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Home , Austrolycopodium, Dendrolycopodium hickeyi, Diphasiastrum, Diphasiastrum digitatum, Diphasium, Huperzia

L01a. HUPERZIACEAE 1

Weakley 2019-06

Autogenerated from the Flora Manager database version 3.58 at 6/26/2019 2:44:49 PM

SECTION 1: LYCOPODIOPHYTA (CLUBMOSSES)

L01a. HUPERZIACEAE Rothmaler 1962 (FIRMOSS FAMILY) [in LYCOPODIALES]

A family of 3 genera and ca. 300 species, mainly tropical and subtropical. Within the lycophytes, and related genera ( and ) form a clade with “an isolated position”, basal to the remainder of the family, and warrant family rank as Huperziaceae (Haines 2003a). See discussion under about family circumscriptions. References: Beitel (1979); Haines (2003a); Lellinger (1985); Mickel (1979); Øllgaard in Kramer & Green (1990); Øllgaard (1987); Øllgaard (1987); Øllgaard & Windisch (2014); Øllgaard, Kessler, & Smith (2018); Snyder & Bruce (1986); Wagner & Beitel in FNA2 (1993b); Wagner & Beitel (1992); Wikström & Kenrick (2000); Wikström & Kenrick (2001).

1 Gemmae present; on soil or rocks; [temperate, of NC, SC, c. GA, and c. AL northwards] ...... Huperzia 1 Gemmae absent; plants epiphytic on trees; [tropical, of s. FL southwards] ...... Phlegmariurus

Huperzia Bernhardi (FIRMOSS, CLUBMOSS)

A of about 10-15 species, north temperate and arctic (and tropical mountains of ). References: Haines (2003a); Øllgaard in Kramer & Green (1990); Testo, Haines, & Gilman (2016); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000); Zhang & Iwatsuki in FoC (2013).

Identification Notes: 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.

Unkeyed taxa: Huperzia ×bartleyi, Huperzia ×protoporophila 1 oblanceolate, the apical portion toothed with 1-8 large, irregular teeth; leaves 6-15 mm long, 1.0-2.5 mm wide; stomates on lower surface only (visible at 10×, or preferably 20-40×, magnification); 23-29 μm in diameter; [mainly of soils] ...... 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; stomates on both leaf surfaces (visible at 10×, or preferably 20-40×, magnification); spores 29-38 μm in diameter; [mainly of rock outcrops]. 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); [of outcrops at low to medium elevations] ...... 2 Leaves ascending to spreading, 2-7.5 mm long, 0.6-0.8 (-1.0) mm wide, not toothed (though sometimes with minute, single cell bumps); stomates relatively many on the upper leaf surface (30-90 on each side of midrib); [of high to medium elevations in our area]......

Huperzia appressa (Desvaux) A. Löve & D. Löve. APPALACHIAN FIRMOSS. 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). Jun- Aug. QC and NL (Newfoundland) west to ON, MI, and MN and south along the Appalachians to w. NC, e. TN, and ne. GA. Previous reports of this species from and e. Asia are based on misidentifications of H. continentalis, H. selago, and other species (Testo, Haines, & Gilman 2016). This species was named in 1992 as H. appalachiana (Beitel & Mickel 1992), but H. appressa (Desvaux) A. Löve & D. Löve is an older combination that applies to the same species (Haines 2003a). Though morphologically only subtly differentiated from the circumboreal H. selago (for distinctions see Beitel & Mickel 1992; Brunton, Wagner, & Beitel 1992; Haines 2003a), the case for the distinctness of H. appressa is confirmed by the production of sterile (abortive-spored) hybrids where it co-occurs with H. selago. [= K3, NE, Tn, Va, Haines (2003a), Testo, Haines, & Gilman (2016); = Huperzia appalachiana Beitel & Mickel – FNA2, K1; < Lycopodium selago Linnaeus – RAB, S, W; >< Lycopodium selago Linnaeus var. appressum (Desvaux) Petrovic – C, F; >< Lycopodium selago var. selago – C, G] Huperzia ×bartleyi (Cusick) Kartesz & Gandhi. BARTLEY'S FIRMOSS. Rock outcrops. Jun-Sep. Reported for NC by Waterway (1986). This hybrid can be told from its 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. porophila). {not keyed}. [= Il, K1, K3, Haines (2003a)] Huperzia lucidula (Michaux) Trevisan. SHINING FIRMOSS, SHINING CLUBMOSS. Moist and ravines. Jun-Aug. NL (Newfoundland) to MB, south to nw. SC, n. GA, n. AL, s. IL, and nw. AR (Peck 2011); disjunct in (Jilin) (FoC). [= Ar, FNA2, K1, K3, Mo1, NE, Pa, Tn, Va, Haines (2003a); = Lycopodium lucidulum Michaux – C, F, G, Md, Pa, RAB, S, W, WV; > Huperzia lucidula var. lucidula – Il] Huperzia porophila (Lloyd & Underwood) Holub. ROCK FIRMOSS, ROCK CLUBMOSS. Rock outcrops and cliffs, especially in the spray of waterfalls, at low to medium elevations. Jun-Sep. Centered in the sedimentary Central Appalachians, H. porophila L01a. HUPERZIACEAE 2

ranges from ne. PA, WV, OH, WI, and MN south to w. NC, nw. SC, ne. GA, nw. AL, and e. MO. Waterway (1986) clarified the distinctions between H. porophila and H. lucidula. [= FNA2, Il, K1, K3, Mo1, Pa, Tn, Va; = Lycopodium porophilum Lloyd & Underwood – C, F, RAB, S, W, WV; < Lycopodium selago var. patens (Palisot de Beauvois) Desvaux – G, misapplied]

Huperzia ×protoporophila A. Haines. Rock outcrops and cliff bases. Known from Chimney Rock Park, Rutherford County, NC (the lowest elevation occurrence of H. appressa in NC) and from Roan Mountain, Mitchell County, NC, and Grandfather Mountain, Avery County, NC. Expected at other cliff bases where the two parents are in proximity. This hybrid can be told from its 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. appressa). An additional useful character is the distribution of gemma-bearing branches: those of Huperzia appressa are abundantly distributed throughout the apical portion of mature plants, while those of the hybrid are confined to 1 or 2 pseudowhorls at the apex of annual growth (i.e., there are large gaps between the pseudowhorls of gemma-bearing branches). {mapped; not keyed}. [= NE, Haines (2003a)]

Phlegmariurus Holub 1964 (HANGING FIRMOSS)

A genus of 250-300 species, of tropical areas of the and Old World. References: Øllgaard in Kramer & Green (1990); Testo et al (2018); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000); Zhang & Iwatsuki in FoC (2013).

Phlegmariurus dichotomus (Jacquin) W.H. Wagner. HANGING FIRMOSS. Cypress swamps, epiphytic on Pond Apple (Annona glabra) and Florida Water Ash (Fraxinus cubensis). Jan-Dec. S. FL (Big Cypress Swamp); West Indies; south through to n. . [= FNA2, Øllgaard, Kessler, & Smith (2018), Testo et al (2018); = Huperzia dichotoma (Jacquin) Trevisan – Fl2, Meso, WH3; = Lycopodium dichotomum Jacquin – S]

L01b. LYCOPODIACEAE Palisot de Beauvois ex Mirbel 1802 (CLUBMOSS FAMILY) [in LYCOPODIALES]

A family of 10-15 genera and about 400 species. Lycopodiaceae+Hperziaceae, along with Selaginellaceae and , have now been shown to be only distantly related to other extant and plants (Pryer et al. 2001). The division of North American Lycopodium into three or more genera has been strongly advocated by Wagner & Beitel (1992), Wagner & Beitel in FNA (1993), Haines (2003a), and nearly all other recent authors. The traditionally broad Lycopodium appears to include a number of natural groups which are strikingly different from one another and have constituted separate lineages for tens to hundreds of millions of years. These natural groups are separable by numerous morphological, developmental, and anatomical characters, karyotype, and inability to hybridize. Wagner & Beitel (1992) divide Lycopodium (sensu latissimo) of our area into six genera in three subfamilies, as follows: Huperzia in Subfamily Huperzioideae, Lycopodium and in Subfamily Lycopodioideae, and , , and in Subfamily Lycopodielloideae. Haines (2003a) further divides Lycopodium (sensu lato) into three genera: , , and Lycopodium (sensu stricto). The reasoning behind this division is very strong, and it is here followed. Profound differences in anatomy, morphology, reproduction, morphology, and karyotype support this separation, in addition to the very great age of these lineages. The chromosome numbers of our genera: Dendrolycopodium (x=34), Diphasiastrum (x=23), Huperzia (x=67, 68), Lycopodiella (x=78), Lycopodium (x=34), Palhinhaea (x=55), Pseudolycopodiella (x=35), and Spinulum (x=34). Øllgaard in Kramer & Green (1990) and Wikström & Kenrick (2000) follow a somewhat broader coarse, recognizing three genera for our species (corresponding to the subfamilies of Wagner & Beitel 1992), and recognizing as sections the genera of Wagner & Beitel (1992). Øllgaard states that the “genera are very distinct, and also the sections within Lycopodiella and Lycopodium seem to represent ancient, independent evolutionary lines”; later, Øllgaard has elevated the sections to generic rank (Øllgaard & Windisch 2014). Wikström & Kenrick (2000, 2001) suggest that the phylogenetic separation of Lycopodium (including Diphasiastrum) and Lycopodiella (including Pseudolycopodiella and Palhinhaea) occurred at least as long ago as the early Jurassic (208 million years before present), and the divergence of Huperzia from Lycopodium and Lycopodiella still longer ago. Based on this deep division between Huperzia and the other genera, some authors additionally advocate the recognition of Huperzia in a separate L01b. LYCOPODIACEAE 3

