Struthiopteris spicant (Hallgrimsson, 1968)

Molino, Sonia, Santos, Guillermo, Vázquez, Rubén, Medina, Rafael & Gabriel, José María, 2024, Monograph of the genera Struthiopteris Scop. and Spicantopsis Nakai (Blechnaceae, Polypodiopsida), Phytotaxa 677 (1), pp. 1-48 : 26-42

Struthiopteris spicant (L.) Weiss ( Figure 9F ; Figure 14E‒F ; Figure 17 ).

Type:— SWEDEN. Scania, no date, no collector ( LINN 1244.14! photo, lectotype, designated in Jonsell & Jarvis 1994).

Synonyms:— ≡ Osmunda spicant L.. Onoclea spicant ( L.) Hoffmann (1796: 11). Acrostichum spicant ( L.) Willdenow (1787: 289). Blechnum spicant ( L.) Roth. Asplenium spicant ( L.) Ehrhart (1788: 95). Lomaria spicant ( L.) Desva.. Lonchitis-aspera spicant ( L.) Farwell (1931: 277).

= Blechnum doodioides Hooker (1840: 263) . Type:—INTERIOR OF N.W. AMERICA. no date, D. Douglas s.n., Herbarium Hookerianum 1867 (K001092778! photo, holotype). Struthiopteris doodioides (Hook.) Trevisan (1869: 571) .

= Blechnum heterophyllum Opiz (1820: 18) . Type:— UNITED KINGDOM. Tunbridge Wells, Kent, 1853, Wollaston s.n. (not found, synonymy based on the current usage of the name).

= Acrostichum lineatum Cavanilles (1799: 106) . Type :— CANADA. Nutka, L. Neé s.n. ( MA228872 !, MA228872-2 ! lectotype, designated here; MA475363 ! isolectotype).

= Acrostichum nemorale Lamarck (1779: 11) . Type:— FRANCE. no date, no collector (not found, synonymy based on the protologue current usage of the name).

= Lomaria crenata Presl (1825: 51) . Type:— CANADA. Habitat in Nutka, no date, no collector (not found, probably at PR, synonymy based on the protologue and current usage of the name); ≡ Blechnum spicant (L.) Roth var. crenatum (C.Presl) Moore (1857a: 205) . Struthiopteris spicant (L.) Weiss var. crenata (C.Presl) Henry (1915: 3) .

= Lomaria spicant (L.) Desv. var. elongata Hooker (1860: 15) . Type:— CANADA. Nutka Sound, no date, Ihcnke s.n.; Juan de Fuca, no date, Brackenridge s.n.; Observatory Inlet, Dr. Scolder; UNITED STATES OF AMERICA. Island of Sitkha, no date, Eschscholtz s.n.; Port Etches, Prince William’s Sound, no date, Hinds s.n. (syntypes, not found, synonymy based on the protologue and current usage of the name). ≡ Blechnum spicant (L.) Roth (1794: 56) var. elongata (Hook.) Boivin (1967: 649) .

= Lomaria spicant (L.) Desv. f bipinnatum Clute (1907: 19) . Type:— CANADA. Vancouver Island, no date, no collector (not found, synonymy based on the protologue and current usage of the name). ≡ Blechnum spicant (L.) Roth f bipinnatum Clute (1911: 72)

= Blechnum spicant (L.) Roth var. serratum Woll in Moore (1857b: 43 C). Type:— UNITED KINGDOM. England, Tunbridge Wells, Kent, no date, Wolaston s.n. (not found, synonymy based on the protologue and current usage of the name) ≡ Lonchitis-aspera spicant (L.) Farwell (1931: 277) var. serrata (Woll ex T.Moore) Farwell (1931: 277) . Blechnum spicant (L.) Roth f. serratum (Woll ex T.Moore) Broun, (1938: 31) . Lomaria spicant (L.) Desv. f. serrata (Woll ex T.Moore) Clute (1907: 19) .

Invalid denominations: Blechnum spicant (L.) Withering (1796: 765) nom. inval.; Blechnum spicant (L.) Smith (1793: 411) nom. inval; Osmunda borealis Salisburi (1796: 402) . nom. sup. Blechnum boreale (Salisb.) Swartz (1801: 75) . nom. sup. Lomaria borealis (Salisb.) Link (1837: 80) . nom. sup.; Blechnum septentrionale Sailer (1841: 306) nom. inval.; Spicanta borealis (Salisb.) Presl (1849: 114) . nom. sup.; Blechnum spicant (L.) Roth f. pectinatum Clute (1911: 73) . nom. nud.

Plants terrestrial; rhizomes thick, short, covered with linear-lanceolate scales, (0.5) 3‒9 x 0.6‒0.8 (3.1) mm, discoloured, dark brown, centre narrowly sclerotized from base to apex, margin entire or toothed, lighter brown; fronds dimorphic, forming rosette; sterile fronds (12) 25 (56) cm, prostrate, stipes short, 0.7‒5 (17)cm, dark brown, black or atropurpureous, laminae narrowly lanceolate to oblong lanceolate, 1.3‒7 x 8‒65 (100) cm, pinnatipartite or pinnatisect, apex pinnatifid, rarely conform, with uniseriate multicellular hairs ending in a red gland, rachis grooved adaxially, green or light brown towards the apex, with glandular hairs like those on the lamina, pinnae tapering towards the base, the middle ones 0.2‒0.5 x 0.8‒6 cm, veins free, simple to 1-furcate, catadromous, ending in adaxial submarginal hydathodes, stomata aligned following veins on abaxial face of pinnae; fertile fronds 36 (75) cm, arising from the central part of the rosette, erect, stipe long, (2) 10‒21 cm, dark brown, black or purplish, red in growth, laminae lanceolate, 2‒4 (8) x 14‒54 cm, pinnatipartite or pinnatisect, apex pinnatifid, rarely conform, with uniseriate multicellular hairs ending in a red gland, rachis grooved adaxially, green or light brown towards apex, with glandular hairs like those on the lamina, pinnae more widely spaced and contracted, the middle ones 0.1‒0.2 x 0.7‒6 cm; sori linear, continuous, forming coenosori on both sides of costa, occupying the entire length of the pinna, decurrent towards the rachis; indusia compound, continuous; sporangia with 12‒18 cells in the arc; spores ovoid, 20‒35 x 30‒50 µm, perispores brown, rugulate, slightly folded.

Habitat and distribution: Europe (from Sweden-Poland-Ukraine-Romania to Iceland), North Africa (where it is rare), the Macaronesian archipelagos (Canary Islands, Madeira and Azores) and the Pacific area of North America (California, Oregon, Washington, Alaska and Canada) ( Figure 2 ). Usually calciferous ( Lawalrée 1964), humid and shady areas ( Ormonde 1986). Usually grows in forests, meadows, meadows, meadows, rocky ravines, and peat bogs ( Rolleri & Prada 2006). From sea level to 3400 m.

Chromosome number:— n = 68, tetraploid ( Manton et al. 1986).

Etymology:— The fertile pinnae give this plant a spiky habit, hence its specific epithet, spicant .

Vernacular names and local uses:—Felecha (“arrow”, Bable), sardina (“sardine”, Bable); blècnum (Catalan); žebrovice různolistá (Czech); kambregne (Dannish); deer fern (English); hard-fern (English); herringbone fern (English); orrazi-iratzea (“comb fern”, Euskera); kampasaniainen (“comb fern”, Finnish); dubbelloof (“double foliage”, Flemish); Fougère-en-épi (“Fern-on-the-cob”, French); blechno (Galician); fenta (“fern”, Galician); rippenfarn (“rib fern”, German); erdei bordapáfrány (Hungarian); skollakambur (“devil’s comb”, Icelandic); lonchite minore ( “minor lonchite”, Italian); varpotoji unksmenė (Lituanian); bjønnkam (“comb fern”, Norwegian); Podrzeń żebrowiec (Polish); feto-pente (“comb fern”, Portuguese); ДербЯнка колосистаЯ (Russian); rebrovka rôznolistá (Slovakian); rebrenjača (Slovenian); lonchite (Spanish); lonquite (Spanish); kambräken (Swedish); Ребрівка Звичайна (Ukranian) ( Ormonde 1986, Salvo 1990, Macía et al. 2018, Niño Ricoi 2019, iNaturalist 2023, Flora of Iceland 2024). According to a specimen collected by Harlod St. John in Washington, U.S.A. (H. St. John 4779, MO1282452), this species is “a favourite of elk”. Quite a few uses have been registered in Spain. For example, in Piloña, Asturias, its occasional use in childbirth has been reported, although it is not specified which part or how it was prepared ( San Miguel 2004). It has also been used as a decorative and ornamental plant ( Verde et al. 2000) and to make floral bouquets together with other ferns (Blanco 1996). Likewise, one of its vernaculars in Bable (Asturian language), “sardina” (sardine), refers to the resemblance of the fronds to a fish scrape, and was used by young girls to play at cooking ( San Miguel 2004). Despite this, it is considered a little-used species with very little known data on its use in past times ( Macía et al. 2018).

Taxonomic notes:— It is common to find individuals with furcate apexes. Indeed, this had led to some authors to describe different forms. Population genetic studies have been conducted both in America ( Soltis & Soltis 1988) and northern Europe ( Korpelainen & Pietiläinen 2008). In both cases it was saw that populations of this species are genetically very similar, which agrees with later studies that have shown it to be a very young species in geological terms, with a divergence age of half a million years ( Molino et al. 2019b). However, to our knowledge, there are no phylogeographic studies that include samples from all the centres of diversity of the species (i.e. America, Europe, Macaronesia and North Africa). We believe that such a study would be very revealing to clarify concisely the evolutionary history of the species. Apparently, the only morphological differences reported between the American and European plants of S. spicant are in size, with the American plants being larger ( Löve & Löve 1966). We have not observed this difference in the specimens studied, however it is possible that there is a collection bias, and that the larger specimens have not often been collected because of the difficulty of being pressed. This is why we think that a field study focused on all distribution centres would also be useful to discern the taxonomic status of the American plants with respect to the rest, since they have even been treated at times as distinct subspecies ( Löve & Löve 1966). It has also been noted by authors such as Löve & Löve (1966) or Hooker (1840) that phenotypic variation within American plants is lower, and in fact much less variation has been described there than in Europe, where up to twentytwo forms and varieties have been described (Aschenson & Graebner 1912). The study of variation within this taxon carried out by Löve & Löve (1966) concluded that within all these forms and varieties, the only ones that had constant and well-defined characters were Blechnum spicant var. fallax (now treated as a distinct species) and Blechnum spicant var. homophyllum . Here we concur with this, adding the variety pradae , which was described many years after this study ( Molino et al. 2019c). To our knowledge, the only variety described in the Pacific populations is Blechnum spicant var. elongata , named by Hooker (1840) and later discarded by himself ( Hooker 1883). A different species was also described from British Columbia, Blechnum doodioides , which has turned out to have no real differences with the species Struthiopteris spicant .

