cliff-fern, woodsia
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wood fern family
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Plants usually on rock. |
Plants perennial, terrestrial or on rock, occasionally hemiepiphytic or epiphytic. |
compact to creeping; ascending or erect (rarely horizontal), stolons absent. |
creeping to erect, rarely arborescent, sometimes climbing, branched or unbranched, dictyostelic, bearing scales. |
monomorphic, dying back over winter or sometimes persistent into the next season. |
circinate in bud, monomorphic or dimorphic. |
1/5–3/4 length of blade, base not conspicuously swollen; vascular bundles 2, arranged laterally, ± round or oblong in cross section. |
usually not articulate to stem, scales usually persistent at base, in cross section with 2–many roundish bundles, or bundles 2 and lunate. |
linear to lanceolate or ovate, 1–2-pinnate-pinnatifid, gradually reduced distally to pinnatifid apex, herbaceous. |
simple to commonly 1–5-pinnate or more divided, leaf buds absent or present. |
not articulate to rachis, segment margins entire to dentate, not spiny; proximal pinnae somewhat reduced, sessile, bases usually ± equilateral; costae often shallowly grooved adaxially, grooves ± continuous from rachis to costae; indument of glandular (occasionally nonglandular) hairs on both surfaces, rarely absent. |
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free, simple or forked. |
pinnate or parallel in ultimate segments, simple or forked, free or anastomosing, areoles sometimes with included free veinlets. |
in 1 row between midrib and margin on ultimate segments, round; indusia basal, dissected into several to numerous filamentous or scalelike segments encircling sorus, persistent but often obscure in mature sori. |
borne abaxially on veins or at vein tips (but usually not marginal), or sporangia acrostichoid and covering abaxial surface, if in discrete sori then variously shaped (round, oblong, or elongate); receptacle not or only slightly elevated, with or without indusium, indusium variously linear, falcate, or reniform, sometimes hoodlike, cuplike, or round. |
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with stalk of 2–3 rows of cells; annulus vertical, interrupted by stalk. |
brownish, cristate, rarely rugose. |
all of 1 kind, usually not green (except Matteuccia, Onoclea), oblong or reniform in outline, monolete, variously ornamented (often broadly winged), 64 per sporangium (32 in apogamous spp.). |
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green, aboveground, cordate, glabrous or often bearing glands or hairs; archegonia and antheridia borne on lower surface, antheridia 3-celled. |
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on blade commonly of glands, hairs, and/or scales, especially on rachis and costae abaxially. |
= 38, 39, 41. |
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Mostly north temperate regions and higher elevations in the tropics |
Worldwide |
Woodsia is a well-marked genus; its morphology and chromosome base number (x = 41) provide evidence of relationships to the dryopteroid ferns. Most authors consider Cystopteris to be its closest ally, and the two genera are often confused in herbarium collections. The resemblance is superficial in many ways, however, and Woodsia is easily distinguished from Cystopteris by its persistent petiole bases, multilobed indusia, and obscure veins that end in hydathodes before reaching the leaf margin. The North American species of Woodsia fall into two natural groups that might be recognized as subgenera. Woodsia ilvensis, W. glabella, and W. alpina have articulate petioles, indusial segments that are uniseriate throughout and composed of cells that are much longer than wide, entire or crenate pinnules, strictly concolored stem scales, and chromosome base numbers of 39–41. They are circumboreal in distribution and show clear affinities to species found only in Eurasia. The remainder of the North American taxa have petioles that are not articulate, indusial segments that are multiseriate at the base and composed of cells that are isodiametric or slightly longer than wide, dentate pinnules, often bicolored stem scales, and a chromosome base number of 38. All of these species are endemic to the New World and probably represent a distinct lineage within the genus. Hybridization is common within these natural groups, but intergroup hybrids are relatively rare. Species ca. 30 (10 in the flora). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
