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LYCOPODIACEAE (Clubmoss Family)

References: Lellinger (1985); Mickel (1979); Wagner and Beitel (1992); Beitel (1979); Snyder & Bruce (1986); Wagner & Beitel in FNA (1993b).

A family of 10-15 genera and about 400 species.

1. Leafy stems erect, simple or dichotomously branched, the ultimate branches vertically oriented; sporophylls like the sterile leaves or only slightly reduced, in annual bands along the stem; vegetative reproduction by leafy gemmae near stem apex; [subfamily Huperzioideae]......Huperzia

1. Leafy stems prostrate or erect, if erect then generally branched, the ultimate branches spreading (horizontal) or ascending; sporophylls differing from sterile leaves, either broader and shorter, or more spreading, aggregated into terminal cones; lacking vegetative reproduction by gemmae; [subfamily Lycopodioideae]......2

3. Shoots flat-branched, 1-3 mm wide (including the leaves); leaves scalelike, dimorphic, overlapping and imbricate, appressed to the stem, in 4 ranks; strobili on long, narrow peduncles......Diphasiastrum

3. Shoots round-branched, usually 5-10 mm wide (including the leaves), leaves awl-shaped, monomorphic (though sometimes differing in size), separate, spreading or ascending, in 6 ranks; strobili sessile at stem apices......Lycopodium

5. Leaves of the prostrate stems 0.5-1.2 mm wide, ciliate-toothed or not toothed; leaves of the erect stem many, overlapping, spiral; leaves of the strobilus (sporophylls) resembling leaves of the prostrate and upright stems in size and shape; upright stems 1.5-15 mm in diameter (including the leaves) ......Lycopodiella

5. Leaves of the prostrate stems 1.3-2.1 mm wide, not toothed; leaves of the erect stem few, not overlapping, whorled; leaves of the strobilus (sporophylls) much reduced relative to leaves of the prostrate and upright stems; upright stems 1.5-3 mm in diameter (including the leaves) ......Pseudolycopodiella caroliniana

Diphasiastrum Holub (Flat-branched Clubmoss, Running Cedar)

A genus of about 15-20 species, mostly north temperate amd subarctic.

1. Foliage dark green, not glaucous; horizontal branchlets 2-4 mm wide (including the leaves); branchlets without conspicuous annual constrictions; rhizomes 0-1 cm deep (which can be determined by pulling up a single upright shoot -- the depth to rhizome is approximately the length of the whitwe portion of the vertical stem); abaxial rank of leaves shorter than lateral ranks (thus the branchlets flat in cross-section)......D. digitatum

1. Foliage blue-green, glaucous; horizontal branchlets 1-2 mm wide (including the leaves); branchlets with conspicuous annual constrictions; rhizomes (1-) 5-12 cm deep; abaxial rank of leaves as long as lateral ranks (thus the branchlets more-or-less square in cross-section)......D. tristachyum

Diphasiastrum digitatum (Dillenius ex A. Braun) Holub, Common Running Cedar, Fan Ground- pine. Mt, Pd, Cp (NC, SC, VA): dry to mesic forests and openings, especially common in disturbed sites, such as successional pine forests; common. July-September. Widespread in e. North America. Hickey & Beitel (1979) and Holub (1975a & 1975b) explain the nomenclatural decision to accept the epithet 'digitatum' over the more familiar 'flabelliforme.' [= FNA; Lycopodium flabelliforme (Fernald) Blanch -- RAB, S; Lycopodium digitatum Dillenius ex A. Braun -- C, K, W; Lycopodium complanatum Linnaeus var. flabelliforme Fernald -- F, G]

Diphasiastrum tristachyum (Pursh) Holub, Blue Ground-cedar. Mt (NC, SC, VA), Pd (NC, VA), Cp (VA): dry forests, glades, barrens, forest openings; uncommon, rare in Piedmont and Coastal Plain (SC Rare List). July-September. Widespread in ne. North America, south in the mountains to SC, GA, and Al. [= FNA; Lycopodium tristachyum Pursh -- RAB, C, F, G, K, S, W]

D. Hhabereri (House) Holub [D. digitatum H tristachyum; Lycopodium H habereri House] is known from widely scattered localities in our area.

