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DRYOPTERIDACEAE (Wood-fern Family)

References: Smith in FNA (1993b); Lellinger (1985).

A family of about 47 genera and 1700 species, cosmopolitan in distribution, but concentrated in temperate and montane areas.

1. Leaves dimorphic, the pinnae or pinnules of the fertile leaves contracted, brown, and more-or-less beadlike......2

1. Leaves monomorphic (the fertile like the sterile), or slightly dimorphic (the fertile differing from the sterile only in size, shape, or orientation)......3

3. Sori elongate, indusia present and flaplike, attached along a long side; vascular bundles in the petiole 2......4

3. Sori round, indusia present or absent, if present reniform, peltate, cuplike, or lateral (but not attached along a long side); vascular bundles in the petiole either 2 or 4-7......6

5. Leaves 1-pinnate-pinnatifid, the pinnae pinnatifid...... Deparia

5. Leaves 1-pinnate, the pinnae entire...... Diplazium

7. Leaf blade pentagonal in outline, the terminal pinna by far the largest; rhizome 5-8 mm in diameter; indusia present, thick, persistent, and reniform; vascular bundles in the lower petiole 4-7; [introduced species, naturalized in moist ravines in SC]...... Arachniodes simplicior

7. Leaf blade broadly triangular in outline, the basal pinnae by far the largest; rhizome ca. 1 mm in diameter; indusia absent; vascular bundles in the lower petiole 2; [native species of mountain peaks of n. NC and VA] ...... Gymnocarpium

9. Veins anastamosing, rejoining to form a netlike pattern; pinnae 4-25 pairs per leaf; [plants non-native, rarely naturalized] ...... Cyrtomium

9. Veins branching dichotomously, free, not rejoining to form a netlike pattern; pinnae 25-50 pairs on larger leaves; [plant a common native species] ...... Polystichum

11. Indusium attached under one side of the sorus, hoodlike or pocketlike, arching over the sorus; petioles glabrous or sparsely beset with scales, the petiole bases not persistent...... Cystopteris

11. Indusium attached under the sorus, cuplike (divided into 3-6 lanceolate to ovate lobes which surround the sorus from below) or of minute numerous septate hairs, which extend out from under the sorus on all sides; petioles often densely beset with scales, the petiole bases persistent...... Woodsia


Arachniodes Blume (East Indian Holly Fern)

References: Smith in FNA (1993b).

A genus of about 50-60 species, of tropical and warm temperate regions, and especially of Asia and America.

* Arachniodes simplicior (Makino) Ohwi, Simpler East Indian Holly Fern. Pd (SC): moist banks in forested creek ravine; rare, introduced from Japan and China. Gordon (1981) discusses this interesting introduced population, apparently established for several decades at the time of its discovery, and likely originating from spores. [= FNA, K]


Athyrium Roth (Lady Fern)

References: Kato in FNA (1993b).

A genus of about 180 species, cosmopolitan in distribution, but concentrated in Asia.

1. Leaf blade widest near middle (the fourth or fifth pair of pinnae from the base the largest); margins of indusium toothed or ciliate (not glandular); rachis glandular; spores yellow or brown, smooth or papillate; petiole scales persistent, up to 1 cm long and 1.5 mm wide......A. angustum

1. Leaf blade widest near base (the second or third pair of pinnae from the base the largest); margins of indusium ciliate and glandular-ciliate; rachis eglandular; spores brown or dark brown, reticulate; petiole scales early deciduous, up to 5 mm long and 1 mm wide......A. asplenioides

Athyrium angustum (Willdenow) K. Presl, Northern Lady Fern. Mt (NC): rock outcrops on grassy balds at high elevations; rare (NC Watch List). June-September. The occurrence of this northern species is not fully documented; it was found in the 1980's by Murray Evans on Hump Mountain, on or near the TN-NC border. A. angustum ranges from Newfoundland and n. QuJbec west to Saskatchewan, south to w. NC, e. TN, OH, MO, and NE. [= S; A. filix-femina (Linnaeus) Roth ex Mertens var. michauxii (Sprengel) Farwell -- C, F, G; A. filix-femina var. angustum (Willdenow) G. Lawson -- FNA; A. filix-femina ssp. angustum (Willdenow) Clausen -- K, W; A. angustum (Willdenow) K. Presl -- S]

