo S, Laínz M, López González G et al (eds) Flora Iberica I, vol. 1. CSIC, Madrid, pp 151–153

    Google Scholar 

  23. Passarelli L (2007) Estudios esporales en especies del grupo Blechnum penna-marina (Blechnaceae-Pteridophyta). Acta Bot Malacit 32:1–19

    Google Scholar 

  24. Passarelli L, Gabriel y Galán JM, Prada C, Rolleri CH (2010) Spore morphology and ornamentation in the genus Blechnum (Blechnaceae, Pteridophyta). Grana 49:243–262.

    Article  Google Scholar 

  25. 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.

    Article  Google Scholar 

  26. PPG1 (2016) A community-derived classification for extant lycophytes and ferns. J Syst Evol 54:563–603.

    Article  Google Scholar 

  27. Prada C, Gabriel y Galán JM, Sáiz P, Passarelli L, Ciciarelli MM, Rolleri C (2016) Caracteres diagnósticos de frondas esporógenas y esporangios de Blechnum (Blechnaceae). Iheringia Bot 71:161–174

    Google Scholar 

  28. Rodríguez-Ríos R (1995) Blechnaceae. In: Marticorena C, Rodríguez R (eds) Flora de Chile, vol. 1. Universidad de Concepción, Concepción, pp 286–303

    Google Scholar 

  29. Rolleri C, Prada C (2006) Revisión de los grupos de especies del género Blechnum (Blechnaceae-Pteridophyta): el grupo B. penna-marina. Acta Bot Malac 31:7–50

    Google Scholar 

  30. Rothfels CJ, Sundue MA, Kuo L-Y, Larsson A, Kato M, Schuettpelz E, Pryer KM (2012) A revised family-level classification for eupolypod II ferns (Polypodiidae: Polypodiales). Taxon 61:515–533

    Article  Google Scholar 

  31. 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 sequences. New Zealand J Bot 45:67–80.

    Article  Google Scholar 

  32. Soltis PS, Soltis DE (1988) Genetic variation and population structure in the fern Blechnum spicant (Blechnaceae) from Western North America. Amer J Bot 75:37–44.

    Article  CAS  Google Scholar 

  33. Tryon AF, Lugardon B (1991) Spores of the pteridophyta. Springer, New York

    Book  Google Scholar 

  34. Tryon RM, Tryon AF (1982) Ferns and allied pants with special reference to tropical America. Springer, New York

    Google Scholar 

  35. Vicent M (2017) Sistemática de la familia Blechnaceae (Polypodiopsida): filogenia de Lomaridium y Parablechnum p.p. y aspectos evolutivos de las pinnas fértiles. PhD Thesis, Universidad Complutense, Madrid

  36. Wagner WHJ, Wagner FS (1977) Fertile–sterile leaf dimorphy in ferns. Gard Bull Singapore 30:251–267

    Google Scholar 

  37. Wasowicz P, Folcik L, Rostanski A (2017a) Typification of Blechnum spicant var. fallax Lange (Blechnaceae). Acta Soc Bot Poloniae 86:3542.

    Article  Google Scholar 

  38. Wasowicz P, Gabriel y Galán JM, Pino R (2017b) New combinations in Struthiopteris spicant for the European flora. Phytotaxa 302:198–200.

    Article  Google Scholar 

  39. Watkins JE, Churchill AC, Holbrook NM (2016) A site for sori: ecophysiology of fertile-sterile leaf dimorphy in ferns. Amer J Bot 103:845–855.

    Article  CAS  PubMed  Google Scholar 

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The Universidad Complutense de Madrid partially supported this research through the funding of the research project PR26/16-20295 and a field trip to Iceland (International Mobility Program 2016).

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Correspondence to Jose M. Gabriel y Galán.

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Appendix 1

Appendix 1

List of materials in alphabetical order of taxa, with indication of location, voucher, and number of individuals per voucher.