D. Hsabinifolium (Willdenow) Holub [D. sitchense (Ruprecht) Holub H tristachyum] ranges south to e. PA, where believed extirpated.

Huperzia Bernhardi (Firmoss, Clubmoss)

A genus of about 10-15 species, north temperate and arctic (and tropical mountains of Asia).

1. Leaves oblanceolate, the apical portion toothed with 1-8 large, irregular teeth; leaves 6-15 mm long, 1.0-2.5 mm wide; plants mainly of forest soils; stomates on lower leaf surface only (visible at 10H or preferably 20-40H); spores 23-29 :m in diameter......H. lucidula

1. Leaves lanceolate (awl-shaped), margins not toothed, or minutely toothed in the apical portion only with 1-3 low teeth; leaves 3-9 mm long, 0.6-1.3 mm wide; plants mainly of rock outcrops; stomates on both leaf surfaces (visible at 10H or preferably 20-40H); spores 29-38 :m in diameter......2

Huperzia appalachiana Beitel & Mickel, Appalachian Firmoss. Mt (NC, VA): rock outcrops at high elevations (very rarely at middle elevations), rarely also in seepage or along banks of small streams at high elevations, and in fens (on hummocks); rare (NC Significantly Rare, VA Rare List). June-August. This species was newly named in 1992 (Beitel & Mickel 1992). Though morphologically only subtly differentiated from the circumboreal H. selago (for distinctions see Beitel & Mickel 1992 and Brunton, Wagner, & Beitel 1992), the case for the distinctness of H. appalachiana is confirmed by the production of sterile (abortive-spored) hybrids where it co-occurs with H. selago. The range of H. appalachiana is Greenland, n. QuJbec and Newfoundland west to Ontario, MI, and MN and south along the Appalachians to NC, TN, and GA. H. selago is circumboreal, ranging south in North America to New England and the Great Lakes region, and disjunct to Oh. It should be sought in the high mountains of our area. It differs from H. appalachiana in having shoots with annual constrictions (vs. lacking constrictions), in producing gemmiferous branchlets strictly in 1 pseudowhorl at the end of the annual growth (vs. producing gemmiferous branchlets in 1-3 pseudowhorls or throughout the annual growth), the gemmae 4-5 mm long, 3-4.5 mm wide, the lateral or "wing" leaves 1.5-2 mm wide (vs. gemmae 3-4 mm long, 2.5-3.5 mm wide, the lateral or "wing" leaves 0.5-1 mm wide). [= FNA; Lycopodium selago Linnaeus -- RAB, S, W, in part; Lycopodium selago Linnaeus var. appressum (Desvaux) Petrovic -- C, F, misapplied, in part; Lycopodium selago var. selago -- C, G, in part; Huperzia selago (Linnaeus) Bernhardi ex Martius & Schrank var. selago -- K, in part]

Huperzia lucidula (Michaux) Trevisan, Shining Firmoss, Shining Clubmoss. Mt (NC, SC, VA), Pd (NC, VA), Cp (VA): moist forests and ravines; common (uncommon in Piedmont and Coastal Plain). June-August. Widespread in ne. North America, south to SC, TN, IN, IL, and MO. [= FNA, K; Lycopodium lucidulum Michaux -- RAB, C, F, G, S, W]

Huperzia porophila (Lloyd & Underwood) Holub, Rock Clubmoss. Mt (NC, SC, VA): rock outcrops and cliffs, especially in the spray of waterfalls, at low to medium elevations; rare (NC Candidate, SC Rare List, VA Rare List). June-August. Centered in the sedimentary Central Appalachians, H. porophila ranges from ne. PA, WV, OH, and WI south to NC, TN, nw. AL, and e. MO. Waterway (1986) clarified the distinctions between H. porophila and H. lucidula. [= FNA, K; Lycopodium porophilum Lloyd & Underwood -- RAB, C, F, S, W; Lycopodium selago var. patens (Beauvois) Desvaux -- G, misapplied]

Several hybrids are known from our area; they usually occur in intermediate habitats (such as in thin soil at the base of cliffs) and generally are found in proximity to both parents, but sometimes occur in the absence of one or both parents. Hybrids can be recognized by their intermediate morphology. The hybrid H. appalachiana H lucidula may be expected at cliff bases on high elevation rocky summits. It is known from Chimney Rock Park, Rutherford County (the lowest elevation occurrence of H. appalachiana in NC) and from Roan Mountain, Mitchell County, and Grandfather Mountain, Avery County. The hybrid H. lucidula H porophila [H. Hbartleyi (Cusick) Kartesz & Gandhi] is reported for NC by Waterway (1986). Both of these hybrids can be told from their parents by the presence of stomates on both surfaces of the leaf (unlike H. lucidula), but their marked lower density on the upper surface (unlike H. appalachiana or H. porophila).