Athyrium asplenioides (Michaux) A.A. Eaton, Southern Lady Fern. Mt, Pd, Cp (NC, SC, VA): moist forests; common. May-September. Ssp. asplenioides ranges from MA, WV, IL, and KS south to n. FL and e. TX. Athyrium and Deparia superficially resemble Dryopteris, and they often grow together. Athyrium and Deparia have linear, flap-like sori (vs. rounded, reniform sori). Sterile individuals can be distinguished by the number of vascular bundles in the petiole; Athyrium and Deparia have 2, Dryopteris has 4-7. [= RAB, S; A. filix-femina (Linnaeus) Roth ex Mertens var. asplenioides (Michaux) Farwell -- C, F, FNA, G; A. filix-femina ssp. asplenioides (Michaux) HultJn -- K, W]


Cyrtomium K. Presl (Net-veined Holly Fern)

References: Yatskievych in FNA (1993b); MacDougal (1976).

A genus of about 15 species, of tropical and temperate regions of the Africa, Asia, and the Pacific Islands. Perhaps better treated as a portion of Polystichum.

1. Leaf coriaceous, the upper surface dark green and shiny; pinnae 4-10 (-12) pairs per leaf, 1.5-3 cm wide......C. falcatum

1. Leaf less coriaceous, the upper surface pale green and dull; pinnae (8-) 10-25 pairs per leaf, 1-2 cm wide......C. fortunei var. fortunei

*Cyrtomium falcatum (Linnaeus f.) K. Presl, Asian Net-veined Holly Fern. Cp (SC): old mortar of brick walls; rare, native of e. Asia. [= FNA, K, S; Polystichum falcatum Linnaeus f.]

*Cyrtomium fortunei J. Smith var. fortunei, Fortune's Net-veined Holly Fern. Cp (SC): old mortar of brick walls; rare, native of se. China. Two other varieties are known; neither appears to be naturalized in North America. [= FNA; C. fortunei -- K, infraspecific taxa not distinguished]

Cyrtomium is closely related to Polystichum. Both species are apogamous triploids.


Cystopteris Bernhardi (Bladder Fern, Brittle Fern)

References: Haufler, Moran, & Windham in FNA (1993b); Haufler, Windham, & Ranker (1990).

A genus of about 20 species, semicosmopolitan in distribution.

1. Lowest pair of pinnae the longest, thus the leaf widest at the base; bulblets often present on the rachis; indusia, rachises, and veins with stalked glands (these sometimes sparse in C. tennesseensis)......2

1. Lowest pair of pinnae shorter than the second or third pair, thus the leaf widest above the base; bulblets never present; indusia, rachises, and veins eglandular......3

3. Leaf blade (2.5-) 3-4H as long as wide; pinnae usually perpendicular to the rachis (or even reflexed); margins of pinnae serrulate, the teeth sharp; basal pinnules sessile, truncate to rounded at the base; indusium up to1 mm long, lanceolate; pinnae usually perpendicular to rachis; [plants on rock outcrops] ......C. fragilis

3. Leaf blade 2-2.5 (-3)H as long as wide; pinnae usually at an acute angle to the rachis, curving toward the blade apex; margins of pinnae crenulate, the teeth rounded; basal pinnules short-stalked or sessile, rounded to cuneate at the base; indusium about 0.5 mm long, ovate to round; pinnae usually at an acute angle to the rachis; [plants on rock outcrops or forest floor]......4

Cystopteris bulbifera (Linnaeus) Bernhardi, Bulblet Fern, Bulblet Bladder Fern. Mt, Pd (NC, VA): moist outcrops and talus of calcareous rocks, rarely up to 1500m elevation; uncommon, rare in Piedmont (SC Rare List). May-August. Ranging from Newfoundland west to MN, south to NC, AL, and AR; also disjunct in UT, AZ, NM, and TX. This species is a diploid involved in the reticulate evolution of Cystopteris in e. North America. It is one parent of C. tennesseensis. Its genome can be symbolized BB. [= RAB, C, F, FNA, G, K, S, W]

Cystopteris fragilis (Linnaeus) Bernhardi, Fragile Fern, Brittle Fern. Mt (NC, VA): cliffs, ascending in our area to 1650m; rare (VA Rare List). June-September. Circumboreal, in North America ranging from Newfoundland west to AK, south to MA, CT, NJ, montane NC, VA, KY, MO, OK, TX, NM, and AZ. This species is a fertile allotetraploid, presumed to be derived from hybridization between C. reevesiana Lellinger and an extinct or currently undiscovered second parent (C. Ahemifragilis@); its genome can be symbolized HHRR (Paler & Barrington 1995). C. fragilis appears to be a complex needing further study; additional entities may be found to warrant taxonomic recognition (see FNA for discussion). [= FNA, K, W; C. fragilis var. fragilis -- C, F, G, S]

Cystopteris protrusa (Weatherby) Blasdell, Lowland Bladder Fern. Mt, Pd (NC, SC, VA): rich woods or on moss- and soil-covered talus in boulderfields, occasionally on ledges of rock outcrops; common (SC Rare List). April-June. Ranging from NY and Ontario west to MN, south to GA, AL, MS, LA, AR, e. KS, and IA. This species is a diploid involved in the reticulate evolution of Cystopteris in e. North America. It is one parent of C. tennesseensis and C. tenuis. Its genome can be symbolized PP. [= RAB, C, FNA, K, W; C. fragilis var. protrusa Weatherby -- F, G, S]

Cystopteris tennesseensis Shaver, Tennessee Bladder Fern. Mt (NC, VA), Cp (NC): moist to dry outcrops of calcareous rocks, including coquina limestone ("marl") in the outer Coastal Plain; rare (NC Significantly Rare, VA Rare List). April-June. Ranging from PA, KY, IL, WI, and IA south to NC, n. AL, AR, and OK. This species is a fertile allotetraploid derived from hybridization between C. bulbifera and C. protrusa. Its genome can be symbolized BBPP. Haufler, Windham, & Ranker (1990) consider this a "successfully fledged and vigorous young species," adapted to a hybrid niche not successfully utilized by either parent [= RAB, C, FNA, K, W]

Cystopteris tenuis (Michaux) Desvaux, Mackay's Bladder Fern. Mt (NC, VA), Pd (VA): moist outcrops and cliffs of metamorphic and sedimentary rocks, occasionally in moist soils near rock outcrops; uncommon (NC Significantly Rare). May-August. Ranging from Newfoundland west to MN and NE, south to VA, IL, and MO, and in the mountains to NC, TN, and n. GA. This species is a fertile allotetraploid derived from hybridization between C. protrusa and an extinct or currently undiscovered second parent (C. Ahemifragilis@); its genome can be symbolized HHPP (Paler & Barrington 1995). [= FNA, K, W; C. fragilis var. mackayi Lawson -- C, F, G]

See Woodsia for suggestions on distinguishing between Cystopteris and Woodsia, similar ferns often confused.

Hybrids frequently occur where two or more species of Cystopteris grow in proximity. The following hybrids may be anticipated in our area:

Cystopteris bulbifera H tennesseensis.

Cystopteris bulbifera H tenuis [C. Hillinoensis R.C. Moran].

Cystopteris fragilis H tenuis.

Cystopteris protrusa H tennesseensis.

Cystopteris protrusa H tenuis.

Cystopteris tennesseensis H tenuis [C. Hwagneri R.C. Moran].

Other species in eastern North America: Cystopteris laurentiana (Weatherby) Blasdell of ne. and nc. North America, ranges south to c. PA.


Deparia Hooker & Greville

References: Kato in FNA (1993b).

A genus of about 40-50 species, primarily in tropical to warm temperate Asia and Africa.

1. Leaf blade narrowed to base; petiole bases swollen, with 2 rows of teeth; [plant a common native species of moist forests]; [section Lunathyrium] ......D. acrostichoides

1. Leaves widest at the base; petiole bases not markedly swollen, lacking teeth; [plant an exotic species, rarely introduced and naturalized]; [section Athyriopsis] ...... [D. petersenii]

Deparia acrostichoides (Swartz) M. Kato, Silvery Spleenwort. Mt (NC, SC, VA), Pd, Cp (NC, VA): moist forests, cove forests; common (uncommon in Piedmont, rare in Coastal Plain). June-September. Ranging from Nova Scotia west to MN, south to NC, SC, n. GA, n. AL, and AR. D. acrostichoides has several very closely related species in e. Asia. Unlike Athyrium, Deparia has the costal groove not continuous with the rachis groove. In addition, Deparia has multicellular hairs on the leaf blades. It stores starch in the swollen, persistent petiole bases. [= FNA, K, W; Athyrium thelypteroides (Michaux) Desvaux -- RAB, C, F, G; Diplazium acrostichoides (Swartz) Butters -- S]

Deparia petersenii (Kunze) M. Kato, native to se. Asia, has been introduced and naturalized in the Southeast, including in c. and s. GA, AL, and FL. It may occur in our area. [= FNA; Deparia japonica (Thunberg) M. Kato -- K, misapplied; Diplazium japonicum (Thunberg) Beddome, misapplied]


Diplazium Swartz (Twin-sorus Fern, Glade Fern)

References: Kato in FNA (1993b).

