OPHIOGLOSSACEAE (Adder's-tongue Family)

A family of 5 genera and about 75 species. Wagner & Wagner in FNA (1993b) suggest that the "Ophioglossaceae may be only distantly related to the ferns and more closely related to Marattiales and certain seedplants, especially Cycadales..."

References: Wagner & Wagner in FNA (1993b).

1. Sterile portion of the leaf blade pinnate, pinnatifid, or more divided; fertile stalks branched, the sporangia sessile or stalked......Botrychium

1. Sterile portion of the leaf simple, unlobed; fertile stalks unbranched, the sporangia embedded in a linear spike......Ophioglossum

Botrychium Swartz (Grapefern, Moonwort, Rattlesnake Fern)

A genus of 50-60 species, subcosmopolitan, but primarily temperate.

1. Fertile stalk joined to stalk of sterile leaf blade near the rhizome, far below the base of the leaf blade, and usually at or below the surface of the ground; leaves evergreen...... 5

2. Sterile portion of the leaf blade 3-pinnate or even more finely divided; plants (9-) 30-50 cm tall; sterile blade herbaceous in texture......B. virginianum

2. Sterile portion of the leaf blade 1-2-pinnate; plants usually less than 20 cm tall; sterile blade fleshy in texture......3

3. Sterile pinnae (or pinnules of 2-pinnate blades) not contracted at the base, thus oblong or elongate; leaf blades pinnate, pinnate-pinnatifid, or 2-pinnate, 1-8 cm long, 0.8-6 cm wide; pinna or pinnule margins entire, lobed, or incised, the pinna apices round, obtuse, or acute...... 4

4. Sterile pinnae apices obtuse to round at the apex, their segments (and undivided pinnae) about as long as wide, round, obtuse, or truncate at the apex; leaf blade mostly short-petioled (rarely sessile), the petiole (0-) 1-3 cm long; leaf blade pinnate to pinnate-pinnatifid......B. matricariifolium

4. Sterile pinnae acute at the apex, their segments (and undivided pinnae) at least twice as long as wide, mostly lanceolate, acute at the apex; leaf blade sessile, leaf blade pinnate-pinnatifid to 2-pinnate......B. lanceolatum var. angustisegmentum

5. Sterile leaf 2-pinnate to 4-pinnate, not finely divided, the ultimate segments ovate or oblong, more than 8 mm wide...... 6

6. Sterile pinnae entirely divided into short, round or acute pinnules; lateral pinnules with an inconspicuous and poorly-developed central vein; plant producing 1 or 2 leaves per season...... 7

6. Sterile pinnae (or their terminal portion) elongate (the sides often nearly parallel), entire to shallowly lobed, not divided into pinnules; lateral pinnules with a conspicuous and well-developed central vein; plant producing 1 leaf per season...... 9

7. Sterile pinna and pinnule apices round to obtuse; ultimate segments cuneate, rounded, or truncate at the base; ultimate segments remote or overlapping.......8

8. Stalk of the basal sterile pinnae (10-) 15-70 mm long; roots irregularly ribbed, blackish; ultimate leaf segments fan-shaped, obovate, longer than wide, pinnately veined, the midrib weakly developed; sporulating August-October......B. jenmanii

8. Stalk of the basal sterile pinnae 4-15 (-20) mm long; roots smooth, yellowish; ultimate leaf segments about as long as wide, subflabellately veined, lacking a midrib; sporulating January-April......B. lunarioides

9. Sterile pinna and pinnule apices acute; ultimate segments mostly oblong or lanceolate-oblong, often more than 2H as long as broad; overwintering leaves bronze (or green if covered by leaves)...... 10

10. Sterile blade mostly 2-pinnate, the segments sharply serrulate......B. biternatum

10. Sterile blade mostly 3-pinnate (or more divided, those forms keyed above), the segments entire to obscurely serrulate or crenulate......B. dissectum

Botrychium biternatum (Savigny) Underwood, Southern Grapefern. Mt, Pd, Cp (NC, SC, VA): moist forests, clearings, old fields; common. August-October. Widespread in se. United States, from MD, PA, s. IN, s. IL, and se. MO south to FL and e. TX. [= RAB, C, FNA, K, S, W; B. dissectum var. tenuifolium (Underwood) Farwell -- F, G]