Additional specimens examined:— ANDORRA. La Massana : Arinsal, 5 July 1992, C. Navarro CN939 (MA525715!);Arinsal, riu Coma Pedrosa, 2 October 2004, I. Aizpuru et al. 10698 (MA720621!). AUSTRIA. Kärnten: 14 September 1937, J.V. Schneider s.n. (MA894642!). Niederösterreich: Höllental, July 1859, F. Himmelseher s.n. (C0379957F!); Krumbach, August 1907, Krieger s.n. (MA001476!); ibidem (MA001479!); Seitenstetten, 1907, Strasser s.n. (MO1877370!). Tyrol: no date, J. Traunsteiner s.n. (C0379901F!); 1884, G.C. Jold s.n. (MO1877383!); Alpes du Stubai, 18 August 1986, D. Podlech 12898 (MA464909!; P01567060!); Kirchwald, 4‒24 July 1977, M. Pull s.n. (C0379949F!); Sankt Johan in Tirol, droite d’un sentier montant en direction du Eifersbacerwasserfall et du Angegeralm, August 1989, G. van Buggenhout 13916 (MA497356!; MO4250370!; P01567062!). Salzburg: no date, no collector 178 (MO1877388!); Hohe Tauern, 20 July 1994, R.K. Eberwein & E. Vitek 64‒193 (C0379940F!; MA767022!); Wald Weslhish, 14 September 1937, J. Kutshosh s.n. (P00785357!). Steiermark: in silvis vallis Schottelgrabe prope pagum Oberwölz, 1 August 1909, B. Fest s.n. (MA001417!). Vorarlberg: Bregenz, 6 October 1906, H. Glück s.n. (MO972759!); Gósifer Wald bei Feldkirch, 30 October 1876, H. Klene s.n. (MA775551!). Unknown location: no date, Poiret s.n. (P01549459!); no date, J. Wilde s.n. (P01417526!). BELGIUM. Amberes: bois de hetres et de chênes dénommé “Peerdsbosch”, 1 October 1937, E. Hostié 32 (P01567057!); Turnhout, Engels Kamp park, 18 June 1973, W. van Cotthem 1123 (MA809713!). Vlaams-Brabant: Louvain, 4 October 1903, illegible collector 6454 (P01549394!). Wallonie: Liege, Heppenbach, 9 August 1972, X. Delannay 4 (MO6604759!); Louette- St. Pierre, July 1860, Crepin 1922 [interpreted] (C0379959F!). CANADA. British Columbia: 3 June 1899, Wm. Trelease 2705 & A. Saunders (MO1876819!); Elphinistone Mt., between Sechelt and Gibson, 26 December 1989, W.H. Lewis & M. Elvin-Lewis 14718 (MO6349329!); Kitlope River watershed, 25 July 1991, B. Boyle 457 (MO4270239!); Selkirk, 6 July 1905, C.H. Swaw 835 (MO1876821!); Still water, 15 miles E of Powell River, 9 August 1950, S. Flowers s.n. (MO3318339!); Vancouver, August 1890, H.E. Freeman s.n. (MO1876845!); Vancouver, 29 July 1909, J.M. Macoun 84123 (C0602113F!); Vancouver, Kingston, 14 July 1903, J. Fowler s.n. (MO828538!); Vancouver Island, 1893, J. Macoun 41 (MO1876817!); Vancouver Island, District of Renfrew, June‒July 1901, C.O. Rosendahl & C.J. Brand 89 (MO1876818!); Vancouver island, Vicinity of Crofton, 1905, Mrs. C.E. Young s.n. (MO840177!); Vancouver island, Vicinity of Ucleulet, 29 July 1909, J. Maccun 84123 (MO744189!). CROATIA. Zagreb: Leskovac, 8 August 1875, v. de Borbás s.n. (P01549383!). CZECH REPUBLIC. Liberec: am wege von Spindelmuhle nach Riesengebirge, 24 July 1889, C. Felsmann s.n. (MA001416!); Riesengebirge, Krkonose, 18 August 1888, A. Poislor 1905 (P01549395!). Zlín: Moravia Orient., Valké Karlovice, ad rivulum in valle Milonov in Abieteto, August 1926, G. Rícan 504 (MO2508748!; MO991327!). DENMARK. Faurhult Heat near Frederikshaun, 26 July 1968, L. Jensen et al. (MA213361!); Jylland, 17 July 1889, illegible collector (MO1877387!); Lys naleskov, Djaulekloften, 16 November 1964, Munkegjerg 25 (MO3941341!). FINLAND. Åland: Sund, 1878, A. Arrhenius s.n. (MA001411!). FRANCE. Alsace: Ballon d’Alsace (Chaine des Vosges), 28 July 1959, B. de Retz 44526 (P01567052!). Auvergne-Rhône- Alpes: au pied du Mont Blanc, Haute-Savoie, July 1888, P. Gave s.n. (C0379919F!); Blot, no date, Moufres s.n. (P01567001!); Cantal, bois de Roquasima à Messillac, près Raulhac, 20 September 1883, J. de Puyfol 746 (C0379926F!; P01566983!; P01567042!); Chères, 29 August 1887, V. Auzat s.n. (P01566988!); France de l’Ouest, Vienne, 28 September 1937, J. Simon 31 (P01566970!; P01567036!); Isère, Saint-Pierre-d’Allevard, 22 September 1898, no collector s.n. (C0379922F!); les bois des montées près Chamonix, no date, no collector s.n. (C0379939F!); près Bonneville, Haute-Savoie, 1865, J. Timothée (C0379963F!); route D 20F de Roybon à Montrigaud (France, Isère), aux bords du ruisseau qui traverse le premier chemin de chars au début de la descente de la D 20 F vers Montrigaud, 24 Septembre 1971, R. Barbezat Vidit & A. Lawalrée 5839 (P01567022!); St. Étienne, 1854, illegible collector 3639 (P01567108!). Bretagne: Saint Malo, August 1887 [interpreted], E. Jeanpert s.n. (C0379921F!); Serent, Morbihan, environs de La Salle, 31 July 1942, B. de Retz 18303 (P01567034!). Centre-Val de Loire: Loir e Cher, Romorantin, 18 February 1871, illegible collector s.n. (P01566979!); Voyer (Cher), 18 October 1901, illegible collector s.n. (P01566986!). Champagne-Ardenne: Forêt de Cahmpagne, no date, Devillcers 14 (P01566998!); Villers-Marmery, no date, M. Remy de Livry s.n. (P01566966!). Corse: Corte, bois humides a la Fontaine de Cadamasse, dans la vallée de la Restonica, 22 July 1932, B. de Retz 25bis (P01567051!); Forêt d’Aïtone, 13 June 1885, E. Reverchon 21 (MA001412!; P01549450!); Forêt d’Aitone, Bocca di Vèrghju, Evisa, 18 August 1996, L. Serra & A. Bort 4865 (MA622949!); Forêt de Vizzavona, Basses pentes au Monte d’Oro, entre le Col et la Cascade des Anglais, 20 July 1954, B. de Retz 34457 (P01567043!); Ghisoni, forêt dom le de Marmano, 6 January 1899, R. Rotges 12 (P01566985!). Franche-Comté: Champagney, Mont du Chanoi, 1974, F.-X. Vendrely s.n. (P01566975!); Champagney, 1889, illegible collector s.n. (P01566981!); Ferté, no date, no collector 1405 (P01566993!); NW de Ronchamp, Champagney, Mourière, 1974, F.-X. Vendrely s.n. (P01566994!); Saône, 14 June 1894, R. Marie s.n. (P01566999!); Vielle-Hutte, 1974, F.-X. Vendrely s.n. (P01566995!). Grand Est: Chaine des Vosges, bords tourbeux du Lac de Lispach, 5 August 1934, B. Retz 9013 (P01567027!); Flornoy (Haute Marne), Regny-bois, 21 August 1904, C. Thomas s.n. (P01567047!); ibidem, 1 September 1932, B. de Retz 2865 (P01567038!); ibidem, 8 August 1961, B. de Retz 48218 (P01567046!); ibidem, B. de Retz 48220 (P01567039!); Le Mussey, 27 June 1874, illegible collector s.n. (P01566982!); Le Rothenbachkopf, Chaine des Vosges, 11 August 1935, B. de Retz 10489 (P01567026!); Moselle, 12 August 1934, M. de Wailly 4304/4313 (P01567041!); Remiremont (Vosges), September 1844, A. Tocquaine 1912 (P01549426!); ibidem, A. Tocquaine194 (C0379906F!; P01566980!);Vosges,1811, J.B.Mougeot Bruyeriensis M.D.&C.Nestler Argentinensis Pharm 110 (C0379950F!; MO1611160!); ibidem, 2 August 1933, Bois s.n. (P01566977!); Vosges, 6 October 1937, B. de Retz 30 (P01567040!); Vosges, massif du faud Ballon, près du rñeservoir de la Lauch en moutaut vers le Marmsteiu, 6 October 1937, B. de Ritz 30 (P01566974!). Haust-des-France: Beauvais (Oise), lande ombragée en lisière du Bois de Belloy, 26 June 1932, B. de Retz 25ter (P01567037!); Foret de Hallotte, Mont Pagnotte, 25 October 1961, H.E. Jeanpert s.n. (C0379913F!); Nord, pentes et vallée sud-ouest du Mont-Noir, près de Bailleul, arr. D’Hazebrouck, 9 June 1853, S.V. Vercier 223 (C0379964F!; P01567000!). Île-de-France: Domont, 19 August 1891, E. Jeanpert s.n. (C0379923F!); Forêt de Marly, Carrefour des Essants, 11 September 1910, H.E. Jeanpert s.n. (C0379936F!); Forêt de Marly, partie de la forêt au S. de la gare de S Nom la Breteche, 22 May 1949, H. Laurence s.n. (P01566978!); Forêt de Montmorency, prés St. Leu, 9 January 1898, H.E. Jeanpert s.n. (C0379937F!); Forêt de Rambouillet, Seine et Oise, bois humides au sud de l’étang d’Angennes, 6 Septembre 1942, B. de Retz 18473 (P01567035!); Vallen de la Guesle, Poigny, 2 July 1939, G. Dillemann s.n. (P01566990!). Limousin: Cantal, le long du GR400 entre Aubusson (au-dessus de St Julien de Jordanne) et le Puy Bassiérou, près d’un ruisseau, 11 June 2009, G. Rouhan & M. Gaudeul 976 (P00577715!). Loire: Forêt de Bercé, 1899, V. Jamin s.n. (P01660190!); ibidem, November 1900, V. Jamin s.n. (P01658100!); ibidem (P01658103!); ibidem (P01660191!): Monte Pierre sur Haute, 14 August 1815, M. Gandoger s.n. (MA001419!); ibidem, July 1878, M. Gandoger s.n. (MO1877367!); Mt. Pilat, July 1843, Mutel 185 (MA135041!). Lorena: Bitche (Moselle), 27 July 1850, F. Schultz 1903 (P01549414!). Midi-Pyrénées: Cauterets (Hautes-Pyrenees), Marcadau (Refuge Wallon), 19 August 1928, B. de Retz 25 (P01567025!); Cauterets (Hautes-Pyrenees), sous le Lac de Gaube, 12 September 1959, B. de Retz 44703 (P01567045!). Nord-Pas-de-Calais: pentes et vallée sud-ouest du Mont-Noir, près de Bailleul, arr. D’Hazebrouck, 9 June 1853, S.V. Vercier 223 (P01567109!). Normandie: Chanteloup, 3 August 1889, no collector s.n. (P01566991!); entre Trouville et Honfleur, September 1875, no collector s.n. (P01566969!); Forêt de Cerisy, 1873, no collector s.n. (P01658102!); ibidem, 26 August 1873, no collector s.n. (P01658101!); Forêt d’Eu, Poreau de la Madeleine, 9 August 1924, M. de Wailly 4313 (P01567048!); Forêt de la Londe (près d’Elbeuf), 13 August 1972, no collector s.n. (P01359500!); Manche, à Saint-Hilaire-du-Harcouët, 11 July and 28 August 1854, Brehier s.n. (P01566987!); ibidem, Brehier 99 (P01566989!); Marais Vernier, September 1945, no collector s.n. (P00917150!); Orne, Saint Bénit, sur le Calcaire, September 1879, E. Reverchon s.n. (C0379918F!). Nouvelle-Aquitaine: Basses Pyrenées, 6 September 1973, no collector s.n. (P01567031!); Basses Pyrenees, dans le Vallon de Laxia, 21 July 1941, B. de Retz 17222 (P01567033!); Biarritz, 19 July 1859, Illegible collector (MO1877389!); Biarritz, 1946, G. Sag 40 (P00041961!); Bois de Guéret, no date, no collector s.n. (P01567028!); Corrèze, Forêt de Gimel, 1854, S. de Daloz s.n. [interpreted] (P01566997!); Deux-Sèvres, Forêt de Secondigny, 28 July 1940, L. Rallet 375 (P01566971!; P01567032!; P01567050!); Forêt de la Braconne, prés la Belle Etoile, 24 July 1921, no collector s.n. (P01617210!); France de l’Ouest, Vendée, St. Hilaire du Bois, 13 June 1937, J. Charrier205 (P01566972!; P01567049!); Gironde, Baigneaux, bois au Sud de Baigneaux, entre Baigneaux et Laurenceae, au N de la route de Martres à Cantois, 4 August 1967 – 19 July 1968, E. Contré & V.J. Callé 5306 (P01567024!); Vendée, St. Cyr des Gâts, 4 Jume 1926, J. Charrier 4510 (P01566973!). Occitanie: Alto Garona, Bagneres de Luchon, Foret de Herran, 7 July 1978, J.M. Montserrat s.n. (MA808488!); Aveyron, Cantoin, 17 July 1919, Caibonal s.n. (MO2357041!); Aveyron, Aubzac (Revel), 1845–1852, T. Puel 1454 (P01566996!); Lozère, sommet de la Margeride, 20 September 1941, E. Licent s.n. (P00708108!); Saleix, Ariège, 14 September 1981, L. Villar s.n. (MA808487!). Pays de la Picardie: Forêt de Villers-Cotterêts, 1846, no collector s.n. (P01567004!); Forêt de Villers-Cotterêts, 6 July 1894, H.E. Jeanpert s.n. (C0379914F!); ibidem (C0379917F!); ibidem (C0379920F!); Serans (Oise), 22 July 1921, P. Bosserdet s.n. (P01001059!). Pays de la Loire: Forêt de Blois, 20 October 1861, S. Mathennet 1921 (C0379968F!); Forêt de Perseigne, 8 August 1879, no collector s.n. (P01567003!); La Ferrière, 1879, J. Hommey s.n. (P01566984!); Marais de Soesmes, August 1879, A.R. Franchet s.n. (C0379930F!); Saône et Loire, La Boulage, no date, F. Chassignol s.n. (P01567023!). Poitou-Charentes: Essards, 9 July 1896, E. Jeanpert s.n. (C0379934F!). Provence-Alpes-Côte d’Azur: Sources Foret de l’Esterel, 01 August 1958, G. Gavelle s.n. (MA186653!). Rhône-Alpes: Route de Roybon a Montrigaud, aux bourds du ruisseau quitraverse le premier che de chars au debut de descente de la D 20F vers Montrigaud, 24 September 1971, R. Barbezat s.n. (MA267596!). Unknown location: no date, no collector s.n. (P01566962!); no date, illegible collector s.n. (P01567030!); 20 June 1869, no collector s.n. (P01566963!); July 1871, Illegible collector (MO1877381!); 1 November 1882, H.F. Munroe s.n. (C0379938F!); 26 July 1896, H.E. Jeanpert s.n. (C0379935F!); October 1901, F.G: Walt 1911 (MO92053!; P01549400!); 11 July 1934, M. Martínez s.n. (MA001422!); 1902, C. Thomas s.n. (P01567044!); illegible location, 1847, illegible collector s.n. (P01566967!); illegible location, June 1870, illegible collector s.n. (P01566992!); illegible location, Hirantz s.n. [interpreted], 1872 (P01566968!); Bois de la couche prés autun [interpreted], 13 June 1842, no collector s.n. (P01567029!); Novau-Vallée de la Couche [interpreted], 7 July 1898, illegible collector 414 (P01567002!). GERMANY. Baden-Wurtemberg: 1906, J. Müller-Knatz 16727 (P01549428!); October 1906, W. Wagner s.n. (P01549420!); 1926, A. Lösch s.n. (C0379952F!); Baden-Baden, 5 October 1899, A. Kunz 6357 (P01417532!); Herrenau, 16 August 1892, Hurritgen & A. Kunz 1902 [interpreted] (P01549413!); Schwarzwald, 5 August 1899, A. Kunz 1892 (P01549448!). Bayern: 8 November 1847, A.M. Einsele s.n. (C0379942F!); August 1891, P. Lydon s.n. (C0379947F!); 14 September 1899, illegible collector 1891 (P01549438!); August 1900, J. Kaulfufs s.n. (P01549439!); 18 August 1912, illegible collector (C0379928F!); 14 October 1916, illegible collector 5165 (C0379944F!); auf Waldwegen und an Graben am Hulberg bei Fischbach nächt Nürnberg, 6 October 1987, J. Kaulpup s.n. (MA001472!); Bodenmais, 11 September 1916, L. Oberneder 5061 (C0379945F!); Elsass, Weissenburg, 24August 1895, Spindler s.n. (P01549421!); Erlangen, no date, illegible collector, (C0379895F!); Feuchte Walber und Nürinberg, 