A genus of about 400 species, primarily tropical and north temperate in distribution.

Diplazium pycnocarpon (Sprengel) M. Broun, Glade Fern. Mt (NC, SC, VA), Pd (NC, VA), Cp (VA): very nutrient-rich, loamy or seepy forests, over calcareous sedimentary (such as limestone or dolostone) or mafic metamorphic or igneous rocks (such as greenstone or amphibolite); uncommon (rare in Coastal Plain and south of VA) (NC Watch List, SC Rare List). July-September. Widespread in e. North America, much more common in limestone areas of the Ridge and Valley than in the primarily acid-soil Blue Ridge and Piedmont. [= FNA, K; Athyrium pycnocarpon Sprengel -- RAB, C, F, G; Homalosorus pycnocarpos (Sprengel) Pichi-Sermolli -- S, W]

Other species in e. North America: Diplazium lonchophyllum Kunze of LA; Diplazium esculentum (Retzius) Swartz of FL and LA.


Dryopteris Adanson (Wood-fern, Shield-fern)

References: Montgomery & Wagner in FNA (1993b); Montgomery & Paulton (1981); Montgomery (1982).

A genus of about 250 species, subcosmopolitan, but concentrated in temperate Asia.

Note: Many Dryopteris species will hybridize with one another to form sterile hybrids. Whenever two or more Dryopteris species are found growing together, there is a good chance that hybrids are present. Hybrids generally show intermediacy between the two parents, and have abortive sporangia or spores. For further information on hybrids, see the discussion of hybrids following the 8 species accounts.

1. Leaves bipinnate-pinnatifid to tripinnate-pinnatifid (or to quadripinnate in the lower pinnae)......2

1. Leaves pinnate-pinnatifid to bipinnate (or to bipinnate-pinnatifid in the lower pinnae)......4

3. Leaf blade ca. 1H as long as the stipe; indusium occasionally glandular; first basal-pointed pinnule of the basal pinna 2.5-5H as long as the first tip-pointed pinnule of the basal pinna...... D. campyloptera

3. Leaf blade 2H as long as the stipe; indusium glabrous; first basal-pointed pinnule of the basal pinna ca. 2H as long as the first tip-pointed pinnule of the basal pinna...... D. carthusiana

5. Leaves dimorphic, the deciduous, fertile leaves erect, 2-3H as long as the spreading, evergreen, sterile leaves; fertile leaves linear-lanceolate in outline, generally 4-8H as long as wide; pinnae mostly 1.5-3H as long as wide, triangular, usually twisted out of the plane of the leaf axes, often nearly to 90 degrees...... D. cristata

5. Leaves not dimorphic, or only slightly so, deciduous or evergreen; fertile leaves lanceolate to ovate in outline, generally 1.5-4H as long as wide; pinnae mostly 3-4H as long as wide, borne in the plane of the leaf axes or only slightly twisted......6

7. Sori submedial, not touching the costule at maturity; leaf blade lanceolate, usually 2-4H as long as wide, gradually tapering at the apex; scales at the stipe base medium to dark brown, with a narrow black central band...... D. celsa

7. Sori medial, touching the costule at maturity; leaf blade ovate to narrowly ovate, usually 1.5-3H as long as wide; abruptly tapering at the apex; scales at the stipe base dark brown, nearly black, with a narrow pale margin...... D. goldiana

Dryopteris campyloptera Clarkson, Mountain Wood-fern. Mt (NC, VA): spruce-fir forests, northern hardwood forests; common (VA Watch List). July-September. Ranging from Newfoundland and n. QuJbec south to extreme n. PA, and from extreme s. PA south through e. WV and w. VA to e. TN and w. NC. This species is a fertile allotetraploid derived from hybridization of D. intermedia and the northern and western D. expansa (K. Presl) Fraser-Jenkins & Jermy, which does not (now) reach our area. The chromosome complement is symbolized EEII. [= RAB, C, K, S, W; D. spinulosa (O.F. Mueller) Watt var. americana (Fischer ex Kunze) Fernald -- F; D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. austriaca -- G]

Dryopteris carthusiana (Villars) H.P. Fuchs, Spinulose Wood-fern, Toothed Wood-fern. Mt, Pd, Cp (NC, SC, VA): acidic, organic-rich bogs, swamps, less frequently in moist rocky ravines and rich forests; common, uncommon to rare south and east of VA Mountains (SC Rare List). June-September. Irregularly circumboreal, in North America ranging from n. QuJbec west to Yukon, south to NC, SC, TN, AR, NE, w. MT, and WA. This species is a fertile allotetraploid derived from hybridization of D. intermedia and "D. semicristata," a hypothetical species which may now be extinct. Its chromosome complement is symbolized IISS. [= C, K, W; D. spinulosa (O.F. Mueller) Watt -- RAB, S; D. spinulosa var. spinulosa -- F; D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. spinulosa (O.F. Mueller) Fiori -- G]