Botrychium dissectum Sprengel, Cut-leaf Grape Fern, Dissected Grapefern. Mt, Pd, Cp (NC, SC, VA): moist forests, clearings, old fields; common (rare in Coastal Plain and Sandhills of NC and SC). August-October. Widespread in ne. North America, from Nova Scotia and QuJbecwest to Ontario and MI, south to FL and TX; also in the West Indies. The two forms have caused much confusion. In our area, forma obliquum is much more common and widely distributed, often confused with B. biternatum. Forma dissectum is fairly common in our area only in VA (rare in NC and SC), occurring primarily in the Mountains. The different distributions of the 2 forms suggest that further research is needed. [= RAB, C, F, FNA, G, K, S, W; including B. dissectum var. obliquum (Muhlenberg ex Willdenow) Clute -- G; including B. obliquum Muhlenberg ex Willdenow -- S]

Botrychium jenmanii Underwood, Alabama Grapefern. Mt (NC, SC, VA), Pd (NC, SC): moist to dryish forests and disturbed areas; rare (NC Significantly Rare, VA Rare List). August-October. Ranging from VA and TN south to FL, AL, and e. LA; also in the West Indies. This species probably arose as a hybrid between B. biternatum and B. lunarioides (Michaux) Swartz, followed by polyploidization, resulting in a fertile taxon functioning as a species. [= C, FNA, K, W; B. alabamense Maxon -- RAB, S]

Botrychium lanceolatum (S.G. Gmelin) Angstr`m var. angustisegmentum Pease & A.H. Moore, Lanceleaf Moonwort, Narrow Triangle Moonwort. Mt (NC, VA): forests and grassy balds; rare (NC Significantly Rare, VA Rare List). July-August. Var. angustisegmentum ranges from Newfoundland and Ontario south to VA, WV, NC, OH, MI, and MN, and in the Rocky Mountains of Canada and MT. Var. lanceolatum is widespread in w. North America. The two varieties are genetically distinct (Farrar & Wendel 1996). [= C, F, G, K, W; B. lanceolatum ssp. angustisegmentum (Pease & A.H. Moore) R.T. Clausen -- FNA]

Botrychium lunarioides (Michaux) Swartz, Winter Grapefern. Cp (SC), Pd (NC): old fields, pastures, young forests; rare (NC Candidate, SC Rare List). January-April. Ranging from w. NC and s. SC south to n. FL, and west to e. TX and se. OK. Wagner (1992) proposes that B. lunarioides be treated in a new monotypic section, Hiemobotrychium, of Botrychium, subgenus Sceptridium. The species is hard to spot, and all the more difficult to find because of its phenology; the leaves appear in late fall and die by early spring. [= RAB, FNA, K]

Botrychium matricariifolium (A. Braun ex Duwell) A. Braun ex W.D.J. Koch, Daisyleaf Moonwort. Mt (NC, VA): forests (often successional) and old fields; uncommon (NC Significantly Rare). June-August. Ranging from Newfoundland and Alberta south to NC, TN, KY, WV, OH, IL, WI, MN, and ND. [= FNA, K, W; B. matricariaefolium -- F, G (orthographic variant); B. matricariaefolium var. matricariaefolium -- C]

Botrychium multifidum (S.G. Gmelin) Treviranus, Leather Grapefern. Mt (NC, VA): grassy balds and high elevation meadows; rare (NC Candidate, VA Rare List). August-September. Widespread in n. North America, from Labrador and Alaska south PA, OH (and in the mountains to VA and NC), IN, IL, IA, NE, CO, NM, and CA. [= C, FNA, K, W; B. multifidum var. multifidum -- F, G; including B. multifidum var. intermedium (D.C. Eaton) Farwell -- F, G]

Botrychium oneidense (Gilbert) House, Bluntlobe Grapefern. Mt (NC, VA), Pd (VA): moist or boggy forests, bogs; rare (NC Candidate, VA Watch List). July-October. Northeastern in distribution and local in its occurrence, from New Brunswick, QuJbec, and Ontario south to NC, TN, KY, IN, and WI. [= RAB, C, FNA, K, W; B. dissectum forma oneidense (Gilbert) Clute -- F ("embarassingly transitional"); B. multifidum var. oneidense (Gilbert) Farwell -- G]

Botrychium simplex E. Hitchcock var. simplex, Least Moonwort. Mt (NC, VA): forests; rare (NC Candidate, VA Rare List). May-June. Widespread in n. North America, from Newfoundland and British Columbia south to NJ, VA, NC, MI, IN, WI, IA, SD, WY, CO, NM, UT, NV, and CA. Wagner & Wagner in FNA (1993b) discuss variation within B. simplex. Farrar & Wendel (1996) indicate that 3 varieties of B. simplex have strong genetic divergence, comparable to that usually distinguishing species. [= C, F, G, K; B. simplex -- FNA, W, infraspecific taxa not distinguished]