8 aug 1903, J. Kaulfup s.n. (MA001473!); Feuchte Waldgräben und Ränder eines Waldbaches beim Hambacher Sauerbrunnen zu Birkenfeld, September 1904, J. Müller-Knatz & F. Wirtgen 462 (P01571950!); Miescach, 1920, A. Lösch s.n. (C0379953F!); Oberbayern, Berchtesgadener Alpen, Watzmannstock, zwischen Herrenroit und Kühroint, Fichtenwald, 10 October 1963, W. Lippert s.n. (MA186638!); Oberbayern, Waldrander sudlich von Grabenstatt am Chiemses, 1 August 1902, H. Stadler 772 (MO1704051!). Berlin: Melzergrund, 26 July 1901, F. Woller s.n. (MA001423!). Hamburg: 29 July 1877, illegible collector s.n. (C0379971F!). Hesse: Hesse-Nassau, 11 October 1896, Illegible collector 1900 (P01549412!; P01549417!; P01549418!); ibidem, August–September 1899, J. Müller-Knatz s.n. (P01417513!); Hasse-Nassau, Taurus, Haidtränkthal bei Oberursel, 29 August 1894, F. Wirtgen 6218 (P01417497!; P01417498!; P01417499!; P01417500!; P01417501!; P01417502!); ibidem, 29 August 1897, F. Wirtgen 6359 (P01417530!); ibidem, 12 September 1897, J. Müller-Knatz 6260 (P01465210!); ibidem, 11 October 1897, J. Müller-Knatz 69 (P01417515!); ibidem, 11 September 1897, J. Müller-Knatz s.n. (P01417487!); ibidem (P01417488!); ibidem, 1899, J. Müller-Knatz s.n. (P01417473!); ibidem (P01417474!); inidem (P01417482!); ibidem (P01417517!); ibidem, J. Müller-Knatz 232 (P01417475!); ibidem, September 1900, J. Müller-Knatz s.n. (P01417511!); ibidem, J. Müller-Knatz 6224 (P01417545!); ibidem, 28 October 1901, J. Müller-Knatz s.n. (MA001478!); ibidem, 18 October 1903, J. Müller-Knatz s.n. (MA001475!); ibidem, 7 October 1906, J. Müller-Knatz s.n. (MA001480!); ibidem, September 1908, M. Durieu [interpreted] 9425 (P01417490!); Frankfurt, 1826, no collector s.n. (MO1877378!); Frankfurt, July 1826, G. Engelmann s.n. (MO1877376!). Niedersachsen: Hameln, no date, Pflümer s.n. (MO872983!); Hannover, 1889, Meyerholz 21029 (P01417476!); ibidem, 1891, C. Luerssen s.n. (P01417520!); Oldenburg, July 1899, illegible collector 6219 (P01417503!); ibidem, 9 September 1904, illegible collector 9428 (P01465207!); ibidem, 10 September 1904, illegible collector 9416 (P0141756!); illegible collector 9423 (P01417507!); ibidem, illegible collector 9433 (P01417529!); ibidem, illegible collector 9434 (P01417540!); Oldenburg, Varel, 21 January 1899, D. Fr. Miller s.n. (P01417514!); ibidem, July 1899, D. Fr. Miller 6356 (P01417533!; P01417534!); ibidem, D. Fr. Miller 6360 (P01417538!); Piesberg,near Osnabrück, 6 June 1857, no collector s.n. (MO1877372!). North Rhine-Westphalia: 9 September 1891, illegible collector 1883 (P01549435!); 3 October 1891, illegible collector 1884 (P01549429!); 27 October 1891, illegible collector 1885 (P01549430!); 21 September 1892, illegible collector 1882 (P01549434!); 7 August 1893, illegible collector1881 (P01549433!); 26 June 1899, illegible collector1886 (P01549432!); 13 September 1899, Illegible collector 1887 (P01549431!); September 1899, F. Writgen 6221 (P01417505!); ibidem, F. Writgen 6361 (P01417539!); 8 October 1899, F. Writgen 6358 (P01417531!); 30 September 1898, W. Sinemkam 1895 [interpreted] (P01549447!); September 1899, F. Wirtgen 1890 (P01549437!); 27 September 1900, F. Wirtgen 1946 (P01417495!); August 1904, F. Wirtgen 9424 (P01417496!); Bergisch, 27 October 1897, illegible collector 1885b (P01549346!); ibidem, 15 September 1898, F. Wirtgen 6355 (P01417536!); ibidem, 28 September 1898, F. Wirtgen 6216 (P01417493!); ibidem, 3 October 1898, F. Wirtgen 6240 (P01417479!); ibidem, 6 October 1900, F. Wirtgen 1947 (P01417522!); ibidem, F.Wirtgen 6239 (P01417481!); ibidem, 2 September 1903, F.Wirtgen 9324 (P01417491!); ibidem, 23 September 1903, F. Wirtgen 9373 (P01417492!); Bielefeld, 1859, H. Wagner s.n. (C0379898F!); Coesfeld, 22 October 1899, F. Wirtgen 6220 (P01417504!); Lieggebiet, Bröl, 13 September 1899, F. Wirtgen s.n. (P01417537!); ibidem, F. Wirtgen 1948 (P014174914); Rheinsprovinz, 20 August 1900, C. Luerssen 5548 (P01549381!); ibidem C. Luerssen 5551 (P01549385!); ibidem, C. Luerssen 5553 (P01549382!; P01417525!); ibidem C. Luerssen 5557 (P01549384!). Rheinland-Pfalz: July 1982, BRD s.n. (MO5496043!); 1860, Haussbueiht s.n. [interpreted] (P01417521!); 1897– 1899, F. Wirtgen 69 (P01417519!); 22 August 1899, C. Luerssen 5332 (P01417512!); 20 August 1900, C. Luerssen 5548 (P01417523!; P01417524!); 28 August 1900, C. Luerssen 5575 (P01417510!); Nahe, October 1898, L. Geisenheyner s.n. (P01417478!). Sachsen: August 1904, K.W. Krieger s.n. (C0379956F!); August 1911, E.R. Missbach s.n. (C0379954F!); Fichtelberg, 6 August 1900, C. Haussknecht 1889 (P01549440!); Prope Königstein, September 1908, W. Krieger s.n. (C0379946F!); Sächsische schweiz, September 1934, no collector s.n. (P00847085!); Sandsteingebirge bei Schöna, August 1906, R. Missbach s.n. (MA001481!); Waldrand Str. Schönanbilhl Baldenchofen, 11 September 1985, R. Bussmann RBM309 (MO6133694!). Sachsen-Anhalt: Dresden Dist., “Sächsische Schweis”, 31 August 1873, C. Luerssen 5479 (C0379933F!); Schellbach, 10 September 1910, Müller-Knatz (P01417508!). Schleswig-Holstein: September 1898, J. Lenmier 63 (P01417477!); Kreis Pinneberg, 15 October 1899, J. Schmidt 32 (P01417518!); ibidem, September 1903, J. Schmidt 9375 (P01417506!); Kreis Pinneberg, an Gräben und Knicks um Tangstedt häufig, September 1902 –1903, J. Schmidt 9 (P01549419!). Thüringen: Varel, 21 January 1899, D. Fr. Miller s.n. (P01417516!); 31 July 1901, B. Hergt 6217 (P01417509!); Schmücke [interpreted], 30 June 1890, Guriegun 1899 [interpreted] (P01549415!; P01549416!). Unknown location: no date, illegible collector (MO1877377!); no date, Becukach s.n. [interpreted] (MO1877380!); no date, ex. Herb. A.N. Desvaux (P01549436!); no date, A. Daenen s.n. (C0379964F!); no date, G. Engelmann s.n. (C0379894F!); ibidem (C0379900F!); no date, F. J. Jung 31 [interpreted] (C0379893F!); no date, I.A. Lapham 43 (C0379967F!); no date, W. Gmelin s.n. (C0379969F!); 10 September 1819, no collector s.n. (P01567061!); August 1849, A.C.V. Schott s.n. (C0379970F!); July 1858, E. Stizenberger s.n. (C0379958F!); 3 October 1860, L. Gommerfeld 17 (MO909691!); 17 August 1864, L.J. Wahlstedt s.n. (C0379903F!); 1866, Fr. Körnicke 1896 (P01549442!); 1865, H. Eggert s.n. (MO1877373!); 1869, H. Eggert s.n. (C0379896F!); 26 August 1877, J. Barth 8298 (P01549392!); 1898, L. Geisenheyner 20912 (P01549386!); 1898, L. Geisenheyner 20916 (P01549387!); 20 August 1900, C. Luerssen 5553 (P01417485!); 24 August 1900, C. Luerssen 5561 (C0379943F!; P01417486!); 1902, F. Writgen s.n. (P01417484!); 28 August 1903, no collector s.n. (P01417483!); August 1907, W. Peschlow 83 (C0379951F!); 10 September 1910, J. Müller-Knatz 12813 (P01465206!); ibidem, J. Müller-Knatz 12814 (P01465205!); 26 July 1919, Schcuattarch s.n. [interpreted] (C0379955F!). GREECE. Creta: Distr. Kydonia, Inter Skines et Nea Rumata, 24 July 1973, K.H. Rechinger 45770 (MO6583303!). Tesalia: Mont Pelión, 1906, R. Marie & M. Petitmengin s.n. (P01567006!). ICELAND. Vestfirðir, 8 July 2016, J.M. Gabriel y Galán & P. Wasowicz s.n. (MACB110659!). IRAN. Juzestan: Ahvaz, bajo Karum, October 1899, F.M. de la Escalera s.n. (MA001420!). IRELAND. Cork: Doneraile, 1963, Mrs J. Bodger s.n. (MO5462759!). Kerry: Iveragh, Caherdaniel, 17 Ju 2016, S. Molino & M. Pachón s.n. (MACB109619!). ITALY. Abruzzo: Teramo, Ceppo, Monte della Laga, bosque della Morricana, 3 August 2002, A. Herrero 1958 (MA698559!). Calabria: Giaja, 1871‒1872, M. Gandoger 1462 (MO1877369!). Liguria: San Romolo, San Remo, 12 November 1900, C. Bicknell 1910 (P01549402!). Piamonte: Stresa, Lago Maggiore, 25 July 1900, L. Pollini & C. Bicknell 1909 (MA001477!; P01549378!; P01549397!); ibidem, 4 September 1906, B. Boigtiaender-Gebner s.n. (P01549401!). Sassari: Mte de Limbara (Sard. September), 1866, Marcucci s.n. (P01549454!). Sicily: Castelbuono, no date, Todaro 1391 (P01549406!); Castelbuono, Mt. Colli, 6 April 1874, Heidenreich 21122 (P01549376!). Toscana: Monte Pisano, July 1862, no collector s.n. (C0379966F!). Trentino- Alto Adigio: Alpi Apuane, versante nord-orientale del M.Altissimo, September 1946, A. Frichi 12120 (MO1610874!). MOROCCO. Unknown location: in monte Yebel Buhaschen (Yebala), 20 June 1928, P. Font Quer 442 (MA001410!); ibidem, P. Font Quer s.n. (MA001421!). NETHERLANDS. North Brabant: Moergestel, near Putven, 24 July 1920, M.R.S. Boetje-van Ruyven s.n. (C0379907F!). Overijssel: Denekamp, 11 August 1916, M.R.S. Boetje-van Ruyven s.n. (C0379932F!). Utrecht: 28 September 1951, K.U. Kramer s.n. (MO1635832!). NORWAY. Hordaland: Dalvik, near Bergen, August 1946, P. Humphreys s.n. (C0379902F!); Norland: Grane, 16August 1927, P. Benum s.n. (P01567058!). Svalbard: Spitzberg, 1838, M. Martins s.n. (P01549411!). Vestflod og Telemark: Notodden, 15 August 2004, P. Sunding s.n. (MA747921!). Unknown location: no date, A. Blytt 599 (MO1877384!); no date, C. Rosenberg s.n. (MA001413!); illegible location, 19 July 1890, A. Holms s.n. (P01539374!). POLAND. Bielsko: Brodu ad Kaluaria, 1 November 1925, J. Trela s.n. (MO1858732!); Carpati Occidentales, Brody, Distr. Wadowice, 15 September 1938, J. Trela 306 (MO1659338!); Carpati occidentales, Jaworze, distr. Nowy Sacz, no date, J. Staszkiewicz & K. Szcepanek 501 (MA267595!; MO1770493!). Liberec: Riesengebirge, 17 August 1878, C. Luerssen 8147 (C0379897F!); Lower Silesian: no date, no collector s.n. (MO872984!). Małopolska: West Carpathian Mts., Beskid Mazy, Tarnawa Górna near Wadowice, on the foot of Mt. Leskowiec, 21 September 1980, J. Kornas &A. Medwecka-Kornas60 (MO2888432!). Silesian: Distr. Poreba, 13 August 1956, N. Ralska s.n. (MO1711825!); West Carpathian Mts., Beskid Slaski, Stozek near Wisla, 24 September 1975, G. Halastra & E. Regdos 27 (MO3868056!). PORTUGAL. Alentejo: Alto-Alentejo, Serra de S. Mamede, Barreirato , 26 April 1957, J.V.C. Malato-Beliz et al. 4082 (MA270372!); Alto-Alentejo, Serra de S. Mamede , prox de Alegrete, ribeira de Arromches, Porto de Azenha, 3 April 1957, J.V.C. Malato-Beliz et al. 4193 (MA269488!); Alto-Alentejo, Serra de S. Mamede, Serra da Selada , junto a estrada dos Galegos, May 1957, J.V.C. Malato-Beliz et al. 4233 (MA269487!); Alto-Alentejo, Serra de S. Mamede , Portalegre, entre S. Julián e a fronteira, margens da ribeira de Xévora, 3 May 1969, J.V.C. Malato-Beliz et al. 7207 (MA269483!); Alto-Alentejo, Serra de S. Mamede , Portalegre, S. Julián, Monte Sete, margens do ribeiro da Lamaceira, 4 June 1969, J.V.C. Malato-Beliz & J.A Guerra 7233 (MA269482!); Alto-Alentejo, Serra de S. Mamede , estrada de Portalegre para S Mamede, prox. Para o cruzamento para o ponto mais alto, 2 July 1969, J.A. Guerra & J.V.C. Malato-Beliz 7763 (MA269481!). Algarve: Serra de Manelique , ribeira de Piroes, 22 April 1968, J.V.C. Malato-Beliz 5959 (MA269485!); Serra de Manelique , estrada para Alferce (ca. 15 km de Monchique), 22 April 1968, J.V.C. Malato-Beliz 5985 (MA269484!); Serra de Monchique , estrada para a Foia, prox do miradouro, Bicas, 28 June 1978, J.A. Guerra & J.V.C. Malato-Beliz 14907 (MA269479!); Serra de Monchique, Foia, J.V.C. Malato-Beliz & J.A. Guerra 14974 (MA269478!); Serra de Monchique , estrada Monchique-Saboia, barranco de Maceira, 20 May 1979, J.A.Guerra & J.V.C.Malato-Beliz 15941 (MA269477!); Serra de Monchique , barranco de Maceira, afluente de Riba de Perna Negra, 30 May 1979, J.V.C. Malato-Beliz & J.A. Guerra 16040 (MA269476!); Serra de Monchique , estrada Monchique-Saboia, barranco de Mamira, prox de Cimalhas, 31 May 1979, J.A. Guerra & J.V.C. Malato-Beliz 16058 (MA269475!). Azores: Isal, 19 July 1894, C.S. Brown 327 (US262512!); Santa Cruz, 29 September 1971, T.L. Monor 15084 [interpeted] (P01567064!); S. Miguel Island, Furnas, Terra Nostra Park, 17 March 2007, J.A. Ramos & L.G. Quintanilla LGQ765 (MACB113885!); Terceira, 27 June 1894, W. Trelease 1166 (US816235!). Aveiro: Vale do Vouga, Ribeiro da Calada, August 1943, P.M. Póvoa dos Reis 908 (MO1637038!). Centro: Beira Alta, Serra da Estrela, 1952 , J.V.C. Malato-Beliz s.n. (MA270374!); Beira Alta, Viseu, Santa Comba-Dao, Chamadouro, ponte de Ouveiro, junto da Barragem, 11 June 1982, A. Marques 2065 (MA385041!); Beira Litoral, Coimbra, Arganil, Benfeita, Pardieiros, mata de Margaraga, 24 July 1982, A. Marques s.n. (MA390255!); entre Oronhe e Travasso, a 2, 5 km de Travasso, na estrada Agueda-Aveiro, 2April 1965, A. Fernández s.n. (MA195094!); Viseu, Vouzela, Serra do Caramulo, pr. Carvalhal de Vermilhas, 24 March 2014, J. Calvo 6394 (MA895113!). Lisboa: Serra do Sintra, February 1840, no collector 303 (P01549452!). Madeira Island: 1838 to 1842, Wilkes Explor. Exped. s.n. (US92905!); entre S. Vicente e a Encumeada, acima do chao de Louros, 8 June 1957, J.V.C. Malato Beliz 1531 (MA266380!);ER228road between Encumeada and S.Vicente, 2May 1998, L.G. Quintanilla LGQ685 (MACB113887!; MACB113888!; MACB1138889!; MACB113890!); Funchal, Chemin entre Choupana et Quinta de Vale de Paraiso, 23 July 2007, G. Rouhan & M. Gaudeul 786 (P00577732!); Levada do Furado, estre o Ribeiro frio e o Santo do Serra, 14 June 1957, J.V.C. Malato-Beliz 1711 (MA266378!); Levada de Casa do Lombo do Mouro, 28 June 2000, C. Navarro 3377 (MA654709!); Montado dos Persegueiros, junto ao Ribeiro da Partilha, 13 June 1957, J.V.C. Malato-Beliz 1647 (MA266379; US229039!!); Paul de Serra, próx. Do Pico da Urze, 10 July 1954, J.V.C. Malato-Beliz 843 (MA266381!); subida de Queimadas para o pico Ruivo, 3 July 1954, J.V.C. Malato-Beliz 761 (MA266382!). Norte: entre Braga e Porto, nas cercanias do Senhor do Perdao, 28 April 1955, J.V.C. Malato-Beliz 2256 (MA270373!); Fafe, Lameira, Mouta das Regadas, Carvalhal da Cerdeira, 24 Jjul 1977, J.V.C. Malato-Beliz & J.A. Guerra 13746 (MA269480!); Fervença-Mondim de Basto, 16 July 1997, D. Soutinho 740 (MA643519!); Mesao Frio, prox ao rio Teixeira, 22 March 1941, P. Lopes & G. Pedro s.n. (MA414816!); Minho, Arcos de Vale de Veg Grosevila, num caminho que liga com a estrada para Mourilhoes e a estrada para Gonte da Barca, 13 July 1944, C. Fontes & M. Myre 9754 (MA001418!); Minho, Ponte de Lima, lieux ombragés humides près de Rebordâes, 4 September 1962, B.V. Rainha & V.J. Callé 5812 (P01567063!); Minho, Viana do Castelo, Ponte de Lima, Arcos, P.N. de las Lagoas de Bertiandos, 7 June 2010, J.L. Fernández-Alonso et al. JFA29196 (MO6423037!); PNA1-Muas-Vila Real, à entrada do PNAL, margem direita da estrada para Lamas de Olo, 6 July 1997, D. Soutinho 616 (MA643615!); Porto, arredores, Ermezinde numa bouça, 18 June 1941, J. Castro s.n. (MA186642!); Serra de Gerês entre baldas e a Pedra Bela, 10 July 1958, J.V.C. Malato-Beliz 4416 (MA186640!; MA269486!); Viana do Castelo, Caminha, Serra do Arga, Arga de São João, Alto do Espiñeiro, 7 June 2010, J.L. Fernández Alonso 29170 (MA821590!); Viana do Castelo, Ponte de Lima, Arcos, P.N. de laos Lagos de Bertiandos, 7 June 2010, J.L. Fernández Alonso 29196 (MA821614!); Vila Nova de Cerveira, Gondarém, entre el Estalagem Boega y el Calvario, 5 June 2010, A. Quintanar Sánchez 3717 (MA824803!). Ribatejo: Santarém, Villa Nova d’Ourem, September 1883, M. Daveau 1033 (P01549396!; P01549410!; P01549449!). Unknown location: no date, Neé s.n. (MA233979!); debajo de la venta de Cody, bajando a Laguna, A.J. Cavanilles s.n. (MA232868!). RUMANIA. Transilvania: 16 August 1877, J. Barth 8298 (P01549388!); Distr. FăgăraȘ, 17 July 1930, E.I. Nyárády 1142 (MO110160!). SPAIN. Andalucía: no date, no collector (MA226680!); Cádiz, Algeciras, 2 July 1979, E. Salvo s.n. (MA235635!); Cadiz, Grazalema, 26 May 1890, E. Reverchon s.n. (MA001415!); Cádiz, Jeréz, June 1908, M. Gandoger s.n. (MO1877368!); Cádiz, Jerez, Sierra de Aljibe, 6 November 1880, Pérez Lara s.n. (MA001429!); Huelva, Cortegana, 10 June 1931, E. Gros s.n. (MA001434!); ibidem, 28 May 1976, B. Cabezudo Artero 1207/76 (MA208547!); Málaga, Sierra de Yunquera, 8 June 1890, E. Reverchon 1913 (P01549399!); ibidem, E. Reverchon 479 (C0379914F!; P01549453!). Aragón: Huesca, Gistaín, Puerto de la Paz, 15 May 1969, P. Montserrat Recorder s.n. (MA808472!); Huesca, Benasque, cerca del hospital, 7 August 1986, L. Villar & P. Montserrat s.n. (MA382740!); Huesca, Panticeya, August 1918, L. Atercito s.n. (MA135039!); Huesca, Sallent de Gállego, barranco de Espelunciecha, 20 July 1981, L. Villar s.n. (MA8008489!); Huesca, Valle de Benasque, July 1873, Frio Trémols s.n. (MA001436!); Huesca, Valle de Tena, July 1881, Zubilla s.n. (MA001428!). Asturias: no date, no collector s.n. (MA159349!); Ajuyán, 15 January 1994, T.E. Díaz González s.n. (MA540468!); Arrás, no date, Lagasca s.n. (MA232663!); Bendición, 1 July 1915, C. Martínez s.n. (MA001449!); Cangas de Narcea, de Rebollar a Peranzaes, entre la Peña del Trayecto y el alto de la Gubia del Portillón, 24 September 1990, R. Gamarra & F. Castilla 542 (MA488315!); Cangas de Narcea, Reserva Biológica de Muniellos, 27 September 1990, R. Gamarra & F. Castilla 603 (MA488333!); Cangas de Onís, Margolles, Peruyes, 21 May 1995, M.C. Fernández Ordóñez s.n. (MA570737!); Castañedo, 20 September 2017, S. Molino et al. s.n. (MACB113877!); Cudillero, Ballota (Santa Marina) , 24 August 1996, T. Campón et al 2778AI (MA594170!); Gijón, 1927, Miranda s.n. (MA001471!); inmediaciones de Gijón, 1 July 1926, F. Miranda s.n. (MA001470!); in valle, supra Pajares, 14 July 1892, A.E. Lomax s.n. (MA001450!); La Bobia, 20 August 1956, E. Carreira s.n. (MA205526!); Ligüeria, Sellón, Piloña, 15 November 2000, E. San Miguel & A. González Vega ESM95 (MA767385!); Luarca, cerca del mirador, 27 May 1971, P. Montserrat Recorder s.n. (MA808483!); Otur, 19 July 2017, J.M. Gabriel y Galán s.n. (MACB113876!; MACB113879!; MACB113880!); Oviedo, Sierra del Naranco, 13 November 1981, C. Aedo Pérez s.n. (MA621935!); pr. Mazo de Mon, 11 October 2009, C. Aedo Pérez 17041 (MA796535!); Rozapanera, Rifabar, Espinaredo, Piloña, 23 August 2001, V. Molina Lobo & E. San Miguel ESM320 (MA767385!); Sierra de la Peral, 19 June 1835, M.C. Durieu 160 (P01549409!; P01549455!); Valdés, Paladeperre, 22 March 2016, J.M. Gabriel y Galán s.n. (MACB109612!); Valdés, Luarca, 12 April 2017 J.M. Gabriel y Galán s.n. (MACB109615!); Valles, San Román de Villa, Piloña, 11 August 2001, E. San Miguel ESM312 (MA768386!). Canarias: 1801, 1801, P.M. Broussonet s.n. (US2360521!); Tenerife, Anaga, 25 October 1989, A. Santos s.n. (MA483040!); ibidem (MACB36152!); Tenerife, Anaga, 14 September 2017, J.M. Gabriel y Galán s.n. (MACB110658!); Tenerife, Cruz de Taganana, October 1897, A.C. Cook 1054 (C0355923F!; US537012!; US537013!; US316649!); Tenerife, in apertis humidis jofi la cumbre de Anaga, October 1926, O. Burchard 308 (C0355922F!); Tenerife, Monte de las Mercedes, sobre el barranco de la Mina, hacia la Peña del Inglés, May 1935, J. Cuatrecasas s.n. (MA190831!). Cantabria: Ahedo, 28 December 1983, Prieto et al. s.n. (MA372712!); Ayuela (Uldias), 9 November 1974, E. Loriente s.n. (MA656337!); Camaleño, Cosgaya, 8 October 2016, J.M. Gabriel y Galán s.n. (MACB109622!); Castro-Urdiales, carretera de Samano a Guriezo, 15 July 1968, L.F. Aparicio s.n. (MA755577!); del monte Carrales cerca de Reinosa, no date, B. Salcedo 158 (MA232562!); de Yera al Pto. De las Estacas de Trueba, 26 April 1982, A. Barra s.n. (MA232253!); Fuente Dé, 11 August 1980, A. Romero s.n. (MA503610!); Monte Corona (Comillas), 15 June 1995, A. Guerra s.n. (MA656340!); Monte San Juan (Ribamontán al Monte), 10 March 1983, E. Loriente s.n. (MA656339!); Ogarrio, P. Sáinz Gutierrez s.n. (MA001455!); Pedreña (Marina de Cudeyo), 2 January 1983, E. Loriente s.n. (MA656338!); Portillo de la Sía, 18 July 1995, M. Herrera s.n. (MA570844!); Reserva de Saja, entre Saja y El Tojo, 3 July 1987, E. Villanueva 1252 (MA499610!); Reserva del Saja, río Saja, 10 August 1984, A. Izuzquiza & I. de Aldecoa s.n. (MA506698!); Saja, Los Tojos, E. Loriente s.n. (MA656336!); San Miguel de Aguayo, Santa María de Aguayo, cerca del cruce con la nacional, 22 February 1998, M. Pardo de Santayana 0359 (MA725198!); Santander, no date, I. Zubia s.n. (MA001453!); Santander, Ramales de la Victoria, Ancillo, 19 September 1987, E. Rodríguez & M. Herrera s.n. (MA412401!); Santander, Resconorio, Pto. De la Magdalena, 12 September 1987, C.J. Valle s.n. (MA412536!); San Vicente de la Barquera, 17 August 1983, C. Aedo s.n. (MA621936!); Seña, entrada a la gruta, 20 July 1968, L.F. Aparicio s.n. (MA755575!); Vega de Pas, Pandillo, 21 September 1998, M. Pardo de Santayana & D. Pérez del Molino 0521 (MA725197!); Vega de Pas, Portilla, 2 August 1997, M. Pardo de Santayana 0116 (MA725199!); Vertiente norte del Pto. De Palombera, 25 April 1982, A. Barra s.n. (MA232247!). Castilla la Mancha: Albacete, Cañada de los Mojones, junto al Arroyo de la Puerta, Villaverde del Guadalquivir, 1 October 1988, S. Ríos s.n. (MA508904!); Ciudad Real, Fuencaliente, umbría de Ventillas Garganta de la Chorrera, 17 June 1998, R. García Río s.n. (MA712311!); Ciudad Real, Montes de Toledo, Navas de Estena-Portillo de Cíjara (río Estena), 27 August 1988, D. Sánchez Mata s.n. (MA483500!); Ciudad Real, Puebla de Don Rodrigo, abedular de Río Frío, 3 November 1991, C.J. Martín-Blanco & Garrido 1096 (MA615760!); Ciudad Real, San Lorenzo de Calatrava, manadero de Hocedilla, 25 October 2001, M. Bellet et al. s.n. (MA721877!); Cuenca, Sierra de Valdemeca, 22 July 1974, G. López s.n. (MA226677!); ibidem (MA232425!); Cuenca, Valdemoro- Sierra, Hoz Malilla, 1 September 2004, O. García Cardo OGC-419 (MA744048!); Guadalajara, Cantalojas, valle del Zarzas, barranco de La Laguna, 11 August 1985, J.M. Cardiel s.n. (MA502078!); Toledo, Castella Nova, San Pablo de los Montes, 13 June 1924, E. Gros s.n. (MA001427!); Toledo, Nacimiento del Río Estena, 19 June 1986, N. Marcos 1046MG (MA343502!). Castilla y León: Ávila, Arenas de San Pedro, río Pelayo, March 1987, F. Gómez 1796 (MA653490!); Ávila, Candeleda, 12 October 1986, P. Vargas & M. Luceño s.n. (MA526015!); Ávila, Solana de Ávila, bajando desde la central del Chorro, 12 July 2000, F. Bajo & R. Álvarez 13100RAD (MA649430!); Burgos, Espinosa de los Monteros, río Trueba, subiendo al puerto de Las Estacas, 5 June 1988, A. Izuzquiza & I. de Aldecoa 1455AI (MA430670!); Burgos, San Millán, 27 September 1975, E. Fuentes s.n. (MA806461!); Burgos, Tablada del Rudrón, 28 April 1985, A. Martín & G. Cela 1003 (MA640526!); Burgos, Valmala, Sierra de la Demanda, barranco de La Genciana, en la umbría del monte Trigaza, 29 June 1994, J.A. Alejandre 432/94 (MA554685!); León, Aira da Pedra, 20 July 1993, M. de Godos s.n. (MA711453!); León, Abelgas, 21 August 1995, E. Puente & M.J. López Pacheco s.n. (MA592474!); León, Alto de Pajares, 12 July 1985, A. Penas s.n. (MA545816!); León, Burón, Monte Mirva, October 1962, C.M. Romero s.n. (MA755576!); León, c. 1.5 km E of Puerto del Pontón, 16 July 1985, P. Frost-Olsen 7334 (MO6589949!); León, Monte 457 de U.P. de Acebedo, 1 October 1962, no collector s.n. (MA205524!); León, Ponferrada, Pinar de Puebla de Lillo, 15 November 1987, A. Penas s.n. (MA412438!); León, Ponferrada, montes Aquilianos, camino de S Pedro a S Adrián, 11 June 1982, Ibañez & G. Nieto Feliner 565GN (MA315755!); León, Pto. del Pontón, 23 August 1983, M.P. Fernández Areces & F.J. Pérez Carro s.n. (MA318284!); León, Puerto Ventana, 13 August 1979; J.A. Devesa s.n. (MA503625!); León, Ravanal de Luna, 30 August 1979, J.A. Devesa s.n. (MA503625!); León, S. Teleno, valle del Pelar, 1 July 1946, F. Bernis Madrazo s.n. (MA001441!); Salamanca, Bejar, 18 March 1974, J. Aldasoro s.n. (MA654364!); Salamanca, Carretera a La Alberca, 14 November 1989, F.J. González et al s.n. (MA790486!); Salamanca, carretera de Las Batuecas, la Alberca, 23 June 1946, A. Caballero s.n. (MA001451!); ibidem (MA001452!); Salamanca, Madroñal, 14 November 1989, M. Laredo s.n. (MA570885!); ibidem (MA505326!); Salamanca, Navasfrias, 18 July 1976, E. Rico s.n. (MA205525!); Salamanca, Pastores, Presa del río Agueda, 19 June 1986, E. Rico s.n. (MA390655!); Salamanca, San Miguel de Valero, fuente del Lancharejo, 17 August 2004, A. Izuzquiza &A.Campón 3363 (MA850548!); Salamanca, Sierra de Francia, June 1904, M.Gandoger s.n. (MO1877366!); Segovia, Cerezo de Arriba, La Pinilla, 11 July 1982, T. Romero s.n. (MA566192!); Segovia, Pinar del Balsain, 10 June 1889, Cutanda s.n. (MA001432!); Segovia, Pr. Hayedo de la Pedrosa, 9 October 2007, J. Calvo JC1659 (MA773246!); Segovia, puerto de la Quesera, 19 October 1978, P. Blanco s.n. (MA450684!); Segovia, puerto de la Quesera, Riaza, 19 November 1984, C. García Rodríguez s.n. (MA881298!); Segovia, Sierra de Ayllón, puerto de la Quesera, 26 June 1973, G. López & E. Valdés s.n. (MA236403!); ibidem (MA844727!); Soria, Covaleda, 18 August 1947, E. Guinea s.n. (MA718298!); Soria, Covaleda, 26 August 1959, A. Segura Zubizarreta 3954 (MA352947!); Soria, Covaleda, sierra de Urbión, cabecera del barranco de la Torneda, 8 August 1992, J.A. Alejandre 693/92 (MA527293!); Soria, Laguna Negra, 21 July 1962, A. Segura Zubizarreta 3955 (MA352932!); Soria, Talveila, 22 July 1985, G. Mateo s.n. (MA318269!); ibidem (MA332552!); Soria, Talveila, 23 July 1987, G. Mateo s.n. (MA332552!); Zamora, Aciberos, 23 September 2017, S. Molino et al. s.n. (MACB110655!); ibidem (MACB113878!); Zamora, Figueruela de Arriba, Flechas, 28 February 1998, P. Bariego Hernández PB40 (MA796587!); Zamora, Micéreces de Tera, Canal del Molino, 18 June 2010, C. Perez Morales et al s.n. (MA895314!); Zamora, Requejo, bosque del Tejedelo, 29 September 2008, D. Losada DL225 (MA889814!); Zamora, San Martín de Castañeda, rio Tera, 20 September 1986, A. Roa s.n. (MA510070!); Zamora, Tábara, Monte Las Fuentes , arroyo del Casal, 12 September 1996, P. Bariego Hernández s.n. (MA652085!). Cataluña: Gerona, Camprodon, August 1878, Bolós s.n. (MA001430!); Lérida, Portilló de Bossost, 31 July 1994, L. Sáez s.n. (MA592754!); Lérida, Valle de Arán, September 1848, A.C. Costa 188 (MA001435!). Euskadi: Álava, Inurrieta, 9 June 1925, J. Ruiz de Azúa s.n. (MA001468!); Álava, Lagán, Sierra de Cantabria, 17 August 1973, P. Montserrat & L. Villar s.n. (MA317972!); Álava, Landa, 22 December, J. Ruiz de Azúa s.n. (MA001484!); ibidem, 21 December 1929, J. Ruiz de Azúa s.n. (MA001483!); Álava, Monte Gorbea (vertiente S.S.E.), 6 July 1925, J. Ruiz de Azúa s.n. (MA001485!); Álava, Monte Gorbea, 17 July 1925, J. Ruiz de Azúa s.n. (MA001465!); Álava, Munguia, no date, J. Ruiz de Azúa s.n. (MA001466!); ibidem, 1 July 1929, J. Ruiz de Azúa s.n. (MA001482!); Álava, Santa Engracia, 19 July 1925, J. Ruiz de Azúa s.n. (MA001464!); Álava, Ulibarri Olleros, E Portucho, 10 October 1987, P.M. Uribe-Echebarria s.n. (MA415386!); ibidem (MACB030899!); Álava, Vitoria-Gasteiz, Montes Altos de Vitoria, fuente de las Neveras, 15 November 2004, P.M. Uribe-Echebarria VIT75754 (MA753174!); Guipúzcoa, Beasain, 27 June 1926, J. Ruiz de Azúa s.n. (MA001459!); Guipúzcoa, cerca de Aya, 21 August 1987, D. Gómez s.n. (MA415399!); Guipúzcoa, Escoriaza, 18 June 1925, J. Ruiz de Azúa s.n. (MA001467!); Guipúzcoa, Gudamendi, 1 August 1978, S. García Caballero s.n. (MA502168!); Guipúzcoa, Irún, Urdanibia, 13 May 1983, P. Catalán & I. Aizpuru 612.83 (MA343491!); Guipúzcoa, Jazquivel, May 1895, Gandoger s.n. (MA001460!); Guipúzcoa, Laurgain, 15 September 1991, P. Garin s.n. (MA505223!); Guipúzcoa, Lizarrusti, 21 August 1987, D. Gómez s.n. (MA808484!); Guipúzcoa, Salinas de Leniz, 22 December 1925, J. Ruiz de Azúa s.n. (MA001474!); Guipúzcoa, San Sebastián, no date, no collector s.n. (MA167008!); Guipúzcoa, San Sebastián, junto a las torres de Arbide, 24 October 1995, P.M. Uribe-Echebarria s.n. (MA570772!); Guipúzcoa, Vergara, 24 September 1926, J. Ruiz de Azúa s.n. (MA001461!);Vizcaya, Aránzazu, 28 July 1951, E. Guinea 1418 (MA389236!); Vizcaya, Bakio, Mte Jata, 24 July 1989, J.A. Alejandre 978/89 (C0379911F!; MA484836!); Vizcaya, Bedia, 15 August 1925, J. Ruiz de Azúa s.n. (MA001463!); Vizcaya, Carretera de Lequeitio a Ondarroa, 16 August 1947, E. Guinea 2187 (MA165111!); Vizcaya, Durango, 7 January 1926, J. Ruiz de Azúa s.n. (MA001458!); Vizcaya, Galdácano, 22 December 1924, J. Ruiz de Azúa s.n. (MA001440!); ibidem (MA001469!); Vizcaya, Lemona, 21 August 1925, J. Ruiz de Azúa s.n. (MA001462!); Vizcaya, Mte. Jata, 7 April 1989, J.A. Alejandre 33/89 (MA484835!); Vizcaya, Ochandiano, 4 September 1925, J. Ruiz de Azúa s.n. (MA001457!); ibidem (MA001486!); Vizcaya, Uberuaga, no date, I. Zubia s.n. (MA001438!); ibidem (MA001439!). Extremadura: Cáceres, Arroyo de Sierra de Valdecaballero, 30 September 1978, J.L. Pérez Chiscano s.n. (MA227193!); Cáceres, Baños, no date, no collector (MA226679!); Cáceres, Borde de fuente, Cañamero, 26 July 1993, J.L. Pérez-Chiscano s.n. (MA531011!); Cáceres, Casas del Miravete, 15 May 1980, D. Belmonte s.n. (MA392824!); Cáceres, entre Alcorneo y Valencia de Alcántara, 17 May 1992, E. Rico s.n. (MA519388!); Cáceres, entre Descargamaría y Martiago, 8 August 1990, R. Gamarra & F. Castilla s.n. (MA488475!); Cáceres, Monfragüe, Ribera del río Tajo, 29 May 1986, E. Rico s.n. (MA390982!); Cáceres, Valencia de Alcántara, El Pino de Valencia de Alcántara, 14 November 1987, M.C. Viera 104/97 (MA503568!); Cáceres, Valencia de Alcántara, Jola, 30 October 1987, G. Fernández s.n. (MA503624!). Galicia: no date, M. Colmeiro s.n. (MA232665!); 1897, Merino s.n. (MA001443!); A Coruña, A Baña, río de Albariña, 29 December 1996, L.G. Quintanilla & B. Pias 22 (MA590970!); A Coruña, A Baña, río de Portalexe, 5 July 1997, L.G. Quintanilla 104 (MA590984!); A Coruña, Abegondo, Robledal del perfil, no date, H. Villar s.n. (MA163807!); A Coruña, As Pontes de García Rodríguez, 17 June 1999, Gil s.n. (MA649754!); A Coruña, Brañas de Brius, 7 April 1964, no collector (MA824008!); A Coruña, Cariño, Landoy, Río das Mestas, bajo Chan da Capela, 20 June 2017, R. Fernández Perales et al 21 (MA917905!); A Coruña, Caveiro (Puentedeume), 17 April 1991, M. Hermida s.n. (MA745386!); A Coruña, Lousame, río de San Xusto, 26 February 1997, L. Gómez-Orellana & L.G. Quintanilla 73 (MA591006!; intermediate form between S. spicant var. spicant and var. homophyllum ); A Coruña, Monfero, Caveiro, 18 October 1986, S. González Crespo s.n. (MA382853!); ibidem (MACB031064!); A Coruña, Monfero, Xestoso, a orillas del río Frai Bermui, 10 November 1991, F.J. Silva-Pando s.n. (MA505181!); A Coruña, Noya, 21 August 1967, Casaseca & Álvarez s.n. (MA185026!); ibidem (MA487075!); A Coruña, Playa de Boiro, 26 August 1969, no collector (MA717419!); A Coruña, río Eume-Fuentedeume, 2 August 1968, Carballal s.n. (MA478950!); A Coruña, Santa Comba, O Pedroso, Fonte Barreña, 22 June 2017, R. Fernández Perales et al 142 (MA918020!); A Coruña, Santa Cruz, September 1904, Bescansa s.n. (MA150175!); A Coruña, Santiago, 24 August 1967, F. Bellot Rodríguez s.n. (MA785146!); ibidem, 29 July 1967, no collector (MA824045!); A Coruña, Santiago de Compostela, Cantaleta, 29 July 1967, Álvarez s.n. (MA198343!); A Coruña, Santiago de Galicia, en el Paular, en San Ildefonso, 29 July 1782, A.J. Cavanilles s.n. (MA232869!); A Coruña, Santiago O. Pino, Arca, rio Braudelos, cuenca del Ulla, 9 July 1991, M.J. Romero s.n. (MA546613!); A Coruña, Zas, Coto de Muiño, 8 November 1989, A. Estévez González & M.P. González Hernández s.n. (MA483063!); Lugo, 24 August 1986, E. Carreira s.n. (MA462832!); Lugo, Mouromorto, 27 May 1971, P. Montserrat Recorder s.n. (MA808482!); Lugo, Murás, Puente Rioboo, 2 August 1987, F.J. Silva Pando et al. s.n. (MA412515!); ibidem (MACB030875!); Lugo, Riberas de León, 25 July 1956, E. Carreira s.n. (MA201118!); Lugo, río Rato, 24 August 1984, E. Carreira s.n. (MA391545!); Lugo, Sierra del Caurel, Mercurín, 10 July 1994, E. Blanco 1182EBC (MA564805!); Lugo, Villardiaz-Fonsagrada, 8 July 1951, E. Carreira s.n. (MA172742!); ibidem 7 August 1954, E. Carreira s.n. (MA172743!); ibidem, 10 July 1957, E. Carreira s.n. (MA722139!); Ourense, Barco de Valdeorras, tras de Gandara, 25 July 1987, C. Prada s.n. (MA412504!); Ourense, Sª de Santa Marta, Castrelo de Minho, 18 July 1935, A. Rodríguez s.n. (MA001442!); Pontevedra, Castro-Loureiro, Ju 1910, A. Casares Gil s.n. (MA001444!); ibidem (MA001445!); ibidem (MA001446!); ibidem (MA001448!); Pontevedra, Ermelo, Bueu, 10 October 1969, S. Castroviejo s.n. (MA198345!); Pontevedra, estación de Redondela, 4 April 1969, E. Valdés-Bermejo s.n. (MA855764!); Huertas de Marin, 8 August 1934, J. González-Albo s.n. (MA001456!); Pontevedra Lourizan, 20 April 1960, A. Rodríguez s.n. (MA201121!); ibidem (MA226674!); ibidem (MA226675!); ibidem (MA226676!); ibidem (MA201121!); ibidem (MA226676!); Pontevedra, Lovios, Sta. Eufemia, 6 November 1991, P. González Hernández & J. Vegas Rodríguez s.n. (MA505180!); Pontevedra, Marin, 1934, J. González-albo s.n. (MA001447!); Pontevedra, Mondariz, 20 May 2016, J.M. Gabriel y Galán s.n. (MACB113881!); ibidem (MACB113883!); ibidem (MACB113884!); Pontevedra, Portela, 27 March 1970, E. Valdés-Bermejo s.n. (MA857233!); Pontevedra, San Adrián, 10 October 1970, Carballal s.n. (MA313241!); Pontevedra, Monte Aloya, Tuy, 15 July 1993, S. Pajarón & E. Pangua s.n. (MA540648!); ibidem (MACB59142!); Pontevedra, Redondela, 1 November 1976, I. Barrera s.n. (MACB004769!); Pontevedra, Vigo, 26 April 1955, E. Guinea s.n. (MA228468!). La Rioja: Daroca de Rioja, barranco de Cabañas en la umbría de la sierra Monvcalvillo, 8 September 1995, J.A. Alejandre 2585/95 (MA565006!); Puerto Piqueras, 8 August 1978, F. Amich s.n. (MA213351!); Rasillo de Carmeros, no date, Iñiguez s.n. (MA001431!); Santa Coloma, sierra de Moncalvillo, junto al arroyo en el barranco del Manadero, 14 October 1995, J.A. Alejandre 2589/95 (MA565007!). Madrid: Dehesa de Somosierra, 31 May 2017, J.M. Gabriel y Galán & S. Molino s.n. (MACB109611!); La Hiruela, abedular de La Dehesa, sobre el Jarama, 18 July 2007, L.M. Ferrero et al LM4199 (MA772506!); Montejo de la Sierra, El Chaparral, 25 August 1908, M. Rodríguez & López-Neyra 2273 (MA346270!); ibidem, M. Rodríguez & López-Neyra s.n. (MA346271!); pista forestal de Peña Cebollera, pr. Somosierra, 26 March 1982, J.J. González AB2138 (MA232435!); Puerto de Canencia, 12 June 1973, F. Bellot Rodríguez s.n. (MA699023!); Peñalara, Sierra de Guadarrama, July 1912, A. Casares-Gil & F. Beltrán s.n. (MA001424!); Sierra de Guadarrama, Cercedilla, July 1914, C. Vicioso Martínez s.n. (MA001425!); Somosierra, no date, E. Reyes Prósper s.n. (MA001426!). Navarra: August 1876, J.M. Laurrgueta s.n. (MA001437!); Castillo Nuevo, Leyre, 27 September 1972, L. Villar & P. Montserrat (MA808485!); Irati, río Urchuria, 3 September 1993, E. Rodríguez & M. Herrera s.n. (MA540678!); Isaba, Belagua, 27 August 1973, L. Villar s.n. (MA808486!); Monte Errekerena, Esteribar, 25 September 1989, J. Loidi s.n. (MA483527!); Puerto de Echegarate, cercanías del Restaurante Alaska, 1 May 1973, S. Castroviejo s.n. (MA226673!); Roncesvalles, subiendo al Puerto de Ibañeta, 5August 1973, E. Fuertes et al s.n. (MA792927!); territorio Vestizararensi, 7 August 1876, J.M. Lacoizqueta s.n. (MA001454!). Unknown location: no date, no collector (MA163806!); July 1782, J.A. Cavanilles s.n. (MA232867!); Sierra de Fontelome, 9 July 1943, O. Rodríguez s.n. (MA346305!). SWEDEN. Dalarna: Falun, 16 August 1895, C. Hammso 19753 [interpreted] (P01549375!). Norrland: Jamtland, 18 July 1883, C. Indebetou s.n. (MA001414!). Götaland: Småland, no date, N.J. Andersson s.n. (MA153040!). Skåne, Hörby, Härröd, August 1924, A. Edv. Gorton s.n. (MO2482457!); Skåne: Smygehamn, 13 July 1974, illegible collector (MO6544346!). Unknown location: no date, no collector (C0379912F!); no date, N.J. Andersson s.n. (MO873120!); ibidem (MO873972!); June 1882, L.J. Wahlstedt s.n. (C0379904F!); 1885, illegible collector (C0379905F!); August 1895, C.F. Curtiss s.n. (C0379960F!); August 1902, A.M. Nordström s.n. (C0379927F!); 19 November 1907, A.G. Eliasson 20 (MO1153566!). SWITZERLAND. Berne: environs du Col de Jaun, 17 September 1964, B. de Retz 52491 (P01567055!). Schwyz: Mt. Rigi, 1883, B. Fink Herb. s.n. (MO1877374!); Unteriberg, no date, Hoppe 21126 (P01549389!) Tesino: Lugano, 21 February 1900, illegible collector (P01549379!). Valais: Haut-Valais, 26 July 1873, A. Thielens s.n. (C0379925F!); Kanton Wallis, zwischen Oberwald und Gletsch im Lärchenwald, August 1903, F.O. Wolk s.n. (MO2360824!); la Crettaz sur Salvan Bois mossus, Epicea et Mélèze, 23 September 1950, C. Oberson 456 (P01567056!). Unknown location: no date, no collector s.n. (C0379924F!); October 1866, R. Jalling [interpreted] (MO873964!); 3 August 1870, Zollikofer s.n. (P01658104!); September 1873, A.J. Skutch s.n. [interpreted] (C0379908F!); 29 September 1906, Missbarh s.n. [interpreted] (P01417528!); 18 July 1960, Binz 3743 [interpreted] (MO3180251!). TURKEY. Rize: 6 km antes de Ikizdere, 28 June 2001, S. Nisa de Oliveira 715 (MA688910!). UKRAINE. Chernivtsí: environs of village Shepit, 12 July 1957, T.J. Omelczuk s.n. (MO5788513!). Leopolis: Galitzia, Juñ 1873, Illegible collector (P01549393!); ibidem, 19 July 1873, Illegible collector (P01549391!). Vorokhta settlement hromada: Ivano-Frankivsk Region, vicinity Vorohta, Goverlyanske forestry, spruce-silver fir forest, 16 July 1957, T.J. Omelczuk s.n. (MO5759224!). UNITED KINGDOM. England: no date, J.Y. Braeas [interpreted] (C0379892F!); 2 September 1903, L.V. Lester-Garland s.n. (C0379929F!); 31 July 1931, no collector (C0379890F!); Dartmoot, Devon, Becky Falls, 10 June 1924, D.S. Johnson s.n. (C0379909F!); Devonshire, 13 July 1924, R.A. Ware s.n. (MO1055891!); Hampstead, M. Lagasca y Segura s.n. (MA167006!); Hastings, October 1866, Miss J.E. Wilkins s.n. (C0379910F!); South Somerset, Seven Wells Wood, 30 July 1968, J.A. Crabbe et al. 11822 (MA195093!; P01567059!); Surrey, Leith Hill, 1914, E. Crespigny s.n. (P01549398!). Scotland: Argyll and Bute, Mid Ebudes, Mull, Tobermory, 6 July 1970, J.A. Crabbe & A.C. Jermy 11837 (MA195091!); Braemar, the Cairnwel, 11 August 1970, S. Castroviejo s.n. (MA695159!); east Sussex, Mark Cross parish, Saxonbury Hill, 6 km S of Tunbridge Wells, 20 July 1967, A.C. Jeremy & J.A. Crabbe 11788 (MA195092!); Granpian Hills, no date, Redfreld s.n. (MO1877371!); Isle of Arran, 2 ½ miles southwest of Lochranza, near track, 10 June 1973, L.J. Tuttle 103 (MO6794402!); Loch Katrine, 26 June 1901, E.T. Harper s.n. (C0379961F!); south Somerset, over stowey parish, Seven Wells Wood, 3 km S of Nether Sowey, 30 July 1968, J.A. Crabbe 11822 (MA195093!). Wales: Cafrmarthen, at Bot Fawr Forest, 20 August 1963, J.P.D.W. Payens 286 (MA477087!); Llangollen, 25 August 1892, A.E. Lomax s.n. (MO1877382!); Merionethshire, v.c. 48, Cader Idris, south-west of Dolgellau, July 1961, R.K. Brummitt 61782 (MA186639!); Pysgodlyn Mawr, Glamorganshire, 7 October 1986, G. Hutchinson & B.A. Thomas s.n. (MA477087!). Unknown location: October 1847, M. Howae s.n. (P01549408!). UNITED STATES OF AMERICA. Alaska: 1865–1866, Western Union Extension Telegraph Exploring Expedition (C0602102F!); Baranoff Island, 15 June 1899, Wm. Trelease 2707 & A. Saunders (MO1876815!); Bluff of Salmon river near bridge, 19 July 1924, K. Whited 1314 (MO926810!); Chomley, 6 August 1929, G.B. Rigg 81 (C0602131F!); Cordova, 10 September 1939, J.H. Steenis 4799 (MO1570445!); Evans island, Port San Juan, 29 September 1948, W.J. Eyerdam 7268 (MO1590419!; MO5460334!); Hope Island, 1867, A. Kellogg s.n. (C0602096F!); Kaasan Bay, 26 July 1901, C.F. Newcombe 162 (C0602101F!); Kake, Luxuriant forest, 12 September 1928, Y. Mexia 2313 (MO970428!); Ketchikan area, Higgins Rd., 7 April 1963, B.J. Neiland 800 (MO3729464!); Kuiu Island, Washington Bay, 1 September 1955, W.J. Eyerdam 8177 (C0602130F!; MO5791710!); Lawson Bay, 1867, A. Kelbert 51* (US375667!); McDonald Lake, 15 July 1921, H.M. Smith s.n. (US1325588!); McDonald Lake region, 19 July 1905, J.L. Burcham 21 (US422146!); New Metlakahtla, 4 June 1899, Wm. Trelease 2706 & A. Saunders (MO1876816!); Port Althrorp, 12 August 1927, H.C. McMillin s.n. (US2255439!); Port Ashton, 14 August 1936, Shulman 54 (MO1967968!); ibidem, Shulman 57 (MO1967967!); Prince of Wales Island, Klawak Lake, 12 September 1915, E.P. Walker 1004 Mrs. E.P. Walker (C0602143F!; MO822061!); Prince William Sound, Orca, 24 June 1899, Wm. Trelease 2708 & A. Saunders (MO1876813!); ibidem, 16 August 1936, I.L. Norberg s.n. (US2291321!); Port Etches (Hinchinbrooke Island, Prince William Sound), 27‒29 August 1837, G.W. Barclay 1426 (C0602149F!; US3100237!); Sitka, 13 August 1892, B.W. Evermann 197 (C0602155F!; MO847389!); Stika and vicinity, July 1891, W.G. Wright 1613 (MO1876812!); Sitka, Harriman Alaska Expedition, 14‒17 June 1899, F.V. Coville & T.H. Kearney 860 (MO1876814!; US421424!); Stevens Pass, Cascades, 18August 1893, J.H. Sandberg & J.B. Leiberg 774 (US275667!); Thum bay, Knoght island, Prince William Sound, 28 September 1939, W.J. Wywedam 3388 (MO1190672!); Windham Bay, 10 September 1905 [interpreted], J.D. Culbertson 4963 (MO1876811!; US666096!); Yakobi Island, 14 July 1915, E.P. Walker 846 & Mrs. Walker (C0602119F!; MO822065!; US886881!); ibidem, 7 August 1921, J.P. Anderson 1325 (C0602137F!); Yakobi Island, Bohemia Basin, 13 June 1957, H.T. Shacklette 3864 (US2416216!). Arizona: Graham, South of San Carlos Reservoir, 19 June 1938, M. Ownbey 1648 & R.P. Ownbey (C0602074F!). California: no date, S.C. Bagg s.n. (C0602087F!); 2 November 1905, H.N. Bolander s.n. (MO1876841!); 2 April 1950, P.H. Raven 1539 (MO1735285!); ¼ mi below fork, Waddell Creek, Santa Cruz Co, 14 September 1941, R.W. Storer 156 (US2256485!); 2 miles SE Mendocino, 17 May 1947, R.W. Storer 401 (US2256636!); along the E. side of Mitchell Creek Road and about 1.3 mi S of the intersection with Simpson Lane, 21 December 2004, D. Goldman 2992 (P01625270!); del Norte Co., Smith River, 28 July 1940, D. Desmaree 21448 (MO1876844!); del Norte Co., Smith River, July 1958, Bronscomb 2 (MO5460329!); Eel River, Humboldt Co., 20 September 1920, R.K. Beattie 5935 (MO5460330!; US2201149!); Humboldt, 8 July 1888, V.K. Chesnut 3712 (C0602081F!); Inverness ridge, northwest of Inverness, Marin County, 2 February 1941, R.F. Hoover 4740 (US2019225!); Little River State Park, 25 August 1957, D. Branscomb s.n. (US2422496!); Marin Co., Tomales Bay, 5 November 1933, J.A. Ewan 8105 (MO1069570!); Mariposa, Mendocino city, 9 August 1901, no collector s.n. (US466493!); Mendocino Co., 27 August 1957, H.N. Bolander 1 (MO5460328!); Mendocino Co., May 1866, H.N. Bolander 6237 (C0602075F!; MO1876842!; US319019!; US319020!); Mendocino Co., 10 July 1931, M. Jones 19180 (MO1013609!); near Sheltter Cove, Humboldt Co, 31 July 1869, A. Kellogg & W.G.W. Halford 1175 (MO1876840!; US324088!); Northern California, Mendocino County, Albion Ridge, July 1909, J. McMurphy 451 (US610983!); on Baltic Creek, about 30 mi NE of Placerville, 5 July 1901, W.G. Watkins 12 (US413126!); Pigmy forest, 1 ½ mi NE Albion, Mendocino Co., R.W. Storer 397 (US2256553!); Sansalito, 24 June 1892, M.A. Howe s.n. (US591517!); Santa Cruz Co., 23 June 1881, M.E. Jones 2232 (P01462469!; US220497!); Santa Cruz Co., August 1888, L.M. Underwood s.n. (US644165!); Santa Cruz Co., 17 January 1921, R.S. Ferris 2107 (MO1876843!); Santa Cruz Co, 2 miles SE of Bonny Doon, Laguna Canyon, 19 December 1965, G. Ledyard Stebbins 6300 (MO6257789!); Santa Cruz Co., along west fork of Waddell Creek about 5 miles inland from the coast, 16 August 1959, J.H. Thomas 8068 (US2852893!); Siskiyou Mts. 3 2/ 10 mi above Smith River (Town) on Rd. Tom High Divide, County del Norte, C.B. Wolf 4020 (US2201135!); Thompson Ranch, 4 miles north of Crescent City, Del Norte Co, 15 august 1946, A. Carter 1772 (C0602038F!; MO5460331!; MO1652670!; US2259134!); Van Damme State Park, Mendocino County, 23 December 1957, D. Branscomb s.n. (US2257262!); Willow Creek, near Eaverville, 15 July 1931, M. Jones 29179 (MO1013476!). Idaho: Clearwater, 12 miles east of Weippe, 17 April 1930, H.J. Rust 1156 (US1534579!). Indiana: LaPorte, Valley of Trail Creek near the town of Trail Creek, 15 August 1934, R.M. Tryon 210 (C0602090F!). Oregon: no date, L. Summers s.n. (US967834!); 1838 to 1842, Wilkes Explor. Exped. s.n. (US98664!); 1875, J. Horwell s.n. (C0602050F!); 1876, J. Horwell et al s.n. (C0602039F!); May 1879, J. Howell & T.J. Howell (C0602026F!); ibidem (MO1876834!); 30 June 1879, J. Howell s.n. (C0602080F!); 13 January 1880, J. Howell & M.O. Rust 9013 (P01462465!); 4 September 1891, M. Craig 1007 (MO1876837!); 1871, E. Hall 680 (C0602062F!; C0602086F!; MO1876832!; P01462461!; US bc1523579!); 15 September 1881, C.G. Pringle s.n. (C0602068F!); 30 November 1883, W.C. Ohlendorf s.n. (C0602069F!); September 1915, C.G. Pringle s.n. (C0602044F!); ibidem (P01462460!); ibidem (P01462463!); September 1924, L.F. Henderson 760 (MO918276!); ½ mile east of Scottsburg, 19 March 1942, R.R. Belton s.n. (US2467153!); Astoria, 1866, A. Wood s.n. (MO873122!); Astoria, Clatsop Co., 31 July to 1 August 1915, W.W. Eggleston 11452 (US768158!); Benton Co., Mary’s Peak, 3 August 1986, H.N. Moldenke 32073 & A. Moldenke (MO3674116!); Cascade mountains, 5 October 1886, J. Howell & T. Howell s.n. (MO1876835!); ibidem (P01462467!); Hillsboro, July 1901, J.E. Kirkwood s.n. (US692259!); Clackamas Co., August 1899, A.D.E. Elmer 1620 (MO1876836!); Clackamas Co., Mount Hood, Government Camp, 9 July 1934, D.S. Johnson s.n. (C0602057F!); ibidem (MO5460332!); ibidem, 31 July 1938, M. Owabay & R.P. Owabay 1805 (MO1190407!); Coos Bay, 20 August 1911, H.D. House 4736 (US623071!); Cypress forests of Oregon, no date, J.G. Lemmon s.n. (C0602027F!); ibidem (US967835!); Grays Harbor Co., about 5 miles south of Lake Quinault, 23 June 1941, H.T. Rogers 819 (MO1282520!); Lane Co, in coniferous woods across the Siuslaw River from Florence, Twp. 18 S, R. 12 W, 28 August 1949, A.J. Cronquist 6116 (US2017759!); Lincoln Co, 3 mi N of Florence at junction with Mercer Lake Rd, 25 August 1962, F.R. Fosberg 43183 (US2675147!); Lincoln Co, along stream, Cape Creek, Perpetua Forest Camp, 29 November 1952, A.N. Steward s.n. (US2414117!); Lincoln Co., east of Waldport, 1.3 mi, E of Hwy 101 on Wakonda Beach Rd., Siuslaw Nat. Forest, 22 June 1977, C. Davidson 6052 (US3586564!); Lincoln Co., Otis Junction, 26 July 1940, D. Demaree 21415 (MO1876839!); Lincoln Co., Yachats, along the Yachats River, 6 August 1928, L. Constance s.n. (MO1876838!); Marion Co., Willamette National Forest, along Forest Service Road 4965, 2 miles E of its intersection with Forest Service Road, 6 miles NE of Detroit, 23 July 1993, Merello et al 855 (MO4402396!); McMinnville, 1876, Summers 127 (US834117!); McMinnville, 17 September 1878, Summers s.n. (US724007!); Multnomah, Portland, Willamette Heights, 24 August 1902, E.P. Sheldon 11262 (C0602051F!); near Summit House, Clackamas Co, 2 September 1897, F.V. Coville & E.I. Applegate 744 (US381024!); near Willamett slough, May 1879, J. Howell& T. Howell s.n. (MO1876833!); Pacific Coast, Foley Springs, 16 July1903, M.W.Gorman 1663 (US525681!); Pacific Coast, Western Oregon, August 1880, T.J. Howell s.n. (C0602020F!); ibidem (C0602032F!); ibidem (C0602045F!); ibidem (C0602056F!); ibidem (C0602416F); ibidem (P01462459!); ibidem (P01462462!); ibidem (P01462466!); ibidem (P01462468!); ibidem (US834115!); ibidem (US834116!); Pacific Slopes, by mountain rills, 15 September 1881, C.G. Pringle 2713 (MO1876830! US724005!; US834114!); Portland, June 1890, J.H. Hicks s.n. (C0602021F!); Portland, 7 May 1905, W. Palmer s.n. (US1488133!); Portland, 10 May 1905, W. Palmer s.n. (US1488129!); Portland, 2 June 1905, W. Palmer s.n. (US1488132!); Portland, 20 October 1905, W. Palmer s.n. (US1488131!); ibidem (US1488130!); Still Creek Camp, Mount Hood National Forest, 5 August 1927, E.L. Evinger s.n. (US2467167!); Tilamook Valley, 1883, Summers 11868 (P01462464!); vicinity of Oregon city, 18 June 1905, M.W. Lyon Jr 79 (US622426!); vicinity of Portland, 10 May 1905, W. Palmer 1450 (US620278!); vicinity of Portland, 22 June 1905, M.W. Lyon 94 (US622437!); Willamette Heights, Portland, 24 August 1902, E.P. Sheldon S11262 (US434884!); Yamhill, 29 June 1893, W.J. Spillman 214 (C0602063F!). Washington: August 1893, J.H. Sandberg & J.B. Leiberg s.n. (MO1876829!); 11 May 1934, O.J. Marie 1110 (MO1086174!); base of Mt. Paddo, 6 August 1885, W.N. Suksdorf s.n. (C0602148F!); ibidem (MO1876826!) ibidem (P01462441!); ibidem (P01462448!); ibidem (P01462451!); ibidem (P01462456!); ibidem, W.N. Suksdorf 830 (P01462449!); Bogachiel River, 4 April 1934, O.J. Marie 1244 (MO1086214!); Callam Co., along state route 112 just northeast of Pysht, 9 August 1996, J.S. Miller et al 8711 (MO05050899!); Callam Co., logged area 12 miles W. of Lake Crescent, 23 June 1940, I.L. Wiggins 9443 (MO1876825!); Chehalis Co., Montesano, August 1919, J.M. Grant s.n. (MO862793!); Chehalis Co., near Montesano, 30 June 1898, A.A. Heller & E.G. Heller 3993 (MO1876827!; US333184!); Clark Co., Camas, June 1927, D.B. s.n. (MO941722!); Clallam Co., Olympic Mts., June 1990, A.D.E. Elmer 2803 (MO1876822!); Clallam Co. Olympic National Forest, Elk Brake, 25 August 1921, H. St. John 4779 (MO1282452!); Clallam Co., Olumpic National Forest, near nature trail, Rain Forest Visitor Center, West Side, 18 June 1966, D.T. Bell 277 (MO6796407!); Clallam Co., Sequim, August 1915, J.M. Grant s.n. (MO787460!); Cumberland, 10 September 1917, W.H. Meyer s.n. (C0602088F!); Grays Harbor, Hoquiam, 11 July 1907, H.C. Cowles 572 (C0602095F!; MO746290!); Grays Harbor Co., Lake Quinault, in vicinity of Willaby Forest Camp, on South Shore Rd., 0.3 mi south of Quinault, 28 June 1970, S. Gleissman 563 SRG (MO6885583!); Grays Harbor Co., near Lake Quinault, 10 July 1931, J.W. Thompson 7324 (MO1010104!); Jefferson, along road to Hoh Rain Forest, off U.S. Route 101 approximately 14 miles south of Forks, 22 August 1969, R.G. Stolze 1340 (C0602106F!); Jefferson Co., Clearwater trail vicinity Braden Cr., 27 July 1924, I.C. Otis 1316 (US1316812!); Jefferson Co., on brench, Clearwater trail bet. Braden Vr.And Hoh River, 1 October 1924, I.C. Otis 1395 (US1316816!); King Co., 20 June‒12 July 1898, T.E. Savage et al. s.n. (C0602154F!); ibidem (MO1876823!); King Co., Brooklyn, 20 June – 12 July 1898, T.E. Savage et al. s.n. (C0602142F!); ibidem (MO1876824!); Co. King, Skykomish, August 1901, J.M. Westgate 958 (C0602118F!); King Co., Snoqualmie Pass, 17 September1932, J.W.Thompson 8928 (MO1030988!); King Co., Seattle, September 1888, C.V. Piper 10007 (C0602125F!); ibidem, September 1892, E.A. Shumway s.n. (MO1876728!); ibidem, 1 July 1915, G.W. Freiberg (MO813660!); King Co., Skykomish, August 1901, J.M. Westgate 903 (C0602112F!); ibidem, August 1901, J.M. Westgate 935 (C0602124F!); Kittitas Co., Easton, 15 July 1930, E.J. Palmer 37900 (MO1003177!); Klickitat Co., Bingen, August 1907, W.N. Suksdorf s.n. (P01462453!); ibidem (P01462454!); Klickitat County, Falcon valley, July 1908, W.N. Suksdorf s.n. (C0602094F!); ibidem (P01462447!); Klickitat, Mount Binger, 30 June 1908, W.N. Suksdorf s.n. (C0602101F!); Olympia, October 1905, E.C. Townsend s.n. (MO1214041!); Olympic Península, 15 June 1973, L.H. Potter s.n. (MO3101388!); Pierce Co., Puyallup, 17 July 1880, G. Engelmann s.n. (MO1876820!); Quiniault, Jefferson, Chehalis and Thurston Co. Granville, 8 June 1902, H.S. Conard 345 (US526740!); Seattle, August 1898, C.V. Piper s.n. (US102542!); Skamania County, 12 August 1899, J.B. Flett 921 (P01462450!); Seattle, 15 October 1910, E.M. Bandell & J.C. Frye s.n. (P01462444!); Snohomish Co, Lee Forest, 0.50 of a mile east of State Highway 1A, approximately 4 miles north of Woodinville, 23 April 1963, D.R. Simpson 61 (C0602136F!); Snohomish Co., Maryville, August 1924, J.M. Grant s.n. (MO922716!); Snohomish Co., Mt. Baker/Snoqualmie National Forest, FR 20, 1 mi N of Barlow Pass, 26 September 1998, A. Colwell & P. Reeves 142 (MO04895651!); Snohomish Co., trail up Perry Creek, ca. 30 miles east of Granite Falls, Mt. Baker National Forest, 22 August 1969, J.D. Montgomery 85030j (MO5460333!); Stevens Pass, Cascade, 18 August 1892, J.H. Sandberg & J.B. Leiberg 774 (P01462438!); Tacoma, 19 September 1902, J.B. Flett 2030 (P01462452!); Thurston Co., moist coniferous woods near the head of Mud Bay, about five miles west of Olympia, 15 August 1936, F.G. Meyer 792 (MO1313447!); Tummel city [interpreted], August 1893, J.H. Sandberg & J.B. Leiberg s.n. (P01462442!); Whatcom Co., below Mt. Baker Lodge, N. Fork Nooksack, 24 July 1930, L. Benson 2367 (MO989086!); Whatcom Co., Mount Baker area, 9 September 1945, P.O. Schallert 5249 (C0602089F!; C0602107F!); Whatcom Co. Timbered slopes below Welcome Pass, Mt. Baker region, 20 August 1931, J.W. Thompson 8057 (MO1009241!). Unknown location: 1872, Hoppe s.n. (P01549390!); April 1883, G.E. Davenport s.n. (P01462458!). EUROPE (unknown location). No date, no collector (P01549451!); ibidem (MO1877!390!); ibidem (MO1877391!); ibidem (MO1877392!); ibidem (MO937727!); no date, illegible collector (MO1877386!); no date, Kühlewein s.n. (C0379941F!); no date, Scott 600 [interpreted] (MO1877385!); 1864, illegible collector (MO872277!); 1895, no collector s.n. (C0379931F!); no date, Hoppe 1823 (P01417527!); no date, H. Wagner s.n. (C0379899F!); 1838, Martubs 223 (P01549405!); June 1846, illegible collector s.n. (P01549377!); 16 July 1889, illegible collector 16334 (P01549380!); no date, Ex Herb. Danty d’Insard (P01549403!).

Struthiopteris spicant (L.) Weiss var. homophyllum ( Merino, 1898: 108) Gabriel y Galán & Pino in Wasowicz et al. (2017b: 198). ( Figure 9G ; Figure 14G ; Figure 18A ).

Type:— SPAIN. Galicia, Fig Ic. Homophyllum blechniforme in Merino (1898: Lám I), lectotype, designated in Wasowicz et al. (2017a).

Synonyms:— ≡ Homophyllum blechniforme Merino. Blechnum spicant ( L.) Roth var. homophyllum (Merino) Merino (1909: 488) .

Blechnum homophyllum (Merino) Sampaio (1934: 66) .

Invalid designations: Blechnum spicant var. (sine nomine) Christ (1917: 8).

Plants terrestrial, rhizomes short, covered with triangular-lanceolate, concolorous, brownish-clear scales, without sclerosed centre or slightly sclerosed, (0.5) 3‒5 x (0.4) 1‒5 mm; fronds (6) 12 (22) cm, subdimorphic, rosette-arranged, sterile and fertile fronds about the same size, shortly petiolate, stipes dark brown, black or purplish, 0.3‒3 cm, laminae lanceolate, pinnatipartite or pinnatisect, 0.8‒3 x 5‒17 cm, apex pinnatifid, rarely conform, with uniseriate, multicellular hairs, topped by a red gland, rachis green or brown towards apex, with glandular hairs like those on the lamina, pinnae tapering towards the base, the middle ones 0.1‒0.4 x 0.6‒2 cm, slightly contracted on fertile fronds (0.1‒0.3 cm); veins free, simple to 1-furcate, catadromous, ending in adaxial submarginal hydathodes, stomata aligned following veins on abaxial face of pinnae; sori linear on both sides of costa, fragmented, decurrent towards rachis; indusia complex, fragmented; sporangia with 12‒18 cells in arc; spores monolete, 20‒40 x 30‒50 µm, perispores brown, rugate.

Habitat and distribution:— Northwest and west of the Iberian Peninsula, Pontevedra, La Coruña, Orense, Salamanca, Segovia, Salamanca, and Álava ( Spain) and Minho ( Portugal) and the Macaronesian archipelagos of Azores (San Miguel) and Madeira ( Figure 2 ). Moist, shady slopes, usually disturbed, almost always in southwestern orientation. Sea level to 500 m.

Chromosome number:— n = 68, tetraploid ( Löve & Löve 1968, Horjales et al. 1990).

Etymology:— The term homophyllum refers to the subdimorphic condition of this taxon.

Taxonomic notes:— This variety can be distinguished from S. spicant var. spicant by its smaller size (20 cm or less) and subdimorphic fronds, with more triangular pinnae and discontinuous sori. It differs from S. spicant var. pradae also in size, as the pradae variety is similar in size to the type variety. In addition, in the variety pradae there are sporophylls with the same appearance as the trophophylls, without subdimorphism, which coexist with dimorphic contracted sporophylls typical of the type variety, a feature that does not occur in the variety homophyllum . Löve & Löve (1966) cultivated some Spanish specimens and reported that, surprisingly, they maintained their characteristics in culture. However, these same authors report that, contrary to what happens with S. fallax , this variety intermingles a lot with the spicant variety, appearing many intermediate forms. We have corroborated on the field that there are a lot of transitional forms between S. spicant var. homophyllum and the type variety with viable spores, one of the reasons why this variety is not elevated to species status.

The original distribution of this variety was restricted to South of Galicia in Spain and north of Portugal. However, this distribution was extended to Salamanca ( Molino et al. 2019a) and latter on to Azores and Madeira ( Molino et al. 2021). In this last work it is noted that the Macaronesian individuals were found in habitats with low nutrient viability, which is consistent with the hypothesis of the positive association between leaf dimorphism and resource-rich habitats (Britton & Watkins 2016).

Also, these recent distribution extensions point out that probably S. spicant var. homophyllum has a more widely distribution than previously thought, and it would be interesting to look for it in more locations, paying special attention to the species’ range edge, where this variety seems to be more abundant ( Molino et al. 2021).

The type, which must be a specimen collected by Merino in San Juan de Tabagón (Galicia, Spain) was not found, so Wasowicz et al. (2017a) designated the illustration presented in the original description ( Merino 1898, Plate I) as lectotype. Here we have found specimens from La Guardia collected by Merino in 1898 (MA152390!, MA170055!), a locality also mentioned in the species description, and hence we consider them here as original material, however the term “ paralectotype ” is not usually used on Botany.

In Wasowicz et al. (2017), the basionym for this variety is said to be Blechnum spicant var. homophyllum Merino in Christ (1904: 79). However, Merino first described the species within a new genus, naming it Homophyllum blechniforme Merino (1898: 108). Later, Merino himself consulted with H. Christ on the status of the taxon and decided to combine it as a variety of S. spicant (sub. Blechnum spicant ), which he published acknowledging Merino (Christ 1904). However, we consider that Homophyllum blechniforme was a correctly published name, and there is no reason not to consider it the basionym of the taxon as it was the first one published.

Additional specimens examined:— PORTUGAL. Azores: San Miguel Island, Algarvia , 17 March 2007, J. A. Ramos & L. G. Quintanilla LGQ763 ( MA939454 !) ; Trilhos Novos, J. A. Ramos & L. G. Quintanilla LGQ764 16 March 2007, ( MA939453 !) . Madeira Island: ER228 road between Encumeada and S. Vicente , 02 May 1998, L. G. Quintanilla LGQ682 ( MA939455 !). Norte: Braga, Vieira do Minho , 1 October 2004, C. Prada s.n. ( MACB109621 !) ; Minho, Pinheiro, N103 Braga-Chaves rd. , 13 July 1974, E. A. Leadlay & B. Petty 392 ( MO2358910 !) ; Ponte do Lima, Anais, Fujaens , 9 September 1963, G. Costa s.n. ( MA633695 !) ; S. Gregorio do Minho , 14 July 1945, J. G. García 757 ( MA270375 !) . SPAIN. Euskadi: Álava, no date, no collector s.n. ( MA226678 !) . Castilla y León: Salamanca, Batuecas , 15 May 2016, J. M: Gabriel y Galán s.n. ( MACB109626 !) ; Segovia, Valle del Tejadilla , no date, E. Reyes Prósper s.n. ( MA001433 !) . Galicia: no date, M. Colmeiro s.n. ( MA232665 !) ; A Coruña, A Baña, San Mamede de Monte church, 13 August 2004, L. G. Quintanilla LGQ726 ( MA939457 !) ; A Coruña, Mariaqueira river , 21 February 2001, L. G. Quintanilla LGQ777 ( MA939456 !) ; A Coruña, Santiago, Cantaleta , 29 July 1967, I. Barrera s.n. ( MACB32367 !) ; Orense, Lovios, Parque Natural do Xurés río Caldo , 4 September 1997, I. Pulgar & L. G. Quintanilla s.n. ( MA595201 !) ; Pontevedra, Bueu, Ermelo , 11 October 1979, S. Castroviejo s.n. ( MA198344 !) ; Pontevedra , between Tabagón and Tomiño, 19 March 2016, J. M. Gabriel y Galán s.n. ( MACB109617 !) ; Pontevedra , La Guardia, B. P. Merino s.n. ( MA152390 !, paratype) ; ibidem ( MA170055 !, paratype) ; Pontevedra, Mondariz , 20 March 2016, J. M. Gabriel y Galán s.n. ( MACB109618 !) ; Pontevedra, Mointe Aloia, regato do Seixal , 31 August 1996, B. Pias & L. G. Quintanilla 23 ( MA591004 !) .

Struthiopteris spicant (L.) Weiss var. pradae S.Molino & Gabriel y Galán in Molino et al. (2019b: 266) ( Figure 9H ; Figure 14H ; Figure 18B ).

Type:— SPAIN. Zamora, Aciberos, c. 3km west of the village, 42.038942º, -6.861643º, on a shady slope near the road, under Fraxinus angustifolia and Quercus pyrenaica , 1,220 m, 20 September 2017, Molino, et al. s.n. ( MACB 110654!, holotype).

Plants terrestrial: rhizomes thick, short, covered with linear-lanceolate scales, 0.45‒1.6 x 4.5‒9.5 mm, discoloured, centre sclerotized from base to apex, dark brown, margin light brown, entire; fronds dimorphic, forming a rosette; sterile fronds (14) 34 (70) cm, stipes short, 0.5‒6 (18) cm, dark brown, black or purplish, laminae narrowly lanceolate to oblong lanceolate, 1.5‒6 x 13‒30 (56) cm, pinnatipartite or pinnatisect, apex pinnatifid, rarely conform, with uniseriate multicellular hairs ending in a red gland, rachis grooved on adaxial side, green or light brown towards apex, with glandular hairs like those on the lamina, pinnae tapering towards the base, the middle ones 0.2‒0.6 x 0.8‒3 cm, veins free, simple to 1-furcate, catadromous, ending in adaxial submarginal hydathodes, stomata aligned following veins on abaxial face of pinnae; fertile fronds of two types, both with glandular hairs like those on the sterile fronds, first type (25) 53 (90) cm, arising from the central part of the rosette, erect, stipes long, 4‒15 (28) cm, dark brown, black or purplish, red in growth, laminae lanceolate, 2‒6 x 20‒40 (63) cm, pinnatipartite or pinnatisect, pinnae more widely spaced and contracted, the middle ones 0.1‒0.3 x 2‒3 cm; sori linear, continuous, forming coenosori on both sides of costa, decurrent towards rachis; indusia continuous; second type of fertile fronds normally a single frond, same aspect as sterile ones but with sori in the middle part of the lamina, linear, fragmented on both sides of the costa; indusia linear, fragmented; sporangia with 13‒20 arc cells; spores monolete, 20‒40 x 30‒60 µm, perispores brown, rugate.

Habitat and distribution:— Known from Northern half of the Iberian Peninsula (provinces of Zamora, Asturias Burgos, León, and Madrid), Germany (States of Baden-Wurtemberg, Hesse, Niedersachsen, Nordrhein-Westfalen, Rhineland-Bergisch, Rhineland-Palatin and Schlesqig-Holstein) and sparce collections of Alsace ( France), Steiermark ( Austria) and Scotland ( Figure 2 ). Probably more abundant. Moist slopes.

Chromosome number:— Unknown.

Etymology:— This variety is dedicated to the reputed Spanish pteridologist Carmen Prada, who has devoted much of her life to the study of ferns, being a teacher, inspiration and mirror of good work for several generations of pteridologists and botany students.

Taxonomic notes:— The detection of several specimens of this variety in herbarium specimens from Germany led us to suspect that these phenotypic variations had been studied in the past. Indeed, we found two 19th century works describing several varieties within the species ( Luerssen 1889, Wilhelm & Laubenburg 1897). Among these varieties, the one described as variety complexa seems to be similar to variety pradae . In particular, intermediate fronds are described, with trophophyllous appearance but with fragmented sori in the middle zone of the frond. However, we have found no evidence in either work to confirm that in these specimens intermediate fronds coexist with normal dimorphic sporophylls, a feature that is fundamental according to the original description of S. spicant var. pradae ( Molino et al. 2019c) . Similarly, high importance is given in both Luerssen (1889) and Wilhelm & Laubenburg (1897) work to the relative length of the stipe, a trait that proved not to be significant in differentiating the different varieties (Molino et al. 2019bc).

Likewise, in the sheet review we have carried out, we have seen that the extent of the variety pradae is much greater than previously reported, as had already begun to be noted in other studies ( Molino et al. 2021). Struthiopteris spicant var. pradae is not, therefore, an Iberian endemism, but extends at least as far as Austria, France, Scotland, and Germany and could probably be even more widespread.

Additional specimens examined:— AUSTRIA. Steiermark: Waldbach , August 1904, F. Wirtgen 9440 ( P01465181 !) . FRANCE. Alsace: Hagenau, June 1834, Grenier s.n. ( P01566964 !; P01566965 !) . GERMANY. Baden-Wurtemberg: Baden-Baden , 5 October 1899, A. Kunz 6285 ( P01465178 !) . Hesse: 2 September 1899, F. Wirtgen 6283 ( P01465185 !) ; Taurus, Heidtränkskal , 29 August 1897, F. Wirtgen 6264 ( P01465204 !) ; ibidem, F. Wirtgen 6288 ( P01465179 !; P01465180 !) ; ibidem, September 1900, J. Müller-Knatz 6298 ( P01465193 !) ; ibidem, J. Müller-Knatz 6303 ( P01465195 !) . Niedersachsen: Oldenburg , 10 October 1904, illegible collector 9439 ( P01465203 !) . Nordrhein-Westfalen: 9 October 1900, Wirtgen 6295 ( P01465191 !) ; 5 July 1901, F. Wirgten 1849 ( P01465194 !) ; ibidem, F. Wirtgen 1850 ( P01465198 !) ; ibidem, F. Wirtgen 1852 ( P01465175 !) ; ibidem F. Wirtgen 1853 ( P01465176 !) ; 6 July 1901, F. Wirtgen 1854 ( P01465177 !) ; Bröl , 13 September 1899, F. Wirtgen 6297 ( P01465190 !) ; Rüdinghausen , 6 July 1906, F. Wirtgen 1855 ( P01465209 !) . Rhineland-Bergisch: Bergisch , 27 October 1897, F. Wirtgen 6286 ( P01465187 !) ; ibidem, 3 October 1898, F. Wirtgen 6287 ( P01465186 !) ; ibidem, 30 August 1900, F. Wirtgen 6282 ( P01465183 !; P01465184 !) ; ibidem ( P01465188 !) ; ibidem, 15 September 1900, F. Wirtgen 6299 ( P01465196 !) ; ibidem, September 1900, J. Müller-Knatz 6300 ( P01465197 !) . Rhineland-Palatinate: Gerolstein, 25 August 1899, Wessam [interpreted] 6284 ( P01465182 !) ; Krautscheid, 1901, F. Wirtgen s.n. ( P01465173 !) ; ibidem ( P01465174 !) . Schlesqig-Holstein: Kreis Pinneberg, September 1902 ‒1903, Schmidt 9329 ( P01465208 !) ; ibidem, September 1905, Misbanth 11716 ( P01465201 !). Unknown location: 9 November 1902, Alf. Hirth s.n. ( C0379948 F!) ; 1906, illegible collector 12773 ( P01465200 !) ; 1906, illegible collector 12777 ( P01465189 !) ; August 1906, illegible collector 12772 ( P01465201 !) ; ibidem, illegible collector 12779 ( P01465199 !) . SPAIN. Asturias: Valdés, Paladeperre , 22 March 2016, J. M. Gabriel y Galán s.n. ( MACB109613 !) ; Ibidem , 3 August 2016, J. M. Gabriel y Galán s.n. ( MACB109615 !) ; Luarca , 26 July 2017, J. M Gabriel y Galán s.n. ( MACB110660 ) ; Otur , 17 August 2017, J. M. Gabriel y Galán s.n. ( MACB110661 !) . Castilla y León: Burgos, Sierra de San Millán , 17 September 1975, Fuentes s.n. ( MACB5994 !) ; León, Hayedo de Busmayor , 19 June 2019, R. Vázquez s.n. ( MA939449 !) ; León, Pinar de Lillo , 20 June 2019, R. Vázquez s.n. ( MA939450 !) ; Salamanca , entre San Miguel de Valero y San Esteban de la Sierra, 15 May 2016, J. M. Gabriel y Galán & M. Puelles s.n. ( P00932168 !) . Euskadi: Álava , no date, no collector s.n. ( MA226678 !) . Madrid: Rascafría, trail near Pradillo dam, 24 February 2019, S. Molino et al. s.n. ( MA939451 !) . Navarra: Belagua, towards Lapatia pass, 26 August 2019, J. M. Gabriel y Galán & M. Puelles s.n. ( MA939452 !) . UNITED KINGDOM. Scotland: Loch Katrine, 26 June 1901, E. T. Harper & S. A. Harper s.n. ( C0379962 F!) .

Figures

FIGURE 2. Distribution of Struthiopteris spicant extracted from GBIF. Purple circles: Struhtiopteris spicant var. spicant; yellow squares: Stuthiopteris spicant var. pradae (added manually); blue pentagons: Struthiopteris spicant var. homophyllum (added manually).

FIGURE 9. Spores of Spicantopsis: A. S. amabilis (J. Faurié, 5612 P01406648). B. S. hancockii (R. Knapp 4311, P02435597). C. S. niponica var. niponica (M. Tagawa & K. Iwatsuki 3014, P01575937). Spores of Struthiopteris: D. S. castanea (Y. Saito s.n., TNS792168). E. S. fallax (P. Wasowicz & J.M. Gabriel y Galán s.n., MACB 109359); F. S. spicant var. spicant (A. Santos s.n., MACB36152). G. S. spicant var. homophyllum (Barrera s.n., MACB32367); H. S. spicant var. pradae (S. Molino et al s.n., MACB110654). Bar = 30 μm in A–D; 22 μm in E–H.

FIGURE 14. Struthiopteris species in the field. A-B. Struthiopteris castanea. A. Habit; B. Detail of the abaxial side of the fertile frond, bearing sori; C-D. Struthiopteris fallax. C. Habit; D. Detail of the abaxial side of the fertile frond, bearing sori; E-F. Struthiopteris spicant var. spicant. E. Habit; F. Detail of the adaxial side of the fertile frond; G. Struthiopteris spicant var. homophyllum, detail of the abaxial side of the fertile frond, bearing sori; H. Struthiopteris spicant var. pradae, detail of the abaxial side of the fertile frond, bearing sori. Pictures: A & B. Shirakawa, Gifu, Japan, by Hiroki Moriwaki; E & H. Zamora, Castilla y León, Spain, by Sonia Molino; F. Soria, Castilla y León, Spain, by Sonia Molino; G. La Coruña, Galicia, Spain, by Sonia Molino.

FIGURE 17. Illustration of Struthiopteris spicant var. spicant. A. Habit (Zollikofer s.n., P01658104); B. Sporangium (J.M. Gabriel y Galán s.n., MACB109622); C. Rhizome scale (S. Pajarón & E. Pangua s.n., MACB59142). Scale: 4 cm in A; 100 µm in B; 1 mm in C.

FIGURE 18. A. Illustration of Struthiopteris spicant var. homophyllum (J.M. Gabriel y Galán s.n., MACB109617); B. Illustration of Struthiopteris spicant var. pradae (S. Molino et al. s.n., MACB110654, holotype). Scale: 2 cm in A; 4 cm in B.