Dryopteris celsa (W. Palmer) Knowlton, W. Palmer, & Pollard ex Small, Log Fern. Cp, Pd, Mt (NC, SC, VA): swamps, seepage bogs; uncommon. June-September. Ranging (scattered) from ne. NJ and ne. NY west to s. IL, e. MO, and AR, south to SC, n. GA, n. AL, TN, and n. LA; disjunct in w. NY and w. MI. This species is a fertile allotetraploid derived from hybridization of D. goldiana and D. ludoviciana; its chromosome complement is symbolized GGLL (Werth 1991). [= RAB, C, F, FNA, K, S, W; D. goldiana (Hooker ex Goldie) ssp. celsa W. Palmer -- G]

Dryopteris cristata (Linnaeus) A. Gray, Crested Wood-fern. Mt, Pd, Cp (NC, VA): bogs, swamp forests; uncommon. July-September. Circumboreal, in North America from Newfoundland to s. Saskatchewan and se. British Columbia, south to NC, TN, OH, IN, n. IL, IA, NE, and ID; disjunct in AL. This species is a fertile allotetraploid derived from hybridization of D. ludoviciana and "D. semicristata," a hypothetical species which may be extinct. Its chromosome complement is symbolized LLSS. Its has also served as a "parent species" of D. clintoniana, a fertile allohexaploid derived from D. cristata H goldiana. Thus, its genome constitutes two thirds of the genome of D. clintoniana. [= RAB, C, FNA, G, K, S, W; D. cristata var. cristata -- F]

Dryopteris goldiana (Hooker ex Goldie) A. Gray, Goldie's Wood-fern. Mt (NC, SC, VA), Pd (VA): boulderfield forests, rich cove forests, seepage swamps, especially over calcareous sedimentary or mafic metamorphic or igneous rocks; uncommon (NC Watch List, SC Rare List). June-September. Ranging from New Brunswick west to s. Ontario and MN, south to nw. SC, n. GA, n. AL, TN, KY, IL, and IA. This species is one of the diploid "parent species" of the e. North American reticulately-evolved Dryopteris complex. Its genome (symbolized GG) forms half of the genome of the tetraploid D. celsa, and one third of the hexaploid D. clintoniana, which does not occur as far south as our area. [= RAB, C, F, FNA, K, S, W; D. goldiana ssp. goldiana -- G]

Dryopteris intermedia (Muhlenberg ex Willdenow) A. Gray, Fancy Fern, Evergreen Wood-fern. Mt (NC, SC, VA), Pd, Cp (NC, VA): cove forests, other moist, rocky forests, over a variety of substrates; common (uncommon in Piedmont, rare in Coastal Plain) (SC Rare List). June-September. Ranging from Newfoundland west to MN, south to GA and AR. This species is one of the diploid "parent species" of the e. North American reticulately-evolved Dryopteris complex. Its genome (symbolized II) forms half of the genome of the tetraploids D. campyloptera and D. carthusiana. [= RAB, C, FNA, K, S, W; D. spinulosa (O.F. Mueller) Watt var. intermedia (Muhlenberg ex Willdenow) Underwood -- F; D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. intermedia (Muhlenberg ex Willdenow) Morton -- G]

Dryopteris ludoviciana (Kunze) Small, Southern Wood-fern. Cp (NC, SC): blackwater swamp forests; rare (NC Watch List). June-September. A Southeastern Coastal Plain species, ranging from e. NC south to s. FL, west to s. AL; disjunct in LA and AR. This species is one of the diploid "parent species" of the e. North American reticulately-evolved Dryopteris complex. Its genome (symbolized LL) forms half of the genome of the tetraploids D. cristata and D. celsa, as well as contributing one third of the genome of D. clintoniana indirectly (via its daughter species D. cristata). [= RAB, FNA, K, S]

Dryopteris marginalis (Linnaeus) A. Gray, Marginal Wood-fern. Mt, Pd, Cp (NC, SC, VA): rock outcrops, boulderfield forests, other rocky forests; common (less common in Piedmont, rare in Coastal Plain). June-September. Ranging from Newfoundland west to s. Ontario and MI, south to SC, GA, AL, TN, AR, and e. OK. D. marginalis has not participated in the reticulate evolution of Dryopteris in e. North America; it does, however, form sterile hybrids with some other species. [= RAB, C, F, FNA, G, K, S, W]

Other species in eastern North America: D. clintoniana (D.C. Eaton) Dowell has sometimes been attributed to our area, apparently by misidentification; it is known as far south as sc. PA and OH and may be documented as occuring in our area. Dryopteris fragrans (Linnaeus) Schott of n. North America, south to NH and VT. Dryopteris filix-mas (Linnaeus) Schott of n. North America, south to IL and VT.