family, Huperziaceae, an opinion followed here. The generic used here follows PPG I (2016). References: Beitel (1979); Haines (2003a); Lellinger (1985); Mickel (1979); Øllgaard in Kramer & Green (1990); Øllgaard (1987); Øllgaard, Kessler, & Smith (2018); PPG I (2016); Snyder & Bruce (1986); Wagner & Beitel in FNA2 (1993b); Wagner & Beitel (1992); Wikström & Kenrick (2000); Wikström & Kenrick (2001).

1 Leafy stems erect or pendant, simple or dichotomously branched, the ultimate branches vertically oriented; like the sterile leaves or only slightly reduced, in annual bands along the stem; vegetative reproduction by leafy gemmae near the stem apex ...... Huperziaceae 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. 2 Leaves herbaceous, pale or yellow-green, dull, deciduous; principal leafy stems creeping (except erect and repeatedly branched in Palhinhaea); dying back annually to an underground vegetative tuber at apex; spores rugulate; [of wetlands, mostly on moist or wet sands or peats]; [subfamily Lycopodielloideae]. 3 Upright shoots repeatedly branched; strobili nodding at the ends of the branches, 4-8 mm long; [known to occur from se. SC southward] ...... Palhinhaea 3 Upright shoots not branched; strobili erect on upright shoots, 1.5-80 mm long; [widespread in our area]. 4 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 (sporophylls) resembling leaves of the prostrate and upright stems in size and shape; strobilis 10-80 mm long; upright stems 1.5-15 mm in diameter (including the leaves) ...... Lycopodiella 4 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, their margins entire to minutely denticulate-fimbriate; strobilis 1.5-9 mm long; upright stems 1.5-3 mm in diameter (including the leaves) ...... Pseudolycopodiella 2 Leaves rigid, bright to dark green, shiny, evergreen; principal leafy stems mainly erect, treelike, fanlike, or creeping (if creeping, then the leaves with elongate, hyaline hair-tips); rhizome perennial, elongate, surficial or subterranean; spores reticulate; [of uplands, mostly in moist to dry soils]; [subfamily Lycopodioideae]. 5 Branches 1-5 mm wide (including the leaves), compressed to quadrangular, with 4 ranks of leaves; branching of strobilus stalks dichotomous ...... Diphasiastrum 5 Branches 4-12 mm wide, terete (to somewhat compressed in Dendrolycopodium obscurum), with 6 or more ranks of leaves; branching of strobilus stalks (when present), pseudomonopodial (falsely appearing to have a main axis from which branches arise). 6 Strobili borne on elongate, sparsely leafy peduncles borne at the tips of leafy, ascending branches; leaves with attenuate, hyaline hair-tips ...... Lycopodium 6 Strobili sessile, borne directly above densely leafy portions of upright branches; leaves acuminate to acute. 7 Erect leafy stems 3-8 mm in diameter (including the leaves), treelike or fanlike, with a definite main axis; leaves acute at the apex; horizontal shoots subterranean, without winter bud constrictions ...... Dendrolycopodium 7 Erect leafy stems 10 mm or more in diameter (including the leaves), branched 1-4 × sub-dichotomously; leaves with a 0.4-1.0 mm long stiff spinule; horizontal shoots at or near the ground surface, with winter bud constrictions ...... Spinulum

Dendrolycopodium A. Haines 2003 (TREE-CLUBMOSS)

A genus of 4 species, temperate and subarctic. Haines (2003a) makes the case for this genus as distinct from Lycopodium s.s. and other relatives. References: Haines (2003a); Hickey (1977); Øllgaard in Kramer & Green (1990); Wagner & Beitel in FNA2 (1993b); Wagner, Beitel, & Moran (1989); Zhang & Iwatsuki in FoC (2013).

1 Leaves of the main vertical axis spreading (30-90° 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 ...... Dendrolycopodium dendroideum 1 Leaves of the main vertical axis appressed (15-30° 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. 2 Abaxial leaves of the horizontal branchlets about the same length as the lateral leaves; leaves of all the ranks spreading at a (21°-) ca. 27° (-36°) angle from the branchlet, thus the branchlet and leaves together 3.5-6 (-7) mm wide ...... 2 Abaxial leaves of the horizontal branchlets about one half to two thirds as long as the lateral leaves; leaves of the abaxial and adaxial ranks generally appressed to the branchlet, the lateral 4 ranks spreading at a (27°-) ca. 40° (-59°) angle from the branchlet, thus the branchlet and leaves together ca. 6-9 mm wide ...... Dendrolycopodium obscurum

Dendrolycopodium dendroideum (Michaux) A. Haines. TREE GROUND-PINE, ROUND-BRANCH CLUBMOSS, PRICKLY TREE- CLUBMOSS. Openings, grassy balds, high elevation spruce- and northern hardwood forests. Jul-Sep. The northernmost of the L. obscurum complex, ranging from n. QC and NL (Newfoundland) west to AK, south to s. NJ, w. NC, MO, MN, SD, CO, MT, ID, and WA; also in Asia. [= Il, K3, NE, Tn, Va, Haines (2003a); = Lycopodium dendroideum Michaux – FNA2, K1, Mo1, Pa, W; < Lycopodium obscurum – C; < Lycopodium obscurum var. dendroideum (Michaux) D.C. Eaton – F, G, Md, RAB, WV] Dendrolycopodium hickeyi (W.H. Wagner, Beitel, & R.C. Moran) A. Haines. DELICATE TREE-CLUBMOSS, GROUND-PINE, HICKEY'S TREE-CLUBMOSS. Grassy balds, bog margins, forest openings, acidic ridgetop forests. Jul-Sep. N. QC and NL (Newfoundland) west to MN, south to NJ, sw. NC, and n. IN. [= Il, K3, NE, Tn, Va, Haines (2003a); = Lycopodium hickeyi W.H. Wagner, Beitel, & R.C. Moran – FNA2, K1, Pa; = Lycopodium obscurum var. isophyllum Hickey – W; < Lycopodium obscurum – C; < Lycopodium obscurum var. dendroideum (Michaux) D.C. Eaton – F, G, Md, RAB, WV] Dendrolycopodium obscurum (Linnaeus) A. Haines. COMMON GROUND-PINE, FLAT-BRANCHED TREE-CLUBMOSS. Acidic forests;. Jul-Sep. NS and NB west to MI and WI, south to n. GA, ne. AL, s. IN, n. IL (?), and c. MN; China, , , e. L01b. LYCOPODIACEAE 4

Russia. [= K3, NE, Tn, Va, Haines (2003a); = Lycopodium obscurum Linnaeus – FNA2, FoC, K1, Pa; = Lycopodium obscurum var. obscurum – F, G, Md, RAB, W, WV; < Lycopodium obscurum – C, S]

Diphasiastrum Holub 1975 (FLAT-BRANCHED CLUBMOSS, RUNNING CEDAR)

A genus of about 15-20 species, mostly north temperate and subarctic. This group is sometimes treated as Lycopodium section Complanata (Øllgaard in Kramer & Green 1990, Øllgaard 1987, Wikström & Kenrick 2000). References: Haines (2003a); Øllgaard in Kramer & Green (1990); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000).

Unkeyed taxa: Diphasiastrum ×habereri, Diphasiastrum sabinifolium 1 Foliage dark green, not glaucous; horizontal branchlets 2-4 mm wide (including the leaves); branchlets without conspicuous annual constrictions; 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 white portion of the vertical stem); abaxial rank of leaves shorter than lateral ranks (thus the branchlets flat in cross-section) ...... 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) ...... Diphasiastrum tristachyum

Diphasiastrum digitatum (Dillenius ex A. Braun) Holub. COMMON RUNNING-CEDAR, FAN GROUND-PINE. Dry to mesic, usually acid forests and openings, especially common in disturbed sites, such as successional pine forests. Jul-Sep. NL (Newfoundland) west to MN, south to SC, GA, AL, MS, and AR. Hickey & Beitel (1979) and Holub (1975a, 1975b) explain the nomenclatural decision to accept the epithet ‘digitatum’ over ‘flabelliforme’. [= Ar, FNA2, Il, NE, Pa, Tn, Va, Haines (2003a); = Lycopodium complanatum Linnaeus var. flabelliforme Fernald – F, G; = Lycopodium digitatum Dillenius ex A. Braun – C, K, Mo1, W; = Lycopodium flabelliforme (Fernald) Blanch – Md, RAB, S, WV] Diphasiastrum ×habereri (House) Holub. Dry forests. Jul-Sep. Known from widely scattered localities in our area; not always in close proximity to its parents. {not keyed}. [= FNA2, Il, NE, Haines (2003a); = Lycopodium × habereri House – K; = Lycopodium ×haberi – WV, orthographic error] Diphasiastrum sabinifolium (Willdenow) Holub. JUNIPER CLUBMOSS, JUNIPER-LEAF GROUND-CEDAR. NL west to ON, south to NY, e. PA, and MI. This taxon is derived from the hybrid of D. sitchense × tristachyum, but is often present with one or both parents absent, so is here treated as a hybrid-derived species. [= K3, Pa; = Diphasiastrum ×sabinifolium (Willdenow) Holub – FNA2, NE; = Lycopodium sabinaefolium Willdenow – C, F, G] Diphasiastrum tristachyum (Pursh) Holub. BLUE RUNNING-CEDAR, GROUND-CEDAR. Dry forests, glades, balds, barrens, forest openings. Jul-Sep. NL (Labrador) west to AB, south to nw. SC, ne. GA, ne. AL, MO, MN, and e. ND. [= FNA2, NE, Pa, Tn, Va, Haines (2003a); = Lycopodium tristachyum Pursh – C, F, G, K, Md, Mo1, RAB, S, W, WV]

Lycopodiella Holub 1964 (BOG CLUBMOSS)

A genus of about 15-20 species, temperate and tropical. 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. This group is variously treated as genus Lycopodiella, or as Lycopodiella section Lycopodiella (Øllgaard in Kramer & Green 1990, Wikström & Kenrick 2000), with a strong trend towards generic rank. References: Haines (2002a); Haines (2003a); Haines (2003b); Haines (2007a); Øllgaard in Kramer & Green (1990); Øllgaard (2012a); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000); Zhang & Iwatsuki in FoC (2013). L01b. LYCOPODIACEAE 5

Identification Notes: 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 southeastern Coastal Plain. Hybrids occur. Juvenile plants, resprouting in spring or after fire, are especially difficult to identify. 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.

Unkeyed taxa: Lycopodiella alopecuroides × appressa, Lycopodiella alopecuroides × prostrata, × inundata, Lycopodiella appressa × prostrata 1 Leaves of the horizontal shoots entire (rarely those toward the shoot apex with a few teeth); horizontal shoots, excluding the leaves, 0.5-0.9 (- 1.0) mm in diameter; each horizontal shoot segment commonly producing a single upright shoot; [in our area, a of the Mountains] ...... 1 Leaves of the horizontal shoots toothed (except when inundated); horizontal shoots, excluding the leaves, 1.5-5.0 mm in diameter; each horizontal shoot segment producing 2-6 upright shoots; [collectively primarily of the Coastal Plain, with some disjunctions inland into the Piedmont and Mountains]. 2 Fertile leaves (sporophylls) 2.9-5.0 (-5.2) mm long, appressed at maturity, entire or with short teeth < 0.3 mm long; strobili 3-6 mm in diameter at maturity ...... Lycopodiella appressa 2 Fertile leaves (sporophylls) 5.5-9 mm long, spreading, with 1-8 teeth per margin, some or all of the teeth exceeding 0.3 mm in length; strobili 10-20 mm in diameter at maturity. 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 toward the apex of the prostrate stems ...... Lycopodiella 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 Upright shoots 13-17 cm tall; horizontal stems 1.8-2.2 mm diam.; strobili 4-9 mm wide with incurved, ascending leaves; margins lacking obvious teeth; [sc. PA, nw. OH, ne. IN northwards] ...... Lycopodiella margueritae 4 Upright shoots 18-35 cm tall; horizontal stems 1.3-1.6 mm diam.; strobili 15-20 mm wide, with incurved, spreading leaves; sporophyll margins with 1-5 teeth; [NC and AR southwards] ...... Lycopodiella prostrata

Lycopodiella alopecuroides (Linnaeus) Cranfill. FOXTAIL CLUBMOSS. Pine savannas, seepages, wet pine flatwoods, wet prairies, bogs, ditches, and other wet, sandy sites. Jul-Sep. Primarily Southeastern Coastal Plain: se. MA south to FL and west to e. TX, and disjunct in the Cumberland Plateau of KY, TN, and VA, the Allegheny Mountains of WV (Morton et al. 2004), the e. Highland Rim of TN, and in ME (Haines 2001); s. Mexico south through Central America to n. South America; Cuba. The tropical portions of the distribution may be considered presumptive at this time; for instance, Øllgaard (2012a) elevates two taxa previously treated as varieties of Lycopodiella alopecuroides to species rank. [= Ar, ETx1, Fl1, FNA2, K1, K3, Meso, NE, Pa, Tn, TxFerns, Va, WH3, Haines (2002a); = Lycopodium alopecuroides Linnaeus – C, F, G, Md, S, W; < Lycopodium alopecuroides Linnaeus – RAB] Lycopodiella alopecuroides × appressa. [= Lycopodiella ×copelandii (Eiger) Cranfill – Ar, K1, K3, NE, WH3, Haines (2002a), Haines (2007a); = Lycopodium ×copelandii Eiger – Haines (2007a)] Lycopodiella alopecuroides × prostrata. [= Lycopodiella ×brucei Cranfill – K1, K3, WH3; = Lycopodium ×brucei (Cranfill) Lellinger] Lycopodiella appressa (Chapman) Cranfill. SOUTHERN BOG CLUBMOSS. Pine savannas, wet pine flatwoods, seepages, bogs, wet prairies. Jul-Sep. Primarily Southeastern Coastal Plain: se. NL (Newfoundland) and MA, south to FL, west to OK, AR, and TX, and disjunct in the mountains of KY, TN, NC, WV, and in sw. MI. [= Ar, ETx1, Fl1, FNA2, Il, K1, K3, NE, Tn, TxFerns, Va, WH3, Haines (2002a), Haines (2007a); = Lycopodium appressum (Chapman) Lloyd & Underwood – C, RAB, S, W; = Lycopodium inundatum Linnaeus var. bigelovii Tuckerman – F, G]

Lycopodiella appressa × inundata. [Lycopodiella ×gilmanii A. Haines – K3, Z]. Earlier tentative reports of Lycopodiella margueritiae J.G. Bruce, W.H. Wagner, & Beitel for the Mountains of Virginia are apparently based on this hybrid. See Haines (2003a, 2003b) for additional information. [= Lycopodiella ×gilmanii A. Haines – NE, Haines (2002a), Haines (2003a), Haines (2003b); = Lycopodiella margueritiae J.G. Bruce, W.H. Wagner, & Beitel – K, misapplied] Lycopodiella appressa × prostrata. Lycopodiella inundata (Linnaeus) Holub. NORTHERN BOG CLUBMOSS. Gravelly or sandy seepage areas, bogs. Jul-Sep. A circumboreal species, ranging south in the Appalachians to NC, where it was first found in 1986 (Weakley, in prep.). [= FNA2, FoC, Il, K, NE, Pa, Va, Haines (2002a), Haines (2003a), Haines (2003b); = Lycopodium inundatum Linnaeus – C, Md, W, WV; = Lycopodium inundatum var. inundatum – F, G] Lycopodiella margueritae J.G. Bruce W.H. Wagner, & Beitel. NORTHERN PROSTRATE BOG CLUBMOSS. Wet, acid sites. Sc. PA; nw. PA and ne. OH; MI and n. IN; WI. [= FNA2, K3] L01b. LYCOPODIACEAE 6

Lycopodiella prostrata (R.M. Harper) Cranfill. FEATHERSTEM CLUBMOSS, PROSTRATE BOG CLUBMOSS. Savannas, wet pine flatwoods, seepages, bogs, wet prairies. Jul-Sep. A Southeastern Coastal Plain endemic: se. NC south to c. FL and west to AR and e. TX, with scattered occurrences disjunct inland (as in n. GA, n. AL, and c. AR). [= Ar, ETx1, Fl1, FNA2, K1, K3, TxFerns, WH3; = Lycopodium prostratum R.M. Harper – C, S; < Lycopodium alopecuroides – RAB]

Lycopodium Linnaeus 1753 (RUNNING CLUBMOSS)

A genus of 5-10 species, mainly temperate and subarctic (but L. clavatum on nearly all continents). The fractionation of Lycopodium has resulted in the creation of more natural genera, more comparable to those in other groups of plants. References: Haines (2002b); Haines (2003a); Hickey (1977); Øllgaard in Kramer & Green (1990); Øllgaard & Windisch (2014); Rumsey (2007); Wagner & Beitel in FNA2 (1993b); Wagner, Beitel, & Moran (1989); Zhang & Iwatsuki in FoC (2013).

1 Strobili (1-) 2-3 (-5), borne on alternate "pedicels" branching from the central "peduncle"; "peduncle" (2.5-) 3-7 (-13) cm long; leaves 4-6 mm (not including the hair point), spreading to loosely ascending; upright shoots each usually bearing 3-6 branches, these disposed in an oblique or spreading posture ...... 1 Strobili 1 (rarely 2, if then, the 2 strobili not on separate "pedicels," but sessile and paired at the top of the "peduncle"); "peduncle" 0-1.2 cm long; leaves 3-5 mm long (not including hair point), ascending to appressed; upright shoots each usually bearing 2-3 branches, these disposed in an ascending or upright posture ......

Lycopodium clavatum Linnaeus. RUNNING CLUBMOSS. Openings, balds, roadbanks, open forests. Jul-Sep. Circumboreal, south in e. along the Appalachians to NC and n. GA; also c. Mexico south through Central America to n. South America (Brazil); West Indies. Additional taxonomic study is needed of what may prove a complex of species. [= FNA2, FoC, Il, K1, K3, Md, NE, Pa, RAB, S, Tn, Va, W, Haines (2002b), Rumsey (2007); = Lycopodium clavatum ssp. clavatum – Øllgaard, Kessler, & Smith (2018); = Lycopodium clavatum var. clavatum – F, G, Il, K, Md, Pa, RAB, Tn, Haines (2002b); < Lycopodium clavatum Linnaeus – C, WV; > Lycopodium clavatum ssp. clavatum – Meso] Lycopodium lagopus (C. Hartman) G. Zinserling ex Kuzeneva-Prochorova. ONE-CONE CLUBMOSS. High elevation heathlands. Jul-Sep. Circumboreal, south in North America to c. PA (Rhoads & Klein 1993), Tucker County, in e. WV (Gottlieb 2002), n. IL, MT, WA, and AK. [= FNA2, Il, K1, K3, NE, Haines (2002b), Haines (2003a), Rumsey (2007); < Lycopodium clavatum Linnaeus – C, WV; > Lycopodium clavatum var. brevispicatum Peck – F; > Lycopodium clavatum var. megastachyon Fernald & Bissel – F, G; > Lycopodium clavatum Linnaeus var. monostachyon Greville & Hooker – F, G]

Palhinhaea Vasconcellos & Franco 1967 (NODDING CLUBMOSS)

A genus of about 25 species, tropical and subtropical. This group has been variously treated as the genus Palhinhaea (Øllgaard 2015; Wagner & Beitel in FNA 1993b; PPG 2016; Øllgaard, Kessler, & Smith 2018) or as Lycopodiella section Campylostachys (Øllgaard in Kramer & Green 1990; Wikström & Kenrick (2000). References: Øllgaard in Kramer & Green (1990); Øllgaard (2015); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000); Zhang & Iwatsuki in FoC (2013).

Palhinhaea cernua (Linnaeus) Vasconcellos & Franco. NODDING CLUBMOSS, STAGHORN CLUBMOSS. Wet savannas, ditches and other disturbed moist areas. Jun-Nov. E. SC south to s. FL, west to s. AR and e. TX; Neotropics; Paleotropics. Some of the L01b. LYCOPODIACEAE 7

occurrences in our area may be adventive and of short duration. MacRoberts et al. (2018) report on occurrences of this species in LA, e. TX, and s. AR. [= Ar, ETx1, FNA2, TxFerns, Øllgaard, Kessler, & Smith (2018); = Lycopodiella cernua (Linnaeus) Pichi Sermolli – Fl2, Meso, WH3; = Lycopodium cernuum Linnaeus – FoC, S; > Lycopodiella cernua (Linnaeus) Pichi Sermolli var. cernua – K1; > var. cernua – K3]

Pseudolycopodiella Holub 1983 (CAROLINA BOG CLUBMOSS)

A genus of about 12 species, sub-cosmopolitan, but especially tropical and subtropical. This group has often been treated as section of Lycopodium (or of Lycopodiella); it appears to warrant status as a genus separate from Lycopodiella (Øllgaard, Kessler, & Smith 2018). 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). Øllgaard in Kramer & Green (1990) and Wikström & Kenrick (2000) retain it as Lycopodiella section Carolinianae. References: Haines (2003a); Øllgaard in Kramer & Green (1990); Wagner & Beitel in FNA2 (1993b); Wikström & Kenrick (2000); Zhang & Iwatsuki in FoC (2013).

Pseudolycopodiella caroliniana (Linnaeus) Holub. CAROLINA BOG CLUBMOSS, SLENDER CLUBMOSS. Pine savannas, seepages. Jul-Sep. MA south to s. FL and west to e. TX; West Indies; Mexico to South America; Asia; . [= Ar, ETx1, FNA2, FoC, K3, NE, TxFerns, Va, Haines (2003a); = Lycopodiella caroliniana (Linnaeus) Pichi Sermolli – WH3; = Lycopodium carolinianum Linnaeus – C, F, G, Md, RAB, S; > Lycopodiella caroliniana (Linnaeus) Pichi Sermolli var. caroliniana – Fl2, K1]

Spinulum A. Haines (BRISTLY CLUBMOSS)

A genus of 3 species, north temperate and subarctic. References: Haines (2003a); Hickey (1977); Øllgaard in Kramer & Green (1990); Wagner & Beitel in FNA2 (1993b); Wagner, Beitel, & Moran (1989); Zhang & Iwatsuki in FoC (2013).

Spinulum annotinum (Linnaeus) A. Haines. STIFF CLUBMOSS, BRISTLY CLUBMOSS. High elevation hardwood or coniferous forests. Aug-Oct. A circumboreal species, south in North America to n. NJ, MN, SD, NM, AZ, and OR, and in the Appalachians to WV, sw. VA, and e. TN (Blount County). 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. [= NE, Tn, Va, Haines (2003a); = Lycopodium annotinum Linnaeus – C, FNA2, FoC, K, Pa, W; > Lycopodium annotinum var. acrifolium Fernald – F, G, WV; > Lycopodium annotinum var. annotinum – F, G, Md, WV; > Lycopodium annotinum var. pungens (La Pylaie) Desvaux – WV]

L02. ISOETACEAE Dumortier 1829 (QUILLWORT FAMILY, MERLIN'S-GRASS FAMILY) [in ]

A family of a single genus and about 250-300 species. References: Jermy in Kramer & Green (1990).

Isoetes Linnaeus 1753 (QUILLWORT, MERLIN'S-GRASS)

P.W. Schafran, L.J. Musselman, and A.S. Weakley

A genus of about 250-300 species, cosmopolitan in distribution. Allopolyploids, derived from crossing between parental species and genome duplication, represent roughly half of the named taxa in eastern North America. Molecular data suggest that many of the southeastern polyploid species contain complexes of morphologically similar but genetically unique taxa. These complexes represent independent polyploidy events between different sets of parental species. The appropriate systematic treatment of these complexes would recognize approximately twice as many species. At present, many polyploid species should be treated only as taxonomic, not evolutionary, entities. References: Boom (1982); Bray, Schafran, & Musselman (2018); Brunton & Britton (1996a); Brunton & Britton (1996b); Brunton & Britton (1997); Brunton & Britton (1998); Brunton & Britton (1999); Brunton & McNeill (2015); Brunton (2015); Brunton (2016); Caplen & Werth (2000a); Caplen & Werth (2000b); Hoot, Napier, & Taylor (2004); Jermy in Kramer & Green (1990); Kott & Britton (1983); Luebke & Budke (2003); Musselman & Knepper (1994); Musselman (2001); Musselman et al (1995); Musselman, Bray, & Knepper (1996); Musselman, Bray, & Knepper (1997); Musselman, Taylor, & Bray (2001); Schafran et al (2016); Schafran et al (2018); Singhurst et al (2011); Taylor et al. in FNA2 (1993b). L02. ISOETACEAE 8

Identification Notes: Hybrids are possible between many combinations of species. Hybrids are readily recognized by having of variable size, shape, and ornamentation (aborted). Leaf width measurements are at midpoint of the leaf length, well above the flared base. Leaf lengths are of undamaged leaves from leaf tip to the top of the rootstock.

