1 Lycopodiaceae




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1  Lycopodiaceae (Draft: N.N. Tzvelev)

S Huperziaceae (incl.)

Comments:

(1) Tzvelev's proposal to separate Huperzia in its own family   Huperziaceae   is not followed here. Opinions? (Elven)



1.1  Huperzia Bernh. (1801), J. Bot. (Schrader) 1800, 2: 126.

S Lycopodium L. (1753) p. p.


1.1.1  Huperzia selago (L.) Bernh. ex Schrank. & Mart. (1829), Hort. Reg. Monac. 3.

B Lycopodium selago L. (1753), Sp. Pl. 1102.

S ?H. haleakalae (Brackenridge) Holub (1985), Folia Geobot. Phytotax. 20: 73 [basionym: Lycopodium haleakalae Brackenridge in Wilkes (1854), U.S. Expl. Exped. 16: 321].

T Burser Herbarium XX:52 (UPS) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 147).

Comments:

(1) Tzvelev's proposal is to recognise three partly zonal subspecies within a widely defined species. This is more or less in accordance with previous European -Russian treatments (e.g., Flora Europaea, Fl. Arct. USSR). These differ only by the number of subspecies. West European authors tend to regard subsp. appressa as intermediates between subsp. selago and subsp. arctica without its own rank, see e.g. Kukkonen in Jonsell (2000, Fl. Nord. 1), and most of them include subsp. appressa with subsp. arctica. Russian authors accept them as three separate subspecies.

The current North American treatment (Wagner Jr. & Beitel 1993, Fl. N. Amer. 2) differs in principle. The authors treat all lycopods as species and hybrids, without applying subspecific categories. An argument for this is presence of aborting spores in putative hybrids, putative from other morphological evidence. This approach is very problematic to apply in the Arctic. The authors place the temperate 'middle' North American material north to the south shore of Hudson Bay and perhaps NW parts of mainland NWT in H. selago s. str., the western material incl. Alaska and Yukon Territory in H. haleakalae (described from Hawaii), and the eastern material incl. Labrador and S Greenland in H. appalachiana. The majority of the American arctic material, from the Canadian Arctic Archipelago and C-N Greenland, is completely absent from their treatment (and maps). In their approach, this material would probably represent a fourth species.

In the checklist draft, Tzvelev lists H. haleakalae as a synonym of H. selago subsp. arctica, H. appalachiana as a synonym of H. selago subsp. appressa. The latter proposal is probably right as the S Greenland material mapped as H. appalachiana by Wagner & Beitel mostly corresponds with Tzvelev's subsp. appressa. The former synonymisation is much more dubious. Presence of this otherwise distinctly arctic race in Hawaii seems very improbable. I have therefore included H. haleakalae as a synonym of the species sensu lato.

In the checklist, we have to choose between the two alternative approaches as they are mutually exclusive. The North American treatment is an unsatisfactory choice for two reasons: (1) it neglects most of the American arctic plants, and (2) it is at present not applicable outside North America. In case of choice of the European -Russian approach, we have to choose between recognition of two or three infraspecific taxa. If subsp. appressa is seen as intermediate, this will also involve the principle concerning how to handle intermediates. (Elven)
1.1.1.1  Huperzia selago (L.) Bernh. ex Schrank & Mart. subsp. selago

S Lycopodium selago L. subsp. selago.

2n= (1) c. 90. (2) 264. (3) c. 272.

2nD (1) Sorsa (1963   Finl). (2) Löve & Löve (1975) list four counts, two as arctic; Fl. N. Amer. 2 (1993). (3) Löve & Löve (1961a Icel, for the collective species).

G RUS CAN GRL

Comments:

(1) Greenland is added because there is a specimen clearly belonging to this subspecies from southernmost Greenland deposited in HbO. (Elven)
1.1.1.2  Huperzia selago (L.) Bernh. ex Schrank & Mart. subsp. appressa (Bach. Pyl. ex Desv.) *** [not D. Löve in Á. Löve & D. Löve (1961), Bot. Not. 114].

B Lycopodium selago L. f. appressum Bach. Pyl. ex Desv. (1827), Ann. Soc. Linn. Paris 6: 180.