The following hybrids are known between species which occur in our area. If the hybrid has been reported from our area, it is so indicated. In addition, the chromosome formulae are indicated, using the conventions listed at the end. These hybrids all have unbalanced chromosome complements which do not allow pairing. Thus, all produce aborted spores (if they produce spores at all), which can be recognized (at 30-40H) by their irregular size, shape, and color. For further information on these hybrids and a key to them, see Montgomery (1982).

D. campyloptera H intermedia. Known from NC. Chromosome formula = EII.

D. campyloptera H marginalis. Known from VA. Chromosome formula = EIM.

D. carthusiana H cristata [D. Huliginosa (A. Braun ex Dowell) Druce]. Known from VA. Chromosome formula = ILSS.

D. carthusiana H intermedia [D. Htriploidea Wherry]. Known from NC and VA. Chromosome formula = IIS. This is one of the commonest Dryopteris hybrids.

D. carthusiana H marginalis [D. Hpittsfordensis Slosson]. Chromosome formula = IMS.

D. celsa H cristata. Known from NC. Chromosome formula = GLLS.

D. celsa H goldiana. Chromosome formula = GGL.

D. celsa H intermedia [D. Hseparabilis (Wm. Palmer) Small]. Known from NC and VA. Chromosome formula = GIL.

D. celsa H ludoviciana [D. Haustralis (Wherry) Small]. Known from NC, SC, and VA. Chromosome formula = GLL.

D. celsa H marginalis [D. Hleedsii Wherry]. Chromosome formula = GLM.

D. cristata H intermedia [D. Hboottii (Tuckerman) Underwood]. Known from VA. Chromosome formula = ILS. This is one of the commonest Dryopteris hybrids.

D. cristata H marginalis [D. Hslossoniae Wherry ex Lellinger]. Known from VA. Chromosome formula = LMS.

D. goldiana H intermedia. Known from NC. Chromosome formula = GI

D. goldiana H marginalis. [D. Hneowherryi W.H. Wagner]. Known from NC and VA. Chromosome formula = GM.

D. intermedia H marginalis. Known from VA. Chromosome formula = IM.

E = D. expansa G = D. goldiana I = D. intermedia L = D. ludoviciana S = D. "semicristata" (hypothetical taxon, perhaps extinct)


Gymnocarpium Newman (Oak Fern)

References: Pryer in FNA (1993b); Pryer & Haufler (1993), abbreviated as Z; Pryer (1992). Key based on FNA.

A genus of about 8 species, north temperate in distribution.

1. Sessile basal basiscopic pinnule of the proximal pinnae with basal basiscopic pinnulet shorter than the adjacent pinnulet; pinnae of second pair sessile, with basal pinnules shorter than the adjacent pinnule (or second basal pinule rarely stalked); spores 27-31 Fm in diameter......G. appalachianum

1. Sessile basal basiscopic pinnule of the proximal pinnae with basal basiscopic pinnulet more or less equal in length to the adjacent pinnulet; pinnae of second pair usually sessile, with basal pinnules more or less equal in length to the adjacent pinnule; spores 34-39 Fm in diameter......[G. dryopteris]

Gymnocarpium appalachianum Pryer & Haufler, Appalachian Oak Fern. Mt (NC, VA): moist, rocky forests, at medium to high elevations; uncommon (US Species of Concern, NC Candidate, VA Watch List). June-September. This species, newly distinguished from G. dryopteris (Linnaeus) Newman, is a diploid, endemic to the c. and s. Appalachians (known from ne. WV, nw. VA, s. PA, and disjunct in nw. NC and OH). Electrophoretic and morphologic analyses show that it is one of the parents of the widespread allotetraploid G. dryopteris. In NC, it is limited to a single site, below the north-facing summit cliffs on Bluff Mountain, Ashe County, where see page results in extensive ice formations which frequently persist until June. [= FNA, K, Z; included within concept of G. dryopteris (Linnaeus) Newman by earlier authors]

Triploids are known from the mountains of VA. Their identity is uncertain; based on geography they are presumably G. appalachianum H dryopteris, but could be G. Hbrittonii (Sarvela) Pryer & Haufler [= G. disjunctum H dryopteris]. Triploids can be distinguished from G. appalachianum by the presence of malformed spores, irregular in shape and size, often intermixed with large round spores (vs. all spores reniform and relatively uniform in size and shape).