The family Dryopteridaceae has been variously circumscribed; it is here delimited in a manner similar to that of R. M. Tryon and A. F. Tryon (1982) but with the inclusion of Nephrolepis. In many works, the family has gone under the illegitimate name Aspidiaceae. Some authorities define Dryopteridaceae more narrowly, to exclude Athyrium, Deparia, Diplazium, Cystopteris, and Gymnocarpium (Athyriaceae or Woodsiaceae), Woodsia (Woodsiaceae), Lomariopsis (Lomariopsidaceae), Nephrolepis (Nephrolepidaceae or Davalliaceae), Onoclea and Matteuccia (Onocleaceae), and Ctenitis and Tectaria (Tectariaceae). Characteristics holding Dryopteridaceae (as circumscribed here) together include the bilateral, monolete spores, often broadly winged perispore, absence of needlelike hairs, scaly stem and petiole bases, abaxial (nonmarginal) sori, base chromosome number of 40 or 41 (also 38 and 39 in Woodsia, 37 in Onoclea, 42 in Cystopteris), and usually indusiate sori. Loss of indusium, dimorphism, areolate venation, and reduced blade dissection have occurred repeatedly along many evolutionary lines in Dryopteridaceae, and in general these characteristics are often not very useful in delimiting genera or assessing intergeneric relationships. In some genera, especially Phanerophlebia and Polystichum, the blade bears very narrow scales (sometimes called microscales) that resemble uniseriate hairs. These scales may be only one or two cells wide. Every intergradation exists between these filiform microscales and more typical, wider scales, and the two types are the same color, generally tan to brownish. Microscales are probably not homologous with true hairs, which may be either unicellular or multicellular, uncolored or sometimes reddish (as in Tectaria and Ctenitis), glandular (as in Woodsia) or not. Hairs in Dryopteridaceae, if present at all, are generally readily distinguishable from the needlelike, transparent ones found in Thelypteridaceae. Genera ca. 60, species perhaps exceeding 3000 (18 genera, 79 species in the flora). (Discussion copyrighted by Flora of North America; reprinted with permission.) |
1. Blades and rachises completely glabrous except for occasional sessile glands; proximal pinnae distinctly fan-shaped, usually wider than long; leaves 0.5-1.2 cm wide, mature petioles green or straw-colored throughout. | W. glabella |
1. Blades and/or rachises with scattered hairs, scales, or stalked glands, rarely glabrescent; proximal pinnae ovate-lanceolate to deltate, usually longer than wide; leaves 1.2-12 cm wide or, if less, then proximal portion of mature petioles reddish brown or dark purple. | → 2 |
2. Petioles articulate well above base, abscission zone visible as swollen node; indusial segments uniseriate throughout, composed of cells that are many times longer than wide; pinnules entire or crenate, without acute teeth on margins. | → 3 |
2. Petioles not articulate above base, swollen abscission zone absent; indusial segments usually multiseriate at base, composed of cells that are isodiametric or slightly longer than wide; pinnules dentate, with acute teeth on margins. | → 4 |
3. Linear-lanceolate scales absent or very rare on abaxial pinnae surfaces; rachises with widely scattered hairs and scales, sometimes nearly glabrous; largest pinnae with 1-3 pairs of pinnules. | W. alpina |
3. Linear-lanceolate scales common on abaxial pinnae surfaces; rachises with abundant hairs and scales; largest pinnae with 4-9 pairs of pinnules. | W. ilvensis |
4. Pinnae with flattened, multicellular hairs concentrated along midrib on both surfaces; mature petioles usually reddish brown or dark purple, relatively brittle and easily shattered. | W. scopulina |
4. Pinnae lacking flattened, multicellular hairs along midrib; mature petioles light brown to straw-colored or, if reddish brown/dark purple (in Woodsia plummerae and W. oregana), somewhat pliable and resistant to shattering. | → 5 |
5. Indusia composed of relatively broad segments, these multiseriate for most of length but often branched or divided distally. | → 6 |