S Lycopodium selago L. subsp. appressum (Bach. Pyl. ex Desv.) Hultén (1968), Ark. Bot., ser. 2, 7, 1: 7; ?H. appalachiana Beitel & Mickel (1992), Amer. Fern J. 82: 45.

T [Described from Greenland and Newfoundland.]

2n=

2nD


G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:

(1) This subspecies might perhaps better be treated as part of an extended subsp. arctica or as intermediates without rank. I am in favour of the former solution. I have been through a large material from Greenland. There is much difference there between the northernmost and southernmost plants but no discontinuity. The discontinuity towards subsp. selago is, however, profound in Greenland as elsewhere. (Elven)
1.1.1.3  Huperzia selago (L.) Bernh. ex Schrank & Mart. subsp. arctica (Grossh. ex Tolm.) Á. Löve & D. Löve (1961), Bot. Not. 114: 35.

B Lycopodium selago L. subsp. arcticum Grossh. ex Tolm. (1960), Bot. Mater. Gerb. Bot. Inst. AN SSSR 20: 39.

S

T [Described from Siberia, Yakutian ASSR: "inter colles ad pagum Pochodskoje, prope fl. Kolyma inferiorem, in Jacutia arctica".]



2n=

2nD


G NOR RUS SIB RFE ALA CAN GRL

Comments:


1.1.2  Huperzia miyoshiana (Makino) Ching (1981), Acta Bot. Yunnan. 3: 303, 304.

B Lycopodium miyoshianum Makino (1898), Bot. Mag. (Tokyo) 12: 36.

S Lycopodium selago L. subsp. miyoshianum (Makino) Calder & R.L. Taylor (1965), Canad. J. Bot. 43: 1388; Lycopodium selago L. subsp. chinense (C. Chr.) Hultén *** [basionym: Lycopodium chinense C. Chr. (1897), Nuov. Giorn. Bot. Ital., n. s., 4: 101]; Huperzia selago (L.) Bernh. ex Schrank. & Mart. subsp. chinensis (C. Chr.) Á. Löve & D. Löve (1965), Univ. Colorado Stud., Ser. Biol., 17: 5.

2n=


2nD

G ALA


Comments:

(1) Should this be accepted as species or subspecies? Acceptance at rank of species is mainly according to the North American principles cited in comment under H. selago. The name is the one applied by Wagner & Beitel (1993). (Elven)



1.2  Lycopodium L. (1753), Sp. Pl. 1100.

Comments:

(1) Lycopodium is here proposed in a more restricted sense than done by, e.g., Øllgaard in Kubitzki (1990), excluding Diphasiastrum. (Elven)
1.2.1  Lycopodium annotinum L. (1753), Sp. Pl. 1103.

S

T Plukenet, Phytographia t. 205, f. 5 (1692) lectotype, selected by Jonsell & Jarvis (1994, Nordic J. Bot. 14: 147).



Comments:

(1) The treatment of subspecies should be discussed in connection with both L. annotinum and L. clavatum; species treatment is an alternative. The North American standpoint (Wagner & Beitel 1993) is to neglect the variation in L. annotinum but to accept the variation in L. clavatum at level of two species. In European  Russian views, these pairs of species/subspecies are comparable. The differences are mainly (or entirely?) quantitative in both cases. My personal preference is treatment as subspecies as both cases are pairs of closely related taxa in S/N direction, and as intermediates seem to occur even if at least the races of L. annotinum fairly often are found together and distinct. (Elven)


1.2.1.1  Lycopodium annotinum L. subsp. annotinum

S L. annotinum L. var. annotinum.

2n= 68 (4x).

2nD Manton (1950 N & W Eur); Löve & Löve (1975) list several non-arctic counts; Fl. N. Amer. 2 (1993).

G RUS SIB RFE ALA CAN GRL

Comments:

(1) Greenland is added to Tzvelev's draft. There are specimens typical of this subspecies in HbO from SE (Akorninarmiut), S (Igaliko) and W (Disko). (Elven)
1.2.1.2  Lycopodium annotinum L. subsp. alpestre (Hartm.) Á. Löve & D. Löve (1958), Nucleus (Calcutta) 1, 1: 7.

B L. annotinum L. var. alpestre Hartm. (1832), Handb. Skand. Fl., ed. 2, 294.