Gymnocarpium dryopteris (Linnaeus) Newman is circumboreal, occurring throughout northern and central Eurasia, Greenland, south in North America to MD (?), s. PA, OH, MI, WI, IA, w. SD, CO, n. NM, and c. AZ. Since it approaches our area from the north and closely resembles G. appalachianum, it should be carefully sought in our area, especially in the mountains of VA. The user is referred to Pryer & Haufler (1993) for a detailed and careful analysis of the distinguishing features between G. appalachianum and G. dryopteris. Gymnocarpium robertianum (Hoffmann) Newman of n. North America, south to MI, MN, and WI.


Matteuccia Todaro (Ostrich Fern)

References: Johnson in FNA (1993b).

A genus of 1 species, north temperate in distribution. Two other species formerly included in Matteucia are better treated in the genus Pentarhizidium Hayata (Gastony & Ungerer 1997).

Matteuccia struthiopteris (Linnaeus) Todaro var. pensylvanica (Willdenow) C.V. Morton, Ostrich Fern. Mt, Pd (VA): alluvial forests and calcareous wetlands; rare (VA Rare List). The species is circumboreal; the North American var. pensylvanica ranges from Newfoundland west to AK, south to VA, MO, SD, and British Columbia. The North American var. pensylvanica is separated from the Eurasian var. struthiopteris on the basis of its concolorous rhizome scales (vs. bicolorous scales) and less truncate pinna lobes. Matteuccia stores starch in its persistent petiole bases. [= FNA, G; M. struthiopteris -- C, K (infraspecific taxa not distinguished); Pteretis pensylvanica (Willdenow) Fernald -- F]


Onoclea Linnaeus (Sensitive Fern)

References: Johnson in FNA (1993b).

A genus of 1 species, of temperate e. North America and e. Asia.

Onoclea sensibilis Linnaeus var. sensibilis, Sensitive Fern, Bead Fern. Mt, Pd, Cp (NC, SC, VA): marshes, swamps, wet disturbed places; common. May-June. The species ranges from Newfoundland west to MN and CO, south to FL, TX, and CO; also in e. Asia. Var. sensibilis is North American; var. interrupta is Asian. The recognition of two varieties is supported by molecular evidence. The genus is monotypic. The specific epithet and common name refer to the fact that the fronds wither at the first touch of frost, not that they respond to touch. The peculiar fertile leaves (with their brown, beadlike, fertile pinnules) are collected for use in dried arrangements. The expanded, persistent petiole bases store starch. [O. sensibilis -- RAB, C, F, FNA, G, K, S, W, infraspecific taxa not distinguished]


Polystichum Roth (Holly Fern)

References: D. H. Wagner in FNA (1993b).

A genus of about 180 species, cosmopolitan in distribution.

Polystichum acrostichoides (Michaux) Schott, Christmas Fern. Mt, Pd, Cp (NC, SC, VA): moist to dry forests and woodlands; common. June-September. Ranging from Nova Scotia west to MN, south to s. FL and e. TX; also in Mexico. One of the most familiar ferns in e. North America. [= RAB, C, F, FNA, G, S, W; P. acrostichoides var. acrostichoides -- K]

Polystichum braunii (Spenner) FJe of circumboreal distribution, south in e. North America to NJ, ne. PA, and NY. Polystichum lonchitis (Linnaeus) Roth of n. North America, south to MI, MN, and WI.


Woodsia R. Brown (Woodsia, Cliff Fern)

References: Windham in FNA (1993b).

A genus of about 30 species, of temperate and cool-temperate regions.

1. Petioles with a distinct joint about 1-3 cm above the base, the petiole bases of former leaves forming a fairly even stubble; leaf blade lacking glands (though bearing both long septate hairs and pale linear scales); indusium of numerous filamentous segments......W. ilvensis

1. Petioles lacking a distinct joint, the petiole bases of former leaves disintegrating irregularly and forming an uneven stubble; leaf blade with stalked glands, at least below on the costae, costules, and veins (and also bearing nonglandular hairs and/or linear scales); indusium of 3-6 lanceolate segments......2

Woodsia appalachiana T.M.C. Taylor, Appalachian Woodsia, Appalachian Cliff Fern, Mountain Woodsia. Mt, Pd (NC, VA): on cliffs of sandstone, shale, granite, granitic gneiss, and hornblende gneiss; uncommon, rare in North Carolina (NC Significantly Rare). June-September. Endemic to the Southern and Central Appalachians of VA, WV, NC, and TN and the Ozarks of AR. This species is similar to W. scopulina of the western mountains of AK south to CO and CA. The eastern plants have been variously treated as a full species, a subspecies or variety of W. scopulina, or as indistinguishable from W. scopulina (see synonymy). It now appears that W. appalachiana may be a rather cryptic but distinct element of a reticulate complex also involving W. scopulina ssp. scopulina (of the Rocky Mountains) and W. scopulina ssp. laurentiana Windham (primarily of the Rocky Mountains but also disjunct eastwards in Ontario and QuJbec). Windham in FNA (1993b) treats these three entities as subspecies, and suggests that ssp. laurentiana is the allotetraploid derivative of hybridization of the eastern and western diploids. If this is indeed so, each of the 3 entities should be recognized at the species level. [= F, K; W. scopulina D.C. Eaton -- RAB, C, S, W (in a broad sense); W. scopulina ssp. appalachiana (T.M.C. Taylor) Windham -- FNA; W. scopulina var. appalachiana (T.M.C. Taylor) Morton -- G]

Woodsia ilvensis (Linnaeus) R. Brown, Rusty Woodsia, Rusty Cliff Fern. Mt (NC, VA): cliffs of amphibolite, greenstone, other rocks; uncommon, rare in NC and apparently only in the northernmost few counties of that state (NC Significantly Rare). June-September. Circumboreal, ranging in North America from Newfoundland and AK south to VA, nw. NC, OH, n. IL, nw. IA, Saskatchewan, and British Columbia. [= RAB, C, F, FNA, G, K, S, W]

Woodsia obtusa (Sprengel) Torrey ssp. obtusa, Common Woodsia, Blunt-lobed Cliff Fern. Mt, Pd, Cp (NC, SC, VA): rock outcrops of various sorts, moist talus, terrestrial near rock outcrops; common. June-September. Ranging from ME, QuJbec, MN, and e. NE, south to FL and TX. [= FNA, K; W. obtusa -- RAB, C, F, G, S, W, subspecies not distinguished]

Woodsia species and Cystopteris species are all small ferns with thin-textured leaves, occurring primarily on or near rock outcrops; they frequently occur together or in proximity to one another and are often confused. Woodsia has the indusium divided into a series of scale-like or hair-like structures, attached below the sorus; Cystopteris has an undivided indusium, pocket-like or hood-like, attached around one side of the sorus. Woodsia has persistent dark petiole bases; in Cystopteris the petiole bases are deciduous. Woodsia has the final veinlets not reaching the margin; Cystopteris veins do reach the margin.

Other species in eastern North America: Woodsia glabella R. Brown ex Richardson in Franklin, of circumboreal distribution, south in e. North America to NH, NY, VT. Woodsia alpina (Bolton) A. Gray of circumboreal distribution, south in e. North America to NH, NY, VT. Woodsia obtusa (Sprengel) Torrey ssp. occidentalis Windham of sc. United States, east to AR and MO. Woodsia oregana D.C. Eaton ssp. cathcartiana (B.L. Robinson) Windham, primarily of w. North America, but east to WI and disjunct in NY.

Other genera and species in eastern North America: Ctenitis sloanei (Poeppig ex Sprengel) C.V. Morton of s. FL. Ctenitis submarginalis (Langsdorff & Fischer) Ching of FL and LA. Lomariopsis kunzeana (C. Presl & L. Underwood) Holttum of s. FL. Maxonia apiifolia (Swartz) C. Christensen var. apiifolia of s. FL. Nephrolepis biserrata (Swartz) Schott of FL. Nephrolepis cordifolia (Linnaeus) C. Presl of FL. Nephrolepis exaltata (Linnaeus) Schott of FL. Nephrolepis multiflora (Roxburgh) F.M. Jarrett ex C.V. Morton of s. FL. Nephrolepis Haveryi Nauman of s. FL. Tectaria heracleifolia (Willdenow) L. Underwood of s. FL. Tectaria incisa Cavanilles of s. FL. Tectaria coriandrifolia (Swartz) L. Underwood of s. FL. Tectaria fimbriata (Willdenow) Proctor & Lourteig of s. FL.


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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.