Botrychium virginianum (Linnaeus) Swartz, Rattlesnake Fern, Sang-find. Mt, Pd, Cp (NC, SC, VA): in a wide range of fairly dry, mesic, and wet forests, cove forests; common. April-June. Widespread in North America, from Newfoundland and British Columbia south to FL and CA. B. virginianum is the only North American representative of section Osmundopteris (J. Milde) R.T. Clausen. [= RAB, C, FNA, G, K, W; B. virginianum var. virginianum -- F; Osmundopteris virginiana (Linnaeus) Small -- S; Botrypus virginianus (Linnaeus) Holub]

Ophioglossum Linnaeus (Adder's-tongue)

1. Underground stem narrowly cylindrical or irregularly elongate, 2-4 mm in diameter; fertile spikes without a sterile portion at the apex or the sterile portion inconspicuous; sterile blade 0.5-10 cm long, 0.2-5.5 cm wide, borne horizontally, ascending, or vertically...... 2

2. Sterile blade 0.2-1 cm wide, the polygonal venation areoles usually lacking both smaller areoles and free included veinlets......O. nudicaule

2. Sterile blade (0.5-) 1.2-5 cm wide, the polygonal venation areoles either with smaller areoles or with free included veinlets...... 3

3. Large areoles ot the sterile blade subdivided into smaller areoles, which lack free veinlets; sterile blade obtuse or acute...... 4

4. Sterile blade ovate-lanceolate, the base obtuse to nearly truncate, broadest less than a quarter of the way from the base to the apex; primary areoles mostly more than 2 mm wide, without included veinlets......O. petiolatum

4. Sterile blade ovate to elliptic, the base cuneate to obtuse, broadest between one quarter and one half of the way from the base to the tip; primary areoles mostly less than 2 mm wide, with included veinlets...... 5

5. Sterile blade ovate, broadest below the middle, obtuse at the base, dark green, shiny, firm in texture; basal frond sheath leathery and tending to persist; spores 35-45 : in diameter......O. pycnostichum

Ophioglossum crotalophoroides Walter, Bulbous Adder's-tongue. Cp (NC, SC): moist ditchbanks and grassy roadside flats; rare (or overlooked). March-September. A Southeastern Coastal Plain species, ranging from e. NC (Dare County) south to FL and west to TX; also in Mexico, the West Indies, Central America, and South America. [O. crotalophoroides -- RAB, FNA, S; O. crotalophoroides var. crotalophoroides -- K]

Ophioglossum engelmannii Prantl, Engelmann's Adder's-tongue, Limestone Adder's-tongue. Mt (NC?, VA): dry barrens and glades over calcareous rocks; uncommon (VA Watch List). March-June. Ranging from w. VA, IN, IL, KS, and AZ south to FL and TX; also in Mexico and Central America. Ascribed to NC by Wagner & Wagner in FNA (1993b), the documentation unknown. [= FNA, K, W]

Ophioglossum nudicaule Linnaeus f., Slender Adder's-tongue. Cp (NC, SC): lawns and other moist, grassy areas; rare or overlooked. Ranging from e. NC south to s. FL, west to TX; also in Mexico, the West Indies, Central and South America, Asia, and Africa. First reported from NC by Thomas & Marx (1979). [= FNA, K; including O. dendroneuron E.P. St. John]

Ophioglossum petiolatum Hooker, Long-stem Adder's-tongue. Cp (NC, SC, VA): maritime wet grasslands, moist ditchbanks, and grassy roadside flats; rare or overlooked (NC Watch List, SC Rare List, VA Rare List). March-November. Widespread in se. United States, from se. VA south to FL and west to TX and OK; also in the West Indies, Mexico, n. South America, and Asia. First reported for NC by Thomas & Marx (1979). Wagner & Wagner in FNA (1993b) suggest that this species is likely introduced in North America. [= RAB, FNA, K; O. floridanum E. St. John]

Ophioglossum pusillum Rafinesque, Northern Adder's-tongue. Mt (NC?, VA): moist streamside meadow; rare (VA Rare List). March-July. Ranging from Nova Scotia west to ND, south to VA, possibly NC, IN, and NE; and in the Pacific Northwest. [= FNA, K; O. vulgatum Linnaeus var. pseudopodum (Blake) Farwell -- F]

Ophioglossum pycnostichum (Fernald) A. & D. L`ve, Southern Adder's-tongue. Pd, Mt, Cp (NC, SC, VA): bottomland forests, moist loamy soils of successsional forests and old fields; uncommon (or overlooked) (SC Rare List). March-July. Fairly widespread in e. North America, mostly south of the Wisconsinan glaciation, from s. NJ, IN, IL, and s. MI south to FL, MS, and e. TX. O. vulgatum is Eurasian. [= W; O. vulgatum Linnaeus var. pycnostichum Fernald -- RAB, C, F; O. vulgatum Linnaeus -- FNA, G, K, S]

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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.