FNA: Woodsia vