9. 1 Athyrium Roth (1799), Tent. Fl. Germ. 3: 31, 58. 1 Athyrium filix femina




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9  Woodsiaceae (Draft: N.N. Tzvelev)

S Athyriaceae

Comments:

(1) Woodsiaceae is here proposed kept apart from Dryopteridaceae. Woodsiaceae 1949 has priority before Athyriaceae 1956, according to Kramer in Kubitzki (1990). (Elven)



9.1  Athyrium Roth (1799), Tent. Fl. Germ. 3: 31, 58.
9.1.1  Athyrium filix femina (L.) Roth (1799), Tent. Fl. Germ. 3: 65.

B Polypodium filix femina L. (1753), Sp. Pl. 1090.

S

T Plukenet, Phytographia t. 180, f. 4 (1692) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 149). Typotype in Sloane Herbarium 85: 127 (BM).



Comments:

(1) This complex could also be treated as an aggregate of three species as we here have to decide between species, subspecies, or perhaps varieties as done by Kato (1993, Fl. N. Amer. 2). As they are at the same ploidy level and seem to represent major, nearly allopatric geographical races, subspecies would be the natural choice in our context. I have therefore chosen that solution as a proposal. Tzvelev proposes species. (Elven)


9.1.1.1  Athyrium filix-femina (L.) Roth subsp. filix-femina

S A. filix-femina (L.) Roth var. filix-femina.

2n= 80 (8x).

2nD Löve & Löve (1975) list several counts, one Icelandic; Jonsell (2000 Icel, Norw, Finl).

G ICE NOR RUS

(1) Greenland is omitted. The S Greenland occurrence is mapped as 'angustum' by Kato (1993). (Elven)


9.1.1.2  Athyrium filix femina (L.) Roth subsp. cyclosorum (Rupr.) C. Chr. in Hultén (1937), Fl. Aleut. Isl. 50.

B A. filix femina (L.) Roth var. (gamma) cyclosorum Rupr. (1845), Distr. Crypt. Vasc. Ross. 41.

S A. cyclosorum (Rupr.) Maxon (1900), Bull. Torrey Bot. Club 27: 637.

2n= 80 (8x).

2nD Löve & Löve (1975) list three non-arctic counts; Kato (1993).

G RFE ALA CAN

Comments:
9.1.1.3  Athyrium filix femina (L.) Roth subsp. angustum (Willd.) Clausen ***.

B Aspidium angustum Willd. (1810), Sp. Pl. 5,1: 277.

S A. angustum (Willd.) C. Presl ***; A. filix femina (L.) Roth var. angustum (Willd.) G. Lawson (1864), Edinb. New Philos. J., N.S. 19: 115;

2n= 80 (8x).

2nD Fl. N. Amer. 2 (1993).

G CAN GRL

Comments:
9.1.2  Athyrium distentifolium Tausch ex Opiz (1820), Tent. Fl. Crypt. Boem. 2, 1: 14.

S

T [Described from Czech Republic, Bohemia.]



Comments:

(1) I wonder about why Fl. N. Amer. 2 (1993) and some other sources accepts A. alpestre as valid name in favour of A. distentifolium. (Elven)

(2) Kato (1993) recognizes all American materials as a separate var. americanum. This should be included as a taxon as it is mapped for S Greenland. In our concept it would be a subspecies as it is a major geographical race. I have changed accordingly from Tzvelev's draft. However, Hultén & Fries (1986, map. 50) separate between var. distentifolium (Eurasia, Greenland), var. gaspense (NE America) and var. americanum (W America), the two latter do not reach the Arctic. (Elven)
9.1.2.1  Athyrium distentifolium Tausch ex Opiz subsp. distentifolium

S Athyrium alpestre (Hoppe) Rylands ex T. Moore (1856), non Clairv. (1811). [Basionym: Aspidium alpestre Hoppe (1805), Bot. Taschenb. 1805: 216.]

2n= 80 (8x).

2nD Löve & Löve (1975) list four counts, one Icelandic.

G ICE NOR RUS

Comments:


9.1.2.2  Athyrium distentifolium Tausch ex Opiz subsp. americanum (Butters) Lellinger ***

B A. alpestre (Hoppe) Clairv. var. americanum Butters (1917), Rhodora 19: 204.

S

2n= 80 (8x).



2nD Kato (1993).

G GRL


Comments:

9.2  Gymnocarpium Newman (1851), Phytologist 4: 371.

S Carpogymnia (H.P. Fuchs) Á.Löve & D.Löve (1966), nom. superfl., Univ. Colorado Stud. Ser. Biol. 24: 8.

Comments:

(1) The Amphi-Atlantic G. robertianum (Hoffm.) Newm. approaches the Arctic in the Ungava Bay area, according to Kato (1993, Fl. N. Amer. 2). (Elven)


9.2.1  Gymnocarpium dryopteris (L.) Newman (1851), Phytologist 4: 371.

B Polypodium dryopteris L. (1753), Sp. Pl. 1093.

S G. dryopteris (L.) Newman subsp. dryopteris; Currania dryopteris (L.) Wherry (1942), Bartonia 21: 15; Dryopteris linnaeana C. Chr. (1905), Index Filic. 275.

T Burser Herbarium XX:32 (UPS) lectotype, selected by McNeill & Pryer (1985, Taxon 34: 142).

2n= 160 (4x).

2nD Löve & Löve (1975) list several counts, one Icelandic; Pryer (1993, Fl. N. Amer. 2); Jonsell (2000 Finl).

G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:

(1) The diploid Gymnocarpium disjunctum is included in Tzvelev's concept of G. dryopteris. I have excluded it as Pryer (1993, Fl. N. Amer. 2) states that G. dryopteris is an allotetraploid from the diploids G. appalachianum Pryer & Haufler and G. disjunctum and therefore should be treated as a separate species. Gymnocarpium disjunctum does not seem to reach the Arctic, at least not in North America. If it reaches the Arctic in Asia, the relevant names are: Dryopteris disjuncta (Rupr.) C.V. Morton (1941), Rhodora 43: 217; G. dryopteris (L.) Newman subsp. disjunctum (Rupr.) Sarvela (1978), Ann. Bot. Fenn. 15: 103; G. disjunctum (Rupr.) Ching (1965), Acta Phytotax. Sin. 10: 304. Opinions? (Elven)

(2) Are the names Currania dryopteris and Dryopteris linnaeana in any current use? (Elven)


9.2.2  Gymnocarpium continentale (Petrov) Pojark. (1950), Soobshch. Tadzh. Fil. Akad. Nauk SSSR 22: 10.

B Dryopteris pulchella (Salisb.) Hayek var. continentale Petrov (1930), Fl. Iakut. 1: 14.

S G. jessoense (Koidz.) Koidz. [(1936), Acta Phytotax. Geobot. 5: 40 [basionym: Dryopteris jessoensis Koidz. (1924), Bot. Mag. (Tokyo) 38: 104] subsp. parvulum Sarvela (1978), Ann. Bot. Fenn. 15: 103; Dryopteris robertiana sensu Porsild & Cody (1980), non (Hoffm.) C. Chr. (1906); Gymnocarpium robertianum sensu Hultén (1968), non (Hoffm.) Newm. (1851).

T [Described from Siberia: Yakutian ASSR.] [Type of G. jessoense subsp. parvulum Canada: NWT, Distr. of Mackenzie, Nahanni National Park, below Virginia Falls, c. 0.3 km from the South Nahanni River, 6136'N, 12543'W, 08.07.1977, leg. T. Ahti 31910 (H) holotype.]

2n= 160 (4x).

2nD Löve et al. (1977); Jonsell (2000 N Finl).

G SIB RFE CAN

Comments:

(1) Sarvela in Jonsell (2000, Flora Nordica 1) are now of the opinion that the tetraploid subsp. parvulum deserves status as species distinct from the diploid (2n=80) G. jessoense s. str., following Pryer et al. (1983, Canad. J. Bot. 61: 2592-2602). Pryer (1993) treats it as a subspecies. It is in accordance with prevailing principles in pteridophyte taxonomy to recognize different ploidy levels as species, as the tetraploids and higher levels usually include several different genomes. (Elven)

(2) Tzvelev includes, as a synonym, G. remote-pinnatum (Hayata) Ching (1935), Bull. Chin. Bot. Soc. 1,2: XIV. This is not obvious as this taxon is treated as a separate species by Sarvela (1978), and as it is considered a narrow endemic of Taiwan and should not appear in any of our standard reference sources. (Elven)


9.2.3  Gymnocarpium x intermedium Sarvela (1978), Ann. Bot. Fenn. 15: 105.

S G. continentale x dryopteris; G. dryopteris x jessoense subsp. parvulum.

T Finland: Utsjoki, northern part of Kevojoki Valley, Puksalskaidi, S. part of Kotkapahta, 16.08.1969, leg. U. Laine (TUR) holotype.

2n= c. 160 (4x).

2nD Jonsell (2000 N Finl, 2 provinces).

G RFE


Comments:

(1) Accepted because it often appears independent and outside the range of at least one of its parents and forms large stands (probably by vegetative means). (Elven)



9.3  Cystopteris Bernh. (1805), nom. cons., Neues J. Bot. 1, 2: 26.
9.3.1  Cystopteris fragilis (L.) Bernh. (1805), Neues J. Bot. 1, 2: 27.

B Polypodium fragile L. (1753), Sp. Pl. 1091.

S C. fragilis (L.) Bernh. subsp. fragilis; C. fragilis (L.) Bernh. var. fragilis; C. filix fragilis (L.) Borbás (1900), Balaton Fl. 314; C. dickieana R. Sim. (1848), Gard. Farmers' J. 2,20: 308; C. fragilis (L.) Bernh. var. dickieana (R. Sim) T. Moore (1848), Handb. Brit. Ferns 81; C. fragilis (L.) Bernh. subsp. dickieana (R. Sim) Hook. fil. (1870), Student Fl. Brit. Isl. 464.

T Plukenet, Almagestum botanicum, tab. 180, fig. 5 (1696) lectotype, selected by Weatherby (1935, Rhodora 37: 376).

2n= (1) 168 (4x). (2) 252 (6x). (3) 336 (8x).

2nD (1) Hämet Ahti et al. (1984   Finland, given for the collective species); Löve & Löve (1975) list numerous counts for C. fragilis s. str., one Icelandic, four for C, dickieana, one as arctic; Haufler et al. (1993, Fl. N. Amer. 2). (2) Vida (1977 C Eur). (3) Vida (1977 C Eur).

G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:

(1) There are good morphological, genetical and ecological reasons for separation of C. fragilis and C. alpina. The higher ploidy levels reported by Vida (1977) might well refer to the latter. (Elven)

(2) The only consistent feature that separates 'dickieana' from 'fragilis' s. str. is the spore ornamentation. In the N Atlantic area, there is no connection between this ornamentation and other features and it is impossible to separate 'dickieana' macromorphologically. It also occurs more or less occasional within populations (see e.g. Berg 1992, Blyttia 50: 143-157). Cystopteris dickieana has been finally 'killed' as a taxon by Dyer et al. (2000, Edinb. J. Bot. 57: 71-81) and Parks et al. (2000, Edinb. J. Bot. 57: 83-105).

Even if the reticulate spore ornamentation has a certain geographic pattern, this is to be considered as a geographical variation in one character, not associated with other characters, and not nearly enough for recognition of a taxon. Vida's proposal (1974) that partly different diploid genomes partake in 'fragilis' and 'dickieana' has not found further support and is countered by Parks et al. (2000). My proposal is to only include it as a comment, and in the synonymy of C. fragilis. (Elven)
9.3.2  Cystopteris alpina (Lam.) Desv. (1827), Mém. Soc. Linn. Paris 6: 264.

B Polypodium alpinum Lam. (1779), Fl. Franc. 1: 22.

S C. regia auct., non (L.) Desv. (1827).

T [Described from the Alps.]

2n= 256 (6x).

2nD Vida (1977 C Eur); Löve et al. (1977).

G NOR

Comments:



(1) The type of Linnaeus Polypodium regium (1753, Sp. Pl. 1091) belongs, according to the Linnaean typification programme of Nordic plants (Jonsell & Jarvis), within C. fragilis s. str. The valid name of this fairly distinct taxon is therefore most probably C. alpina. It reaches the Arctic at least in NE Norway (Varanger Peninsula). (Elven)
9.3.3  Cystopteris montana (Lam.) Desv. (1827), Mém. Soc. Linn. Paris 6: 264.

B Polypodium montanum Lam. (1779), Fl. Franc. 1: 23.

S Rhyzomatopteris montana (Lam.) Khokhr. ***

T [Described from France.]

2n= 168 (4x).

2nD Löve & Löve (1975) list four non-arctic counts; Fl. N. Amer. 2 (1993).

G NOR RUS RFE ALA CAN GRL

Comments:

(1) The separation of this species as a Rhyzomatopteris ***, as proposed in Tzvelev's draft, should be justified. The genus is not accepted in recent surveys of genera of ferns (see, e.g., Kubitzki 1990). I have therefore tentatively retained the species inside Cystopteris but am very willing to reconsider when a justification is presented (and information on when and where Rhyzomatopteris is published). (Elven)

9.4  Woodsia R. Br. (1810), Prodr. 158.
9.4.1  Woodsia ilvensis (L.) R. Br. (1815), Trans. Linn. Soc. London 11: 173.

B Acrostichum ilvense L. (1753), Sp. Pl. 1071.

S

T Linnaean Herbarium 1245.13 (LINN) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 149).



2n= (1) 82 (2x). (2) 80 82 (2x).

2nD (1 2) Löve & Löve (1975) list several counts, one Icelandic; (1) Windham (1993, Fl. N. Amer. 2); Hämet Ahti et al. (1984   Finl); Jonsell (2000 Sweden).

G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:


9.4.2  Woodsia alpina (Bolton) Gray (1821), Nat. Arr. Brit. Pl. 2: 17.

B Acrostichum alpinum Bolton (1790), Fil. Brit. 76.

S

T [Described from Britain: "mountains of Wales".]



2n= (1) 156. (2) 164 (4x).

2nD (1) Jonsell (2000 Icel). (2) Löve & Löve (1975) list two counts, one Icelandic; Windham (1993); Jonsell (2000 Icel, 2n= c. 160 Sweden).

G RUS SIB RFE ALA CAN GRL

Comments:


9.4.3  Woodsia glabella R. Br. ex Richardson in Franklin (1823), Narr. Journey Polar Sea (Bot. App.) 754.

S W. glabella R. Br. ex Richardson subsp. glabella.

T [Described from Canada: Mackenzie District.]

2n= (1) 78 (2x). (2) c. 80 (2x).

2nD (1 2) Löve & Löve (1975) list several counts, two as arctic; (1) Windham (1993): Hämet Ahti et al. (1984 Finl).

G NOR RUS SIB RFE ALA CAN GRL

Comments:

10  Dryopteridaceae (Draft: N.N. Tzvelev)

S Aspidiaceae



10.1  Dryopteris Adans. (1763), nom. cons., Fam. Pl. 2: 20, 551.

Comments:

(1) Order of species has been changed from Tzvelev's draft to keep relatives closer together. One species is added, D. marginalis. With only five species, the three sections of Tzvelev's draft are omitted: sect. Dryopteris (D. filix-mas, D. marginalis), sect. Fragrantes (H. Ito) Tzvelev (D. fragrans) and sect. Lophodium (Newm.) C. Chr. (D. expansa, D. carthusiana). (Elven)
10.1.1  Dryopteris fragrans (L.) Schott (1834), Gen. Fil. 1: t. 9.

B Polypodium fragrans (1753), Sp. Pl. 1089.

S

T Linnaean Herbarium 1251.32 (LINN) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 15: 149).



2n= 82 (2x).

2nD Löve & Löve (1975) list four non-arctic counts; Montgomery & Wagner (1993, Fl. N. Amer. 2).

G RUS SIB RFE ALA CAN GRL

Comments:


10.1.2  Dryopteris filix mas (L.) Schott (1834), Gen. Fil. 1: t. 9.

B Polypodium filix mas L. (1753), Sp. Pl. 1090.

S D. abbreviata sensu Jalas & Suominen (1972) and Löve & Löve (1975), non (DC.) Newman ex Manton (1950).

T Clifford Herbarium 475, Polypodium 10 (BM) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 15: 149).

2n= 164 (4x).

2nD Löve & Löve (1975) list numerous counts, one as arctic; Montgomery & Wagner (1993); Jonsell (2000 Icel, Finl).

G ICE NOR RUS GRL

Comments:


10.1.3  Dryopteris marginalis (L.) A. Gray (1848), Manual 632.

B Polypodium marginale L. (1753), Sp. Pl. 1091.

S D. abbreviata sensu Böcher et al. (1978).

2n= 82 (2x).

2nD Montgomery & Wagner (1993).

G GRL


Comments:

(1) This entity seems to be the base of repeated reports of D. 'abbreviata' from Greenland. It is restricted to the southernmost parts of Greenland and is a border case. (Elven)


The Dryopteris dilatata aggregate (D. carthusiana, D. expansa)

Comments:

(1) The need for this aggregate disappears if D. carthusiana is omitted. (Elven)
10.1.4  Dryopteris expansa (C. Presl) Fraser Jenk. & Jermy (1977), Brit. Fern Gaz. 11: 338.

B Nephrodium expansum C. Presl (1825), Reliq. Haenk. 1: 38.