Lycopodiella Holub (Bog Clubmoss)

A genus of about 15-20 species, temperate and tropical.

1. Leaves of prostrate stems entire to sparsely ciliate-toothed; surface of the erect stems visible, not hidden by the leaves......2

1. Leaves of prostrate stems conspicuously ciliate-toothed; surface of the erect stems hidden by the leaves......3

3. Prostrate stems arching, not in contact with the ground (and rooting) all along their length, 8-11 mm wide (including leaves), the stem (stripped of leaves) 2-4 mm in diameter; leaves of the prostrate stem of one size and shape, spreading to ascending, 5-7 mm long, 0.5-0.7 mm wide; erect stems many, equally spaced along the prostrate stems, progressively shorter and sterile towards the apex of the prostrate stems......L. alopecuroides

3. Prostrate stems creeping, in contact with the ground (and rooting) all along their length, 12-19 mm wide (including leaves), the stem (stripped of leaves) 1-2.2 mm in diameter; leaves of the prostrate stems dimorphic, spreading to reflexed, the upper leaves smaller (4-5 mm long, 0.4-0.6 mm wide) than the lateral leaves (7-8 mm long, 0.7-1.8 mm wide); erect stems few, clustered well behind the apex of the prostrate stems, mostly fertile and subequal in length......4
Lycopodiella alopecuroides (Linnaeus) Cranfill, Foxtail Clubmoss. Cp, Pd, Mt (NC, SC, VA): savannas, seepages, and other wet, sandy sites; common (rare in Mountains and Piedmont). July-September. The species is essentially a Southeastern Coastal Plain endemic, ranging from se. MA south to FL and west to se. TX, and disjunct in the Cumberland Plateau of KY, TN, and VA, and e. Highland Rim of TN. [= FNA, K; Lycopodium alopecuroides Linnaeus -- RAB, C, G, S, W]

Lycopodiella appressa (Chapman) Cranfill, Southern Bog Clubmoss. Cp, Pd (NC, SC, VA), Mt (NC, SC): savannas, seepages, bogs; common (rare in Mountains and Piedmont). July-September. This species is primarily Southeastern Coastal Plain, ranging from se. Newfoundland and MA, south to FL, south to OK, AR, and TX, and disjunct in the mountains of KY, TN, NC, and in sw. MI. [= FNA, K; Lycopodium appressum (Chapman) Lloyd & Underwood -- RAB, C, S, W; Lycopodium inundatum var. bigelovii Tuckerman -- F, G]

Lycopodiella inundata (Linnaeus) Holub, Northern Bog Clubmoss. Mt (NC, VA): gravelly or sandy seepage areas in bogs at middle to high elevations; rare (NC Significantly Rare, VA Rare List). July-September. A circumboreal species, ranging south in the Appalachians to NC, where it was first found in 1986 (Weakley, in preP.). [= FNA, K; Lycopodium inundatum var. inundatum -- F, G; Lycopodium inundatum Linnaeus -- C, W]

Lycopdiella margueritae J.G. Bruce, W.h. Wagner, & Beitel, Northern Prostrate Clubmoss. Mt (VA): moist seepage areas; rare (VA Rare List). This newly described tetraploid species is known only from a few scattered occurrences, in MI and VA; it is likely to be more widespread. VA plants are best assigned here pending additional study; it is possible that VA plants will prove to be of a different genetic origin than the MI plants, and should then be considered a distinct taxon. [= FNA, K]

Lycopodiella prostrata (Harper) Cranfill, Featherstem Clubmoss, Prostrate Bog Clubmoss. Cp (NC, SC): savannas, seepages; uncommon. July-September. A Southeastern Coastal plain endemic, ranging from se. NC south to FL and west to TX. [= FNA, K; Lycopodium alopecuroides -- RAB, in part; Lycopodium prostratum Harper -- C, S]

All pairwise combinations of sympatric species form fertile hybrids (only L. inundata and L. prostrata are entirely allopatric and not known to hybridize). The following hybrids should be expected where the parents grow together. Which occur in our area is uncertain at present. L. alopecuroides H appressa [L. Hcopelandii (Eig.) Cranfill] L. alopecuroides H inundata L. alopecuroides H prostrata [L. Hbrucei Cranfill] L. appressa H inundata L. appressa H prostrata

Additional research on this genus in our area is needed. Two fertile tetraploid species were recently named from MI (Bruce, Wagner, & Beitel 1991), and additional cryptic or semicryptic species may be found in the Southeastern Coastal Plain.

Species of this genus are difficult to identify. They often grow together; it is not uncommon to find two or more species at a single site in the Coastal Plain. Juvenile plants, resprouting in spring or after fire are especially difficult. In contrast to the other species, Pseudolycopodiella caroliniana and, to a lesser degree, L. prostrata, are dorsiventrally flattened (or apparently distichous), but it seems that juvenile sprouts of all species are somewhat flattened.

Lycopodium Linnaeus (Ground-pine, Running Cedar) (see also Diphasiastrum, Huperzia, Lycopodiella, and Pseudolycopodiella)

References: Wagner, Beitel, & Moran (1989); Hickey (1977).

A genus of 15-25 species, mainly temperate and subarctic.

1. Erect leafy stems 10 mm or more in diameter (including the leaves), not treelike or fanlike......2

1. Erect leafy stems 3-8 mm in diameter (including the leaves), treelike or fanlike......3

3. Leaves of the main vertical axis spreading (30-90 degree angle to stem) in the vicinity of the lower lateral branches, prickly to the touch; branchlets round in cross-section, the 6 ranks of leaves (2 lateral ranks, 2 adaxial ranks, and 2 abaxial ranks) equal in length and spreading to ascending......L. dendroideum

3. Leaves of the main vertical axis appressed (15-30 degree angle to stem) in the vicinity of the lower lateral branches, soft to the touch; branchlets slightly to strongly dorsiventrally flattened in cross-section, the 6 ranks of leaves (4 lateral ranks, 1 adaxial rank, 1 abaxial rank) round or slightly to very unequal, the abaxial leaves more appressed and mostly shorter than (to equal to) the spreading lateral leaves......4

Lycopodium annotinum Linnaeus, Stiff Clubmoss, Bristly Clubmoss. Mt (VA): high elevation hardwood or coniferous forests; uncommon (NC Watch List). August-October. A circumboreal species, south in North America to NJ and MN, and in the Appalachians to WV and sw. VA. Two varieties have been considered to reach our area in VA: var. acrifolium Fernald and var. annotinum. They are doubtfully distinct but need further study. This species was reported for NC by Lellinger (1985) and FNA, and is apparently indicated as occurring in NC on the range map in Mickel (1979); there is apparently no documentation for these reports, though the species occurs in Grayson County, VA, a county adjacent to NC. There is also an old collection from the Great Smoky Mountains of TN. Its occurrence in NC is certainly plausible, and it should be sought. [= C, F, FNA, G, K, W; including L. annotinum var. acrifolium Fernald -- F, G, K]

Lycopodium clavatum Linnaeus, Running Clubmoss. Mt (NC, SC, VA), Pd, Cp (VA): openings, balds, roadbanks, open forests; uncommon (rare in Piedmont and Coastal Plain). July-September. Circumboreal, south in e. North America along the Appalachians to NC. [= RAB, FNA, W; L. clavatum var. clavatum -- C, F, G, K, S]

Lycopodium dendroideum Michaux, Tree Ground-pine, Round-branch Clubmoss, Prickly Tree-clubmoss. Mt (NC, VA): openings, balds, high elevation spruce-fir and northern hardwood forests; rare (NC Watch List, VA Watch List). July-September. The northernmost of the L. obscurum complex, ranging from n. QuJbec and Newfoundland west to AK, south to NC, MO, MN, SD, CO, MT, ID, and WA; also in Asia. [= FNA, K, W; Lycopodium obscurum var. dendroideum (Michaux) D.C. Eaton -- RAB, F, G; presumably included in concept of L. obscurum -- C]

Lycopodium hickeyi W.h. Wagner, Beitel, & R.C. Moran. Pennsylvania Ground-pine, Hickey's Tree-clubmoss. Mt (NC, VA): balds, bog margins, forest openings; rare (NC Watch List, VA Watch List). July-September. Ranging from n. QuJbec and Newfoundland west to MN, south to NJ, w. NC, and n. IN. [= FNA, K; Lycopodium obscurum var. isophyllum Hickey -- W; presumably included in concept of L. obscurum -- C]

Lycopodium obscurum Linnaeus, Common Ground-pine, Flat-branched Tree-clubmoss. Mt, Pd, Cp (NC, SC, VA): acidic forests; common (uncommon in Piedmont and Coastal Plain). July-September. The southernmost of the L. obscurum complex, ranging from Nova Scotia and New Brunswick west to MI and WI, south to n. GA, n. AL, and IN. [= FNA, K; Lycopodium obscurum var. obscurum -- RAB, F, G, W; L. obscurum -- C, S, in part only]

L. lagopus (Laestadius ex C. Hartman) G. Zinserling ex Kuzeneva-Prochorova [= L. clavatum Linnaeus var. monostachyon Greville & Hooker] ranges south to c. PA.

Palhinhaea Vasconcellos & Franco (Nodding Clubmoss)

A genus of 10-15 species, tropical and subtropical.

Palhinhaea cernua (Linnaeus) Vasconcellos & Franco, Nodding Clubmoss, Staghorn Clubmoss. Cp (SC): disturbed moist areas; rare, perhaps adventive from further south this species is pantropical, occurring in the both the Neotropics and the Paleotropics. Its occurrence in our area may be adventive. [= FNA; Lycopodiella cernua (Linnaeus) Pichi Sermolli var. cernua -- K; Lycopodium cernuum Linnaeus -- S]

Pseudolycopodiella Holub (Carolina Bog Clubmoss)

A genus of about 12 species, subcosmopolitan.

Pseudolycopodiella carolinian (Linnaeus) Holub, Carolina Bog Clubmoss, Slender Clubmoss. Cp (NC, SC, VA): savannas, seepages; uncommon, rare in VA (VA Rare List). July-September. This species occurs in se. North America, the West Indies, and is widespread in the Southern Hemisphere. In North America, it ranges from MA south to FL and west to e. TX. It is often placed in a separate and monotypic section, and warrants status as a genus separate from Lycopodiella. In addition to the morphologic distinctions, this species has considerable anatomical differences, a different base chromosome number than the four species of Lycopodiella (x = 35 vs. x = 78), and does not hybridize with Lycopodiella (Wagner & Beitel 1992). [= FNA; Lycopodium carolinianum Linnaeus -- RAB, C, F, G, S; Lycopodiella caroliniana (Linnaeus) Pichi Sermolli var. caroliniana -- K]

The division of North American Lycopodium into three or more genera has been strongly advocated by Wagner & Beitel (1992) and numerous other recent authors. The traditionally broad Lycopodium appears to include a number of natural groups which are strikingly different from one another and have likely constituted separate lineages for many millions of years. These natural groups are separable by numerous morphological, developmental, and anatomical characters, karyotype, and inability to hybridize. Wagner & Beitel divide Lycopodium of our area into six genera in three subfamilies, as follows: Huperzia in Subfamily Huperzioideae, Lycopodium and Diphasiastrum in Subfamily Lycopodioideae, and Lycopodiella, Palhinhaea, and Pseudolycopodiella in Subfamily Lycopodielloideae. The reasoning behind this division is very strong, and it is here followed. Profound diferences in anatomy, morphology, reproduction, gametophyte morphology, and karyotype support this separation. The chromosome numbers of our genera: Diphasiastrum (x=23), Huperzia (x=67, 68), Lycopodiella (x=78), Lycopodium (x=34), Palhinhaea (x=55), Pseudolycopodiella (x=35).

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Flora of the Carolinas and Virginia, Working Draft of 27 October 1997 -- KEY TO PTERIDOPHYTE GENERA. Alan S. Weakley. The Nature Conservancy, Southeast Regional Office, Southern Conservation Science Department.