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]......Huperzia1. 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
2. Leaves herbaceous, pale or yellow-green, dull, deciduous; principal leafy stems creeping (except erect and repeatedly branched in Palhinhaea); rhizome dying back annually to an underground vegetative tuber at apex; spores rugulate; [plants of wetlands, mostly on moist or wet sands or peats]; [subfamily Lycopodielloideae]......4
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 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. lucidula1. 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
2. Leaves spreading, (3-) 5-9 mm long, ca. 1 mm wide, usually sparsely toothed; stomates relatively few on the upper leaf surface (1-25 on each side of midrib); [plants of outcrops at low to medium elevations]......H. porophila
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
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
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).
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.