S Dryopteris assimilis S. Walker (1961), Amer. J. Bot. 48: 607; Dryopteris austriaca auct., non Woynar ex Schinz & Thell. (1915); Dryopteris dilatata auct., non (Hoffm.) Gray (1848); D. dilatata (Hoffm.) Gray subsp. americana (Fisch.) Hultén (1968), Ark. Bot., ser. 2, 7, 1: 4.

T [Described from North America.]

2n= 82 (2x).

2nD Löve & Löve (1975) list numerous counts, one as arctic; Montgomery & Wagner (1993); Jonsell (2000 Norw, Finl, Sweden).

G ICE NOR RUS SIB? RFE ALA CAN GRL

Comments: See D. carthusiana.


10.1.5  Dryopteris carthusiana (Vill.) H.P. Fuchs (1959), Bull. Soc. Bot. France 105: 339.

B Polypodium carthusianum Vill. (1786), Hist. Pl. Dauphiné 1: 292.

S Dryopteris spinulosa O. Kuntze (1891), Revis. Gen. Pl. 2: 813.

T France: Isère, 1978, leg. C.R. Fraser-Jenkins 7335 (BM) neotype, selected by Fraser-Jenkins (1980, Taxon 29: 610).

2n= 164 (4x).

2nD Löve & Löve (1975) list numerous non-arctic counts; Montgomery & Fraser (1993); Jonsell (2000 Finl, Sweden).

G RUS? SIB?

Comments:

(1) Presence of this species in the Arctic needs confirmation. Russian sources map this species far north in Kola Peninsula and further east. On the Finnish and Norwegian sides of the border it is a southern species, rare already in the middle boreal zone. A superficial examination of specimens in LE showed that what went by the name D. carthusiana from the northern parts of European Russia mainly confirmed with D. expansa as this plant is understood on the Finnish/Norwegian side. The few exceptions seemed to be from non-arctic areas. There is an obvious difference in criteria used for identifying the two species in NW Europe and Russia, and others have also commented on the different Russian concept of D. carthusiana (e.g., Jalas & Suominen in Atlas florae europaeae 1). (Elven)

WARNING! Will be excluded if occurrences in arctic Russia not confirmed.

10.2  Polystichum Roth (1799), Tent. Fl. Germ. 3, 1: 31, 69.
10.2.1  Polystichum lonchitis (L.) Roth (1799), Tent. Fl. Germ. 3, 1: 71.

B Polypodium lonchitis L. (1753), Sp. Pl. 1088.

S

T Clifford Herbarium 475, Polypodium 5 (BM) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 149).



2n= 82 (2x).

2nD Löve & Löve (1975) list several counts, one Icelandic; Wagner (1993, Fl. N. Amer. 2).

G ICE NOR RUS CAN GRL

Comments:



11  Polypodiaceae (Draft: N.N. Tzvelev)

11.1  Polypodium L. (1753), Sp. Pl. 1082.
11.1.1  Polypodium sibiricum Sipliv. (1974), Nov. Sist. Vyssh. Rast. 11: 329.

S P. virginianum auct., non L. (1753); P. vulgare L. subsp. virginianum sensu Hultén (***); P. vulgare L. subsp. columbianum (Gilb.) Hultén (1962), Circumpolar Pl. 1: 176; P. hesperium auct., non Maxon (1900); P. vulgare sensu Böcher et al. (1978).

2n= 74 (2x).

2nD Haufler et al. (1993, Fl. N. Amer. 2).

G ALA CAN GRL

Comments:

(1) There is perhaps some need for comments on the confusing concept and synonymy of this species. (Elven)
11.1.2  Polypodium vulgare L. (1753), Sp. Pl. 1085.

S

T Burser Herbarium XX:44 (UPS) lectotype, selected by Jonsell & Jarvis (1993, Regnum Veg. 127: 78).



2n= 148 (4x).

2nD Löve & Löve (1975) list numerous counts, two as arctic; Haufler et al. (1993); Hämet Ahti et al. (1984   Finland).

G ICE NOR RUS

Comments:


12  Blechnaceae (Draft: R. Elven)

12.1  Blechnum L. (1753), Sp. Pl. 1077.
12.1.1  Blechnum spicant (L.) Roth (1794), Ann. Bot. (Usteri) 10: 56.

B Osmunda spicant L. (1753), Sp. Pl. 1066.

S

T Sweden (Skåne), Linnaean Herbarium 1244.14 (LINN) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 150).



2n= 68 (4x).

2nD Löve & Löve (1975) list several counts, some Icelandic; Nauman (1993, Fl. N. Amer. 2).



G ICE

Comments:



(1) Added to Tzvelev's draft as it occurs in arctic parts of Iceland. (Elven)


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