Unkeyed taxa: echinospora, Isoetes saccharata, Isoetes septentrionalis, Isoetes species 2, 1 Plants with narrow (0.5-1.0 mm wide), stiff leaves 3-20 cm long; leaves ± erect; emergent in sand and fine gravel of shallow, ephemeral bedrock pools and seeps in woodland glades or on open outcrops. 2 Leaves grass-green; surface granular or unornamented, > 525 μm in diameter; [in basic (limestone) substrates]; [2n: n. AL, nw. GA and northward] ...... 2 Leaves gray-green to bright-green; megaspore surface ridged or tuberculate, < 525 μm in diameter; [in acidic substrates]. 3 Plants 10-15 cm tall, individual plants apparent; leaf bases often with shiny black phyllopodia; velum coverage < 30%; megaspores white to light gray, 400-525 μm in diameter. 4 Megaspores typically 440-510 μm in diameter; ornamentation pattern wrinkled to low tuberculate; [plants on granite outcrops]; [4n: Piedmont of e. AL (and w. GA?)] ...... Isoetes graniticola 4 Megaspores typically < 450 μm in diameter, ornamentation pattern sparsely wrinkled, short-ridged (some with pseudo-reticulate configuration) or densely tuberculate; [plants on granite or sandstone outcrops]; [2X: Piedmont of e. AL to s. VA; 4n populations reported in s. GA and se. VA] ...... Isoetes piedmontana 3 Plants 2-10 cm tall; individuals densely clustered or mat-forming, often not readily discernable; leaf bases pale (black phyllopods absent); velum coverage > 70% (as little as 25% in Isoetes species 1); megaspores light gray to black, < 400 μm in diameter. 5 Plants 3-8 (-10) cm tall; velum coverage 40-60% (occasionally as little as 25%); megaspores olive-gray to brown; [2n: Lancaster County, Piedmont of SC] ...... Isoetes species 1 5 Plants 2-4 (-7) cm tall; velum coverage 70-100%; megaspores pale gray to black. 6 Plants in large, sometimes densely clustered colonies, 3-5 (-7) cm tall, with globose rootstocks (some with elongated rootstalks); megaspores ± dull, densely ornamented with fine, often low and/ or obscure tubercles; [2n: Piedmont of GA] ...... 6 Plants in dense, continuous mats; frequently joined, 2-3 (-4) cm tall, with elongated rootstalks (no globose rootstocks); megaspores ± shiny, unornamented or obscurely wrinkled; [2n: Piedmont of GA] ...... 1 Plants with broad (1.25-2.25 mm wide), ± flaccid leaves > 15 cm long; submerged or emergent in mineral or organic substrate of swamps, marshes, ponds, rivers, shores or open graminoid swales. 7 Megaspores > 575 μm in diameter, with reticulate ornamentation. 8 Velum coverage < 30%; plants with leaves 15-20 cm long in fast-flowing open streams; [of the Mountains of TN or VA]. 9 Megaspore muri unevenly tall in congested, broken-reticulate pattern; [10n: n. VA] ...... 9 Megaspore muri evenly tall in uncongested, evenly-reticulate pattern; [8n: se. TN] ...... Isoetes tennesseensis 8 Velum coverage > 30%; plants with leaves 20-45 cm long in slow-flowing woodland stream channels; [of the Coastal Plain of GA, FL, and AL]. 10 Megaspore ornamentation congested with thinner but still moderately thick muri; velum coverage ± 30%; [6n: s. GA, ne. FL, se. AL] ...... Isoetes boomii 10 Megaspore ornamentation open with thick muri; velum coverage ± 60%; [6n: s. GA] ...... Isoetes georgiana 7 Megaspores < 575 μm in diameter; with reticulate to almost spiny or low tuberculate and/or vermiform ornamentation. 11 Velum coverage > 40% (uncommonly to ± 30% in ). 12 Plants with broad (1.25-2.25 mm wide), light yellow-green, ± firm, reflexed leaves; velum coverage 45-70% (uncommonly only 30%); megaspores with tall, uneven, thin muri in ragged-reticulate pattern; [2n: found in woodland seepages (less commonly in pools); [primarily of Mountains, uncommonly of Piedmont and Coastal Plain, PA to s. AL; also ne. MS] ...... Isoetes valida 12 Plants with narrow (1.0-1.5 mm wide), dark-green, straggly to widely reflexed leaves; velum coverage 80-100%; megaspores with low tubercles or broad ridges (mounds) of even height in broken to evenly reticulate pattern; [of Coastal Plain, in ± permanently flooded swamps (nearly endemic to FL)]. 13 Megaspores ca. 500 μm in diameter, with dense ornamentation pattern of small tubercles; leaves 1.25-1.5 mm wide; [2n: of Jackson Co., FL] ...... Isoetes chapmanii 13 Megaspores < 450 μm in diameter, with coarse ornamentation pattern of broad tubercles or loosely interconnected vermiform mounds; leaves ca. 1.0 mm wide; [2n: of FL, sw. GA, se. AL Coastal Plain] ...... Isoetes flaccida 11 Velum coverage < 40%. 14 Megaspore ornamentation of low vermiform muri in semi-reticulate pattern or with low tubercles. 15 Megaspore ornamentation of semi-reticulate pattern or with (usually obscure) tubercles; megaspores 440-480 (-520) μm in diameter (tetraploids). 16 Leaves firm, ± stiffly erect (Juncus-like); heavily brown-streaked and with 20-40% velum coverage; megaspore surface smooth, prominently ornamented with broad, often vermiform muri; [4n: of s. GA Coastal Plain] ...... Isoetes junciformis 16 Leaves reflexed to loosely erect; sporangium ± unmarked and with ca. 10% velum coverage; megaspore surface mealy, sparsely to (uncommonly) densely ornamented with ± obscure tubercles and/or narrow muri; [4n: of Piedmont, s. VA and n. NC] ...... Isoetes virginica 15 Megaspore ornamentation plain or with irregular pattern of distinct, often numerous tubercles; megaspores < 440 μm in diameter (diploids). 17 Megaspores < 360 μm in diameter; megaspores ornamented with dense pattern of ± narrow, short muri; velum coverage 15- 35%. 18 [2n: freshwater tidal marshes of the e. VA Coastal Plain] ...... Isoetes mattaponica 18 [2n: blackwater streams of the s. MS Coastal Plain] ...... Isoetes mississippiensis 17 Megaspores > 360 μm in diameter; megaspores plain or ornamented with vermiform muri and/or broad tubercles; velum coverage 5-15%; [ephemeral inland fresh water pools and swales]. L02. ISOETACEAE 9

19 Broad (± 1.3 mm wide) light green leaves, bases never becoming black-brown and hardened; megaspores ± 410 (to 440) μm in diameter in diameter; conspicuously ornamented with numerous low tubercles or ridges; [2n: woodland pools of Piedmont of VA to AL and se. MS ...... ssp. silvatica 19 Narrow (± 1 mm wide) gray-green leaves with shiny, black-brown bases when spores are mature; megaspores ± 380 μm in diameter in diameter; typically with plain and mealy surface or obscurely ornamented with scattered tubercles and ridges; [2n: wet prairies and open graminoid swales of s. MS, w. AL and westward Isoetes melanopoda ssp. melanopoda 14 Megaspore ornamentation in distinctly reticulate (‘honeycomb’) pattern or with dense pattern of tall, short-crested, almost echinate muri. 20 Plants emergent or aquatic; megaspores ± 460 (occasionally to 500) μm in diameter; evenly-reticulate ornamentation pattern of even-topped muri continuous to the equatorial ridge without an equatorial band; [2n: Atlantic Coastal Plain, Appalachian Mts and irregularly northward, south to c GA, ne AL] ...... Isoetes engelmannii 20 Plants primarily emergent or amphibious; megaspores > 520 (rarely to over 600) μm in diameter; unevenly-reticulate and interrupted ornamentation pattern of ragged-topped muri, with an equatorial band of few to numerous spines; [4n or 6n]. 21 Leaves narrow (< 2.0 mm wide), blackish-green to olive green; megaspore ornamentation broken-reticulate to almost spiny with numerous tubercles often with short, mostly stand-alone muri or with broken ridges, usually with an equatorial band of numerous spines. 22 Leaves of mature plants 20-45 cm long, 1.0-1.5 mm wide, slender, wiry, dark (blackish) green; velum coverage 10-20%; moderately congested, almost echinate megaspore ornamentation pattern; aquatic in blackwater streams; [4n: Coastal Plain, VA to AL] ...... Isoetes hyemalis 22 Leaves of mature plants 10-20 cm long, 1.25-2.0 mm wide, olive-green; velum coverage 25-30%; densely congested (uncommonly uncongested and low-reticulate) megaspore ornamentation pattern; emergent on tidal beaches. 23 Megaspore ornamentation open with few tubercles and with low, broad muri with even crests; [4n: Coastal Plain, s. VA to DC] ...... var. reticulata 23 Megaspore ornamentation densely congested with tubercles and moderately tall, narrow, short muri with uneven crests; [4n: Coastal Plain, VA northward] ...... Isoetes riparia var. riparia 21 Leaves broad (> 2.0 mm wide), dark green to bright green; megaspore ornamentation moderately to evenly reticulate with few or no standalone short muri or tubercles, usually with equatorial band absent or a plain band with few spines. 24 Irregularly reticulate megaspore ornamentation pattern of both interconnected, longer muri and short, stand-alone muri; velum coverage ± 30% (occasionally less); plants usually deeply rooted (> 20% of leaf length) in clay or clayey-sand; [4n: Coastal Plain; LA, MS, AL] ...... 24 Regularly reticulate megaspore ornamentation pattern of interconnected muri with few or no stand-alone muri; velum coverage 10-25%; plants usually shallowly rooted (10-20% of leaf length) in sand or silty-sand substrate. 25 Velum coverage 20-25%; megaspores with ± open, reticulate ornamentation pattern; [4n: Appalachian Mts and VA to AL Coastal Plain] ...... Isoetes appalachiana 25 Velum coverage 6-10 (-20)%; megaspores with ± congested, densely reticulate ornamentation pattern; [6n: NC Coastal Plain] ...... Isoetes microvela

Isoetes appalachiana D.F. Brunton & D.M. Britton. APPALACHIAN QUILLWORT. Seepages, small woodland streams, ephemeral wetlands, backwaters. A tetraploid species (2n=44), apparently derived from a northern I. engelmannii entity and I. valida (Hoot, Napier, & Turner 2004), genotype=NNVV. A southern variant is derived from a southern I. engelmannii entity and I. valida, genotype=SSVV. See Brunton & Britton (1997) for additional information. [= Fl1, K, Tn, Va, WH3, Musselman (2001); < Isoetes engelmannii A. Braun – C, FNA2, Pa, RAB, W, WV; < Isoetes engelmannii var. engelmannii – F, S; > Isoetes engelmannii var. georgiana Engelmann] Isoetes boomii N. Luebke. BOOM'S QUILLWORT. Shallow water of slow-moving streams. Known from Laurens County, GA, AL, and FL. A hexaploid species (2n=66). [= FNA2, K, WH3; < Isoetes boomii N. Luebke – Musselman (2001), (also see I. georgiana)] Isoetes butleri Engelmann. BUTLER'S QUILLWORT. Seepage areas on calcareous glades, sometimes also in seepage on non- calcareous rocks. May-Jun. KY, IL, MO and e. KS south to e. TX and c. TX; disjunct eastwards in sc. KY, c. TN, n. GA and n. and c. AL. A diploid species (2n=22), genotype=BB. [= Ar, C, ETx1, F, FNA2, G, Il, K1, K3, Mo1, S, Tn, TxFerns, Musselman (2001), Singhurst et al (2011)] Isoetes chapmanii (Engelmann) Small. CHAPMAN’S FLORIDA QUILLWORT. Springs, stream bottoms, river bottoms, ditches. FL Panhandle. This taxon should be recognized at species rank based on the polyphyly of Isoetes flaccida s.l. (Schafran et al. 2018). A diploid species (2n=22). [= Isoetes flaccida (Shuttleworth ex A. Braun) var. chapmanii Engelmann – K1, S; < Isoetes flaccida – Fl1, FNA2, K3, WH3, Musselman (2001)]

Isoetes echinospora Durieu. SPINY- QUILLWORT. In acid lakes, ponds, and rivers (submerged to emersed), tidal mud flats. Circumboreal, in North America from , NL (Labrador), and AK south to DE, n. OH, MI, WI, CO, and CA. [= FNA2, K3, Pa; = Isoetes tenella Léman – K1; > ssp. muricata – NE; > Isoetes echinospora var. braunii (Durieu) Engelmann – G; > Isoetes echinospora var. echinospora – F, G; > Isoetes echinospora var. muricata (Durieu) Engelmann – C, F, G; > Isoetes muricata Durieu] L02. ISOETACEAE 10

Isoetes engelmannii A. Braun. Usually in permanent water bodies with active current. May-Oct. A diploid species (2n=22). Apparently there are two cryptic taxa currently combined under the name I. engelmannii (Hoot, Napier, & Taylor 2004), genotype NN and genotype SS. [= Ar, Il, K, NE, Tn, Va, Musselman (2001); < Isoetes engelmannii A. Braun – C, FNA2, G, Pa, RAB, W, WV, (also see I. appalachiana; < Isoetes engelmannii var. engelmannii – F, Mo1, S] Isoetes flaccida Stuttleworth ex A. Braun. WINGED FLORIDA QUILLWORT. Springs, stream bottoms, river bottoms, ditches. S. GA and se. AL south to s. FL. A diploid species (2n=22). [= Isoetes flaccida var. flaccida – Brunton (2015); < Isoetes flaccida – Fl1, FNA2, K3, WH3, Musselman (2001), Singhurst et al (2011); > Isoetes flaccida var. flaccida – K1, S] Isoetes georgiana N. Luebke. QUILLWORT. Streams. Known only from GA (Colquitt, Dodge, Irwin, Tift, Turner, and Worth counties). A hexaploid species (2n=66). See Brunton & Britton (1996b) for additional information. Musselman (2001) indicated that this may be conspecific with I. boomii, but molecular data suggest both taxa are polyphyletic. [= FNA2, K; < Isoetes boomii N. Luebke – Musselman (2001)]

Isoetes graniticola D.F. Brunton. Granite flatrocks. Mar-Jun. Endemic to ec. AL and adjacent GA. A tetraploid species (2n=44). See Brunton (2016) for detailed information. [= Brunton (2016)] Isoetes hyemalis D.F. Brunton. WINTERGREEN QUILLWORT. Blackwater streams and sandy streambanks. Sc. VA south through e. and c. NC to GA, AL, and FL Panhandle (Nelson 2000), in the Coastal Plain and lower Piedmont. A tetraploid species (2n=44), apparently derived from 2 unknown or extinct species, X and Y (Hoot, Napier, & Taylor 2004). See Brunton, Britton, & Taylor (1994) and Brunton & Britton (1996a) for additional information on this species. [= Fl1, K, Va, WH3, Musselman (2001); < Isoetes engelmannii A. Braun – C, G, RAB; < Isoetes engelmannii var. engelmannii – F, S] Isoetes junciformis D.F. Brunton & D.M. Britton. RUSH QUILLWORT. Ephemeral wetland swales in bottomland hardwood swamps. In sw. GA Coastal Plain (Tift and probably Calhoun counties, GA). A tetraploid species (2n=44). See Brunton & Britton (1999) for additional information. [= Brunton & Britton (1999), Musselman (2001)] Isoetes lacustris Linnaeus. LAKE QUILLWORT. Submerged in lakes and permanent streams. Jul-Sep. Apparently extirpated from the historic locality in VA. A decaploid species (2n=110). [= C, FNA2, K1, K3, Va; > Isoetes macrospora Durieu – F, G, W]

Isoetes louisianensis Thieret. LOUISIANA QUILLWORT. Small streams, braided swamps, seepage areas. S. AL, MS, and LA. A tetraploid species (2n=44). [= FNA2, K1, K3] Isoetes mattaponica L.J. Musselman & W.C. Taylor. MATTAPONI RIVER QUILLWORT. Tidal rivers. Apparently endemic to freshwater tidal marshes along rivers flowing into the Chesapeake Bay. A diploid relative of I. acadiensis (2n=22). See Musselman, Taylor, & Bray (2001) for additional information on this species. [= K3, Va, Musselman, Taylor, & Bray (2001)] Isoetes melanopoda Gay & Durieu ex Durieu ssp. melanopoda. BLACKFOOT QUILLWORT. Floodplains. May-Sep. S. IN, IL, and MO south to ne. LA; probably represented eastward to c. TN and s. MS (the available material ambiguous) (Brunton & Britton 2006). A diploid species (2n=22). (Brunton & Britton 2006). This taxon should be treated at species rank relative to I. melanopoda ssp. silvatica, based on the polyphyly of Isoetes melanopoda s.l. (Schafran et al. 2018). [= K3, Tn, TxFerns; < Isoetes melanopoda – Ar, C, ETx1, FNA2, G, Il, K1, Mo1, Musselman (2001)] Isoetes melanopoda Gay & Durieu ex Durieu ssp. silvatica D.F. Brunton & D.M. Britton. EASTERN BLACKFOOT QUILLWORT. Clay soils in low woods, seeps on sandstone or granitic rocks, in NJ in clay-based depressions on Cape May. VA south (in the Piedmont and Coastal Plain) to sw. GA, sc. AL, and s. MS; disjunct in s. NJ. A diploid species (2n=22), genotype= PP. This taxon should be raised to species rank based on the polyphyly of Isoetes melanopoda s.l. (Schafran et al. 2018). [= K3; < Isoetes melanopoda – C, FNA2, G, K1, RAB, Va, Musselman (2001)]

L02. ISOETACEAE 11

Isoetes melanospora Engelmann. BLACK-SPORED QUILLWORT. In pools on granite flatrocks. Endemic to SC and GA. A diploid species (2n=22). [= S, Musselman (2001); < Isoetes melanopoda – RAB; < Isoetes melanospora Engelmann – FNA2, K] Isoetes microvela D.F. Brunton. Banks of rivers in the outer Coastal Plain, associated with calcareous soils (at least in some cases). May-Jul (-Sep). Endemic to e. NC. A hexaploid species (2n=66). See Brunton & Britton (1998) for additional information. [= K1, K3, Brunton & Britton (1998)] Isoetes mississippiensis S.W. Leonard, W.C. Taylor, L.J. Musselman, & R.D. Bray. MISSISSIPPI QUILLWORT. Submersed in sluggish Coastal Plain streams. June. Endemic to s. MS. A diploid species (2n=22). See Schafran et al. (2016) for detailed information. [= Schafran et al (2016)] Isoetes piedmontana (N.E. Pfeiffer) C.F. Reed. PIEDMONT QUILLWORT. In seepage on or near granitic flatrocks, rarely on sandstone (Altamaha grit and Ridge and Valley). VA south and west to ec. and n. AL; material in c. TX has been placed here, but is likely to warrant description as a separate taxon. [= K1, K3, Va, Musselman (2001), Singhurst et al (2011); < Isoetes melanopoda – RAB; < Isoetes virginica N.E. Pfeiffer – C, F, FNA2, G]

Isoetes riparia Engelmann ex A. Braun var. reticulata (A.A. Eaton) Proctor. RETICULATE SHORE QUILLWORT. A tetraploid species (2n=44). Traditionally lumped in I. riparia s.l. Isoetes riparia Engelmann ex A. Braun var. riparia. SHORE QUILLWORT. Shores. It is extirpated from most of its historic localities. A tetraploid species (2n=44). Isoetes riparia is believed to be a mid-Atlantic relative of I. septentrionalis. [= K3] Isoetes saccharata Engelmann. Tidal waters, lakes. [= K; = Isoetes riparia var. palmeri (A.A. Eaton) Proctor – G; < Isoetes riparia Engelmann ex A. Braun – C, FNA2, Va] Isoetes septentrionalis D.F. Brunton. NORTHERN QUILLWORT. Freshwater ponds and shores. Sep. ME, QC, and ON south to n. NJ and se. and c. PA. A tetraploid species (2n=44). See Brunton & McNeill (2015) for detailed information. [= Brunton & McNeill (2015)]

Isoetes species 1. Pools on granite flatrocks. Forty Acre Rock, Lancaster County, SC. Under current taxonomic study. Isoetes species 2. 'S QUILLWORT. On Altamaha grit sandstone. Coastal Plain of GA. [= Isoetes snowii in prep.] Isoetes tegetiformans Rury. MERLIN'S-GRASS. In shallow pools on granite flatrocks. Endemic to a few granite flatrocks in ec. GA (notably Heggies Rock), near the SC line. A diploid species (2n=22), genotype=TT. [= FNA2, K1, K3, Musselman (2001)] Isoetes tennesseensis N.T. Luebke & J.M. Budke. Rocky river shoals. Endemic to Polk County, TN, near the North Carolina-Georgia state line, in the Hiwassee River. An octoploid species (2n=88). See Luebke & Budke (2003) for additional information. [= K3, Tn, Luebke & Budke (2003); < Isoetes lacustris Linnaeus – FNA2, K1]

L02. ISOETACEAE 12

Isoetes tuckermanii A. Braun. TUCKERMAN'S QUILLWORT. Shores, freshwater tidal marshes. NS south to n. NJ (reports of this species further south are apparently in error). A tetraploid species (2n=44), apparently derived from hybridization of I. viridimontana M.A.Rosenthal & W.C.Taylor and an unknown or extinct species, Z (Hoot, Napier, & Taylor 2004), genotype=??ZZ. [= C, F, G; > – FNA2, K1, K3; > Isoetes tuckermanii A. Braun – FNA2, K1, K3] Isoetes valida (Engelmann) Clute. MOUNTAIN QUILLWORT, CAROLINA QUILLWORT. Bogs (growing in ), pools, ponds, and seeps. PA to s. AL. A diploid species (2n=22). Genotype=VV. [= K, Tn, Va, Musselman (2001); = Isoetes caroliniana (A.A. Eaton) N. Luebke – FNA2; = Isoetes engelmannii A. Braun var. caroliniana A.A. Eaton – F, S; < Isoetes engelmannii A. Braun – C, RAB, W, WV] Isoetes virginica N.E. Pfeiffer. VIRGINIA QUILLWORT. In woodland streams. Jul-Sep. See Brunton, Britton, & Wieboldt (1996) for additional information. [= C, K, Va; < Isoetes melanopoda Gay & Durieu ex Durieu – RAB; < Isoetes virginica N.E. Pfeiffer – C, F, FNA2, G, W, (also see I. piedmontana)]

L03. SELAGINELLACEAE Willkomm 1854 (SPIKEMOSS FAMILY) [in SELAGINELLALES]

A family of 1-several genera (the generic circumscriptions still unclear), and about 700 (or more) species. Selaginellaceae, along with Lycopodiaceae and Isoetaceae, now appear to be only distantly related to other extant pteridophytes and seed plants (Pryer et al. 2001). There has been some recent tendency to split based on groups that represent very old clades (comparable to the recognition of multiple genera in Lycopodiaceae), though this remains controversial (Soják 1992; Škoda 1997; Korall, Kenrick, & Therrien 1999; Korall & Kenrick 2002). By a moderate approach to generic segregation, we have two to three genera, in which case Selaginella itself (sensu stricto) is restricted to the type species and a close relative. PPG I (2016) and Weststrand & Korall (20161, 2016b) retained Selaginella as a single, broadly defined genus including all members of the family, with recognition of seven major clades worldide at subgeneric rank. References: Buck (1977); Jermy in Kramer & Green (1990); Lellinger (1985); Somers & Buck (1975); Tryon (1955); Valdespino in FNA2 (1993b); Weststrand & Korall (2016a); Weststrand & Korall (2016b).

1 Vegetative leaves monomorphic, spirally arranged around the stems; leaves acuminate and with a white or translucent apical hair-tip (the hair- tip rarely lost); fertile branch tip only slightly differentiated from the sterile portions of the stems ...... Bryodesma 1 Sterile leaves dimorphic, in 4 ranks, the ventral pair spreading laterally, the dorsal pair ascending, the pairs different in shape and/or size (anisophyllous); leaves acute, mucronate, lacking a white or translucent apical hair-tip; fertile branch tips strongly differentiated (into strobili) from the sterile portions of the stem 2 Rhizophores dorsal; stems articulate ...... Gymnogynum species 1 (=kraussiana) 2 Rhizophores ventral; stems not articulate ...... Lycopodioides

Bryodesma Soják 1992 (SPIKEMOSS)

A genus of about 100 species, widespread in distribution, but with highest diversity in the Americas. References: Buck (1977); Jermy in Kramer & Green (1990); Lellinger (1985); Somers & Buck (1975); Tryon (1955); Valdespino in FNA2 (1993b).

1 Apical hair-tip of the leaves twisted-contorted, 1.2-1.7 mm long (sometimes deciduous); strobili 3-6 mm long, 1.5-2 mm wide; leaves 0.15-0.3 mm wide, the marginal cilia absent, toothlike, or as much as 1/6 as wide as the leaf blade; budlike “arrested” branches present ...... Bryodesma tortipilum 1 Apical hair-tip of the leaves straight, 0.3-1.4 mm long (sometimes deciduous); strobili (5-) 10-35 mm long, 1-1.5 mm wide; leaves 0.2-0.45 mm wide, the marginal cilia 1/4-1/3 as wide as the leaf blade; budlike “arrested” branches present or absent. 2 Stems mostly creeping or turned up at the apex, forming mats 1.5-4 cm high; rhizome or rhizomatous stem absent; aerial present all along the stems; budlike “arrested” branches absent ...... Bryodesma rupestre L03. SELAGINELLACEAE 13

2 Stems mostly erect or ascending, forming compact clumps usually > 4 cm high; rhizome or rhizomatous stem present; aerial roots present only at or near the base of the erect stems; budlike “arrested” branches present. 3 Leaves of the underground (rhizomatous) stems not scalelike; rhizophores mostly aerial; sporophyll base pubescent; leaf and sporophyll apices often pubescent ...... Bryodesma acanthonota 3 Leaves of the underground (rhizomatous) stems scalelike; rhizophores mostly subterranean; sporophyll base glabrous; leaf and sporophyll apices glabrous. 4 Leaves mostly tightly appressed; base conspicuously pubescent; strobili distinctly larger in diameter than the subtending stem; sporophyll apex often recurved ...... Bryodesma arenicola ssp. arenicola 4 Leaves mostly loosely appressed; base usually glabrescent; strobili not distinctly larger in diameter than the subtending stem; sporophyll apex usually straight ...... Bryodesma arenicola ssp. riddellii