S L. dubium auct., non Zoëga (1772); L. pungens Bach.Pyl. (1827), Mém. Soc. Linn. Paris 6: 182; L. annotinum L. var. pungens (Bach.Pyl.) Desv. ***; L. annotinum L. subsp. pungens (Bach.Pyl.) Hultén (1968), Ark. Bot., sr. 2, 7, 1: 7.

T Sweden, "Jämtl. et Lapp." herb. Hartman (UPS) lectotype, selected by Kukkonen in Jonsell (1996, Nordic J. Bot. 16: 3).

2n= (1) 66 (4x). (2) 68 (4x).

2nD (1) Harmsen in Löve & Löve (1948). (2) Löve & Löve (1975) list two counts, one as arctic.

G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:

(1) Is the name L. dubium used in standard sources? (Elven)

(2) As to Iceland, in HbO we have material from N Iceland of clear subsp. alpestre and of intermediates but not of subsp. annotinum. (Elven)


1.2.2  Lycopodium clavatum L. (1753), Sp. Pl. 1101.

S

T Burser Herbarium XX:49 (UPS) lectotype, selected by Jonsell & Jarvis (1993, Regnum Veg. 127: 63).



Comments: See L. annotinum.
1.2.2.1  Lycopodium clavatum L. subsp. clavatum

S

2n= 68 (4x).



2nD Manton (1950 W Eur); Löve & Löve (1975) list several counts, one as arctic; Fl. N. Amer. 2 (1993).

G RUS SIB ALA CAN

Comments:
1.2.2.2  Lycopodium clavatum L. subsp. monostachyon (Grev. & Hook.) Selander (1950), Acta Phytogeogr. Suec. 28: 7.

B L. clavatum L. var. monostachyon Grev. & Hook. in Hook. (1831), Bot. Misc. 2: 375.

S L. clavatum L. var. lagopus Laest. ex C. Hartm. (1858), Handb. Skand. Fl., ed. 7: 313; L. lagopus (Laest. ex C. Hartm.) G. Zinserl. ex Kuzen. (1953), Fl. Murm. Obl. 1: 80.

T [Described from Canada: "the Rocky mountains, north of Smoking River in lat. 56.]

2n= 68 (4x).

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

G NOR RUS SIB RFE ALA CAN GRL

Comments:

(1) Acta Phytogeogr. Suec. 28, p. 22, erroneously cited in some sources. (Elven)
1.2.3  Lycopodium dendroideum Michx. (1803), Fl. Bor. Amer. 2: 282.

S L. obscurum L. var. dendroideum (Michx.) D.C. Eaton in A. Gray (1890), Manual, ed. 6, 696.

2n= 68 (4x).

2nD ?


G ALA CAN

Comments:



1.3  Diphasiastrum Holub (1975), Preslia 47: 104.

S Lycopodium L. (1753) p. p.; Diphasium auct., non C. Presl. ex Rothm. (1944).


1.3.1  Diphasiastrum complanatum (L.) Holub (1975), Preslia 47: 108.

B Lycopodium complanatum L. (1753), Sp. Pl. 1104.

S Diphasium complanatum (L.) Rothm. (1944), Feddes Repert. 54: 64.

T [Type not selected] or [European specimens have been selected as the type of this species (Wilce 1961, Nova Hedwigia 19)]

Comments:

(1) Kukkonen in Jonsell (2000, Fl. Nord. 1) recognizes D. complanatum with two subspecies, complanatum and montellii, but accepts D. tristachyum as species. The reason is the occurrence of intermediates with its own distribution between D. complanatum (subsp. complanatum) and D. tristachyum (named D. x zeilleri (Rouy) Holub). I find this rather a good argument for inclusion of D. tristachyum as a subspecies in an extended D. companatum. The problem is, however, of less relevance for the checklist as the Russian occurrences of D. tristachyum indicated by Tzvelev and others certainly belong within D. complanatum subsp. montellii (see, e.g., Jalas & Suominen 1972 and Hultén & Fries 1986). (Elven)


1.3.1.1  Diphasiastrum complanatum (L.) Holub subsp. complanatum

S Diphasium complanatum (L.) Rothm. subsp. complanatum.

2n= (1) 46 (2x). (2) 44 48 (2x). (3) c. 48 (2x).

2nD (1 3) Löve & Löve (1975) list several counts. (1) Fl. N. Amer. 2 (1993). (2) Hämet Ahti et al. (1984   Finland).