Struthiopteris fallax (Lange) S. Molino, Gabriel y Galán & Wasowicz.

ICELAND Deildartunguhver, Wasowicz and Gabriel y Galán7 Jul 2016, (MACB 109359) (3 individuals).

Struthiopteris spicant (L.) Weiss var. spicant

AUSTRIA. Salzburg, Hohe Tauern, 20 Jul 1994, Eberwein and Vitek (MA 767022). BELGIUM. Turnhout, Engels Kamp park2, 18 Jun 1973, W. Van Cotthem (MA 809713). FRANCE. Sources Foret de l’Esterel, 01 Aug 1958, G. Gavelle (MA 186653). Serans (Oise), 22 Jul 1921, P. Bosserdet (P 1001059). ICELAND. Vestfirðir, 8 Jul 2016, Gabriel y Galán and Wasowizc (MACB 110659). IRELAND. Iveragh, Caherdaniel, 17 Jul 2016, Molino and Pachón (MACB 109619). NORWAY. Telemark, Nottoden, 15 Aug 2004, P. Sunding (MA 747921). SPAIN. Asturias: Valdés, Luarca, 12 Apr 2017, Gabriel y Galán (MACB 109615) (7 individuals). Canary Islands: Anaga, Tenerife, 25 Oct 1989, Santos (MACB 36152); Anaga, Tenerife, 14 Sep 2017, Gabriel y Galán (MACB 110658) (2 individuals). Cantabria: Camaleño, Cosgaya, 8 Oct 2016, Gabriel y Galán (MACB 109622) (4 individuals). Madrid: Dehesa de Somosierra, 31 May 2017, Gabriel y Galán and Molino (MACB 109611). Zamora: Aciberos, 23 Sep 2017, Molino et al. (MACB 110655). Pontevedra: Monte Aloya, Tuy, 15 Jul 1993, Pajarón and Pangua (MACB 59142). UNITED KINGDOM. South Somerset, Seven Wells Wood, 30 Jul 1968, J.A. Crabbe (MA 195093). Mid Ebudes, Mull, Tobermory, 06 Jul 1970, Crabbe and Jermy (MA 195091).

Struthiopteris spicantvar. homophyllum (Merino) Gabriel y Galán & R.Pino

PORTUGAL. Braga, Vieira do Minho, 1 Oct 2004, Prada (MACB 109621). SPAIN. La Coruña: Santiago, Cantaleta, 29 Jul 1967, Barrera (MACB 32367). Pontevedra: between Tabagón y Tomiño, 19 Mar 2016, Gabriel y Galán (MACB 109617); Mondariz, 20 Mar 2016, Gabriel y Galán (MACB 109618). Salamanca: Batuecas, 15 May 2016, Gabriel y Galán (MACB 109626) (4 individuals).

Struthiopteris spicantvar. pradae S.Molino & Gabriel y Galán

SPAIN. Asturias: Valdés, Paladeperre, 22 Mar 2016, Gabriel y Galán (MACB 109613) (3 individuals); 3 Aug 2016, íbidem (MACB 109615) (3 individuals); Luarca, 26 Jul 2017, Gabriel y Galán (MACB 110660); Otur, 17 Aug 2017, Gabriel y Galán (MACB 110661). Burgos: Sierra de San Millán, 27 Sep 1975, Fuentes (MACB 5994). Salamanca: San Miguel de Valero, 14 May 2016, Gabriel y Galán (MACB 109624) (3 individuals). Zamora: Aciberos, 23 Sep 2017, Molino et al. (MACB 110654).

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Molino, S., Gabriel y Galán, J.M., Wasowicz, P. et al. The Struthiopteris spicant (Blechnaceae, Polypodiopsida) complex in Western Europe, with proposals for taxonomic and nomenclatural changes. Plant Syst Evol 305, 255–268 (2019).

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  • Endemism
  • Morphology
  • Pteridophytes
  • Taxonomy