Bryodesma acanthonota (Underwood) Škoda. SPINY SPIKEMOSS, SAND SPIKEMOSS. Sandhills, Altamaha Grit glades. Jun- Aug. S. acanthonota ranges from se. NC south to s. FL, west to w. Panhandle FL. The complex comprising S. acanthonota, S. arenicola, and S. corallina has been treated variably. The complex ranges from se. NC south to s. FL and west to c. TX; see Tryon (1955) and Valdespino in FNA (1993b) for additional information on the complex. [= Selaginella acanthonota Underwood – FNA2, K1, K3, S; = Selaginella arenicola Underwood ssp. acanthonota (Underwood) R. Tryon; < Selaginella arenicola – Fl1, RAB, WH3] Bryodesma arenicola (Underwood) Soják ssp. arenicola. SAND SPIKEMOSS. Scrub, sandhills. E. GA south to s. FL, se. GA, and e. Panhandle FL. [= Selaginella arenicola Underwood – K3, S; = Selaginella arenicola Underwood ssp. arenicola – FNA2, K1; < Selaginella arenicola – Fl1, WH3] Bryodesma arenicola (Underwood) Soják ssp. riddellii (Van Eseltine) Škoda. RIDDELL’S SPIKEMOSS. Dry sands (longleaf pine sandhills, Florida scrub, etc.), granite outcrops, sandstone outcrops, Altamaha grit glades. Jan-Dec. E. and c. GA west to AR, s. OK, and c. TX. See Wilbur & Whitson (2005) for an explanation of the nomenclatural change at species rank. [= S. arenicola Underwood ssp. riddellii (Van Eseltine) R.M. Tryon – Ar, ETx1, FNA2, K1; = Selaginella corallina (Riddell) Wilbur & Whitson – K3, TxFerns] Bryodesma rupestre (Linnaeus) Soják. ROCK SPIKEMOSS. Granite flatrocks, other, mostly acidic, rock outcrops, occasionally on greenstone or calcareous shales. Jun-Sep. S. Greenland and NS west to BC, south to GA, AL, AR, OK, and WY. Valdespino in FNA (1993b) suggests that two or more cryptic or semicryptic species are present within what is currently called S. rupestris; additional study is needed. Reports for KY are unconfirmed. [= Va; = (Linnaeus) Spring – Ar, C, F, FNA2, G, Il, K1, K3, Ky, Md, Mo1, NE, Pa, RAB, S, Tn, W, WV]

Bryodesma tortipilum (A. Braun) Soják. TWISTED-HAIR SPIKEMOSS. Rock outcrops, mostly at high elevations. Jul-Sep. Endemic to the Southern Appalachians (rarely into the Piedmont) of NC, SC, and GA. Occurring close to TN and VA; it should be sought there. [= Selaginella tortipila A. Braun – FNA2, K1, K3, RAB, S, W]

Lycopodioides Boehmer 1760 (SPIKEMOSS)

A genus of ca. 650 species, primarily tropical and subtropical. References: Buck (1977); Jermy in Kramer & Green (1990); Lellinger (1985); Somers & Buck (1975); Tryon (1955); Valdespino in FNA2 (1993b); Zhang, Nooteboom, & Kato in FoC (2013).

1 Main stems erect, the plants to 5 dm tall. 2 Stems erect; stems with dense, stiff hairs; median leaves long-acuminate to bristle-tipped ...... Lycopodioides braunii 2 Stems vinelike, climbing; stems lacking hairs; median leaves acute ...... Lycopodioides willdenowii 1 Main stems creeping or ascending. 3 Margins of lateral leaves entire; lateral branches of the stems further branching 2-3 times ...... Lycopodioides uncinatum 3 Margins of lateral leaves dentate-serrate; lateral branches of the stems further branching 1-2 times 4 Median leaves 0.7-1.2 mm long, with bristle-tipped apex up to 1/3 the length of the leaf; plants < 6 cm in diameter; [s. FL] ...... Lycopodioides species 1 (=eatonii) 4 Median leaves (1-) 1.25-1.5 (-1.8) mm long, with an acute or acuminate apex; plants > 6 cm long or in diameter; [more widespread, c. FL northwards]. 5 Leaves with margins of 3-5 rows of transparent (hyaline) cells; stomates of lateral leaves confined to near the midrib on the upper surface ...... Lycopodioides ludovicianum 5 Leaves with margins undifferentiated or with 1-2 rows of slightly paler cells; stomates distributed over entire leaf surface 6 Apices of median leaves acute to attenuated, usually keeled but with vein not extending almost to tip; lateral leaves 1.4-2.2 mm × 0.8-1.3 mm; megaspores 0.29-0.35 (-0.38) mm in diameter, dull, closely reticulated ...... Lycopodioides apodum 6 Apices of median leaves long-attenuate to brisyled, veined, the apices frequently recurved; lateral leaves ca. 1-2 mm × 0.5-1.3 mm; megaspores 0.33-0.40 mm in diameter, shiny, more loosely reticulated ...... Lycopodioides species 2 (=eclipes) L03. SELAGINELLACEAE 14

Lycopodioides apodum (Linnaeus) Palisot de Beauvois. MEADOW SPIKEMOSS. Seepages, bogs, spray cliffs, stream margins, wet meadows, marsh edges, wet spots in lawns, other moist habitats. Jun-Oct. S. ME, NY, OH, s. IN, AR, and e. OK south to FL, GA, AL, MS, LA, and e. TX; c. Mexico south to Guatemala. Often overlooked by botanists as a or liverwort. L. ludovicianum of the Gulf Coast east to GA, and S. eclipes W.R. Buck, more northern/midwestern, are superficially very similar (see key). [= Va; = Diplostachyum apodum – S; = – Ar, FNA2, Il, K1, K3, Ky, Md, Mo1, NE, Pa, RAB, Tn, W, WV; = Selaginella apoda var. apoda – ETx1, Fl1, TxFerns, WH3; < Selaginella apoda (Linnaeus) C. Morren – C, F, G, Meso] * Lycopodioides braunii (Baker) Kuntze. TREELET SPIKEMOSS, BRAUN'S SPIKEMOSS. Naturalized around graveyards or gardens; rare, introduced, native of China. [= Selaginella braunii Baker – FNA2, FoC, K1, K3] Lycopodioides ludovicianum (A. Braun) Small. GULF SPIKEMOSS, LOUISIANA SPIKEMOSS. Limestone outcrops, ravine slopes, calcareous swamos and hammocks. Jul-Oct. Gulf Coastal Plain from ne. FL and sw. GA west to e. LA. [=; = Diplostachyum ludovicianum – S; = Selaginella apoda var. ludoviciana (A. Braun) B.F. Hansen & Wunderlin – Fl1, WH3; = Selaginella ludoviciana (A. Braun) A. Braun – FNA2, K1, K3] Lycopodioides species 1 (=eatonii) (Hieronymus ex Small) Small. EATON’S SPIKEMOSS, PYGMY SPIKEMOSS. Hammocks and sinkholes on limestone. Jan-Dec. S. FL; Bahamas. [= Diplostachyum eatonii – S; = Lycopodioides species 1; = Selaginella armata Baker var. eatonii (Hieronymus ex Small) B.F. Hansen & Wunderlin – Fl1, WH3; = Selaginella eatonii Hieronymus ex Small – FNA2, K3]

Lycopodioides species 2 (=eclipes). HIDDEN SPIKEMOSS. Moist, often calcareous areas. Jun-Oct. W. CT, w. MA, NY, and ON west to WI, south to s. OH, nc. KY, s. IN., s. IL, and c. AR. [= Lycopodioides species 2; = Selaginella eclipes W.R. Buck – Ar, FNA2, Il, K1, K3, Mo1, NE; < Selaginella apoda (Linnaeus) Fernald – C, F, G] * Lycopodioides uncinatum (Desvaux ex Poiret) Kuntze. BLUE SPIKEMOSS. Moist forests, disturbed hammocks; native of China. Introduced in sw. GA and other places in the Southeastern . [= (Desvaux ex Poiret) Baker – Fl1, FNA2, FoC, K1, K3, WH3] * Lycopodioides willdenowii (Desvaux ex Poiret) Kuntze. VINE SPIKEMOSSWILLDENOW'S SPIKEMOSS. Moist, disturbed hammocks (spread from cultivation); native of Asia. [ (Desvaux ex Poiret) Baker; = Selaginella willdenowii (Desvaux ex Poiret) Baker – Fl1, K3, Meso]

Bibliography 15

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