5. Indusia composed of narrow, usually filamentous segments, these uniseriate for most of length. | → 8 |
6. Proximal portion of mature petioles reddish brown or dark purple; blades densely glandular, often somewhat viscid; vein tips not enlarged, barely visible on adaxial surface. | W. plummerae |
6. Proximal portion of mature petioles light brown or straw-colored (sometimes darker at very base); blades sparsely to moderately glandular, rarely viscid; vein tips usually enlarged to form whitish hydathodes visible on adaxial surface. | → 7 |
7. Indusial segments branched or divided distally to form narrow, filamentous lobes; glandular hairs of blade with thin stalks and slightly expanded tips; pinnule margins usually thickened, lustrous on adaxial surface. | W. cochisensis |
7. Indusial segments often glandular along distal edge but otherwise nearly entire, not divided into narrow, filamentous lobes; many glandular hairs of blade with thick stalks and distinctly bulbous tips; pinnule margins not noticeably thickened or lustrous. | W. obtusa |
8. Pinnule margins (viewed from abaxial surface) smooth to somewhat ragged but usually lacking translucent projections or filaments; proximal portion of mature petioles reddish brown or dark purple; indusial filaments generally inconspicuous, concealed by or slightly surpassing mature sporangia. | W. oregana |
8. Pinnule margins (viewed from abaxial surface) with translucent projections or filaments on teeth; proximal portion of mature petioles usually light brown or straw-colored (sometimes darker at very base); indusial filaments generally apparent, often greatly surpassing mature sporangia. | → 9 |
9. Translucent projections on pinnule margins mostly 1-2-celled, occasionally filamentous; spores averaging 44-52 µm; largest pinnae divided into 3-7 pairs of closely spaced pinnules, pinna apices usually abruptly tapered to rounded. | W. neomexicana |
9. Translucent projections on pinnule margins mostly multicellular, often prolonged to form twisted filaments; spores averaging 37-44 µm; largest pinnae with 7-18 pairs of discrete, widely spaced pinnules, pinna apices often attenuate to narrowly acute. | W. phillipsii |
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1. Leaves strongly dimorphic, either fertile or sterile, the 2 types very dissimilar. | → 2 |
1. Leaves monomorphic, fertile and sterile similar in size and dissection, occasionally with somewhat contracted fertile pinnae on same leaf as sterile pinnae (as in Polystichum acrostichoides). | → 5 |
2. Sporangia completely covering abaxial surface of blade, not in discrete sori or hidden by revolute segment margins; sterile blades 1-pinnate, pinnae serrate. | Lomariopsis |
2. Sporangia not covering abaxial surface of blade, in discrete sori or hidden by revolute segment margins; sterile blades variously divided, not 1-pinnate with pinnae serrate. | → 3 |
3. Plants hemiepiphytic, rooted in ground and with stems climbing trees; sterile blades 3-4-pinnate; indusia thick, conspicuous. | Maxonia |
3. Plants terrestrial, not climbing; sterile blades deeply pinnatifid to 1-pinnate-pinnatifid; indusia thin, fragile, hidden by revolute segment margins. | → 4 |
4. Sterile blades pinnatifid to 1-pinnate at base; venation areolate; fertile blades 2-pinnate, sori enclosed in small, globose, hardened pinnules. | Onoclea |
4. Sterile blades 1-pinnate-pinnatifid; veins free; fertile blades 1-pinnate, sori on linear pinnae and enclosed by hardened pinna margin. | Matteuccia |
5. Stolons present, wiry, arising from stem; blades 1-pinnate; pinnae articulate to rachis, sometimes deciduous with age; indusia lunate to reniform or circular with narrow sinus. | Nephrolepis |
5. Stolons absent; blades variously divided; pinnae not articulate to rachis, or rarely proximal pinnae weakly articulate but not deciduous; indusia various or absent. | → 6 |
6. Indusia completely surrounding receptacle and composed of filaments or scalelike segments arranged in cuplike fashion around sorus; petiole base with 2 vascular bundles; scales absent on costae abaxially. | Woodsia |
6. Indusia attached centrally or laterally, not completely surrounding receptacle, or indusia absent; petiole base with 2 or more vascular bundles; scales absent or present on costae abaxially. | → 7 |
7. Veins areolate or copiously anastomosing. | → 8 |
7. Veins free or only casually and sparingly anastomosing. | → 10 |
8. Sori and indusia linear; petiole base with 2 vascular bundles. | Diplazium |
8. Sori round, with indusia round or round-reniform; petiole base with many vascular bundles. | → 9 |
9. Costae adaxially rounded or flattened, bearing multicellular reddish hairs; margins of pinnae lacking spinules or teeth. | Tectaria |
9. Costae adaxially grooved, lacking reddish multicellular hairs but sometimes with tan, very reduced, filiform scales; margins of pinnae spinulose to denticulate or crenate. | Cyrtomium |
10. Indusia round, attached at center (peltate); sori round; petiole base with 3 or more vascular bundles. | → 11 |
10. Indusia round-reniform, reniform, linear, or absent, attached laterally at sinus; sori round or elongate; petiole base with 2 or more vascular bundles. | → 12 |
11. Blades 1-pinnate with terminal pinna similar to lateral pinnae; sori in 2-4 rows between costa and margin; s Arizona to w Texas. | Phanerophlebia |
11. Blades 1-3-pinnate with gradually reduced and pinnatifid apex; sori in 1(-2) rows between costa and margin or between costule and margin; widespread. | Polystichum |
12. Sori elongate, straight or hooked at one end, indusiate; petiole base with 2 vascular bundles. | → 13 |
12. Sori round or nearly so, indusia present or absent; petiole with 2 or more vascular bundles. | → 15 |
13. Adaxial grooves of costae shallow, not decurrent into rachis groove; multicellular hairs borne along costae, especially adaxially; stems moderately long-creeping; blades 1-pinnate-pinnatifid. | Deparia |
13. Adaxial grooves of costae deep, decurrent into rachis groove; multicellular hairs absent on costae; stems short-creeping to erect; blades 1-pinnate to 2-pinnate-pinnatifid. | → 14 |
14. Blades commonly 2-pinnate or more divided; proximal pinnae often slightly to greatly reduced; sori usually hooked at distal end. | Athyrium |
14. Blades 1-pinnate, 1-pinnate-pinnatifid, or 2-pinnate (if 2-pinnate then veins anastomosing); proximal pinnae not or slightly reduced; sori ± straight, not hooked at distal end. | Diplazium |
15. Costae rounded or flat adaxially, bearing dense, obviously multicellular hairs with reddish crosswalls. | Ctenitis |
15. Costae grooved adaxially, lacking hairs. | → 16 |
16. Indusia attached at distinct sinus, round-reniform; petiole base with 3 or more vascular bundles. | → 17 |
16. Indusia absent or laterally attached and hoodlike, arching over sori; petiole base with 2 vascular bundles. | → 18 |
17. Stems short-creeping, nearly erect or erect; blades lanceolate to ovate, not pentagonal; widespread. | Dryopteris |
17. Stems moderately long-creeping; blades pentagonal, with basal basiscopic pinnules decidedly longer than next pair; South Carolina. | Arachniodes |
18. Stems long-creeping; blades deltate to pentagonal, proximal pinnae by far the largest; petioles mostly 1/3-3 times length of blades. | → 19 |
18. Stems short-creeping to ascending; blades ovate to lanceolate, proximal pinnae occasionally slightly longer than adjacent pair; petioles mostly shorter than blades. | → 20 |
19. Indusia present but often inconspicuous in mature leaves, laterally attached and arching over sori; segment margins serrate-dentate (Cystopteris montana). | Cystopteris |
19. Indusia always absent; segment margins entire or crenate but never serrate-dentate. | Gymnocarpium |
20. Indusia absent from all sori; stems erect to ascending, surface obscured by petiole bases and scales, scales more than 5 mm. | Athyrium |
20. Indusia present but often inconspicuous in mature leaves, laterally attached and arching over sori; stems decumbent, usually creeping, surface often visible through petiole bases and scales, scales less than 5 mm. | Cystopteris |
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FNA vol. 2. Author: Michael D. Windham. |
FNA vol. 2, p. 246. Author: Alan R. Smith. |
Dryopteridaceae |
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Arachniodes, Athyrium, Ctenitis, Cyrtomium, Cystopteris, Deparia, Diplazium, Dryopteris, Gymnocarpium, Lomariopsis, Matteuccia, Maxonia, Nephrolepis, Onoclea, Phanerophlebia, Polystichum, Tectaria, Woodsia |
R. Brown: Prodr. 158. (1810) |
Herter |
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