G NOR RUS SIB ALA CAN GRL?

Comments:

(1) Greenland with a question mark because of doubts. Are there two or three Greenland taxa of the group, and which? See Böcher et al. (1978) and Fl. N. Amer. 2. (Elven)
1.3.1.2  Diphasiastrum complanatum (L.) Holub subsp. montellii (Kukkonen) Kukkonen in Hämet Ahti & Kukkonen (1984), Ann. Bot. Fenn. 21: 210.

B Diphasium complanatum (L.) Rothm. subsp. montellii Kukkonen (1970), Ann. Bot. Fenn. 7: 142.

S Lycopodium tristachyum auct. ross., non Pursh (1814); Diphasium tristachyum auct. ross. et Löve & Löve (1975), non (Pursh) Rothm. (1944).

T Finland: Kittilän Lappi, Muonio, 06.09.1915, J. Montell (H) holotype.

2n= c. 48 (2x).

2nD Hämet Ahti et al. (1984   Finland).

G NOR RUS ALA CAN? GRL?

Comments:

(1) According to Hultén & Fries (1986) and Jalas & Suominen (1972), the N Russian occurrences considered as D. tristachyum are to be included here. The reports in Tzvelev's draft from Canada and ?Greenland might also refer to this subspecies. Material that I collected in Seward Peninsula (W Alaska) in 1998 falls very well within subsp. montellii.

If D. tristachyum, in spite of this, is to be retained, the data for it is: Diphasiastrum tristachyum (Pursh) Holub (1975), Preslia 47: 108. Basionym: Lycopodium tristachyum Pursh (1814), Fl. Amer. Sept. 2: 653 [or: 635?]. Synonyms: Diphasiastrum complanatum (L.) Holub subsp. chamaecyparissus (A. Braun ex Mutel) Kukkonen in Hämet Ahti & Kukkonen (1984), Ann. Bot. Fenn. 21: 210; Lycopodium chamaecyparissus A. Braun ex Mutel (1837), Fl. France 4: 192 [to be included as basionym if subspecific rank is chosen]; Diphasium complanatum (L.) Rothm. subsp. chamaecyparissus (A. Braun ex Mutel) Kukkonen [possibly (1970), Ann. Bot. Fenn. 7: ca 142]; Diphasium tristachyum (Pursh) Rothm. (1944), Feddes Repert. 54: 65. Described from USA, Virginia. 2n=46 (2x); Löve & Löve (1975) list two non-arctic counts. Reported from RUS. (Elven)


1.3.2  Diphasiastrum sitchense (Rupr.) Holub (1975), Preslia 47: 108.

B Lycopodium sitchense Rupr. (1845), Beitr. Pflanzenk. Russ. Reiches 3: 30.

S Lycopodium sabinaefolium Willd. var. sitchense (Rupr.) Fern. (1923), Rhodora 25: 166; Lycopodium sabinaefolium Willd. subsp. sitchense (Rupr.) Calder & R.L. Taylor (1965), Canad. J. Bot. 43: 1387.

2n= 46 (2x).

2nD ?

G CAN GRL?



Comments:

(1) The tentative report of this species from Greenland in Tzvelev's draft needs some justification. It is probably based on a 'dot' in the map in Fl. N. Amer. 2. (Elven)


1.3.3  Diphasiastrum alpinum (L.) Holub (1975), Preslia 47: 107.

B Lycopodium alpinum L. (1753), Sp. Pl. 1104.

S Diphasium alpinum (L.) Rothm. (1944), Feddes Repert. 54: 65.

T Linnaean Herbarium 1257.19 (LINN) lectotype, selected by Wilce (1965, as "1275.19", Beih. Nova Hedwigia 19: 32).

2n= (1) 46 (2x). (2) 44 48 (2x). (3) 48-50.

2nD (1 2) Löve & Löve (1975) list several counts. (1) Fl. N. Amer. 2 (1993); Jonsell (2000 Sweden). (2) Hämet Ahti et al. (1984   Finland). (3) Manton (1950 C Eur).

G ICE NOR RUS SIB RFE ALA CAN GRL

Comments:



(1) There is a rumour that this species also is found on Jan Mayen. A specimen might be in C. To be checked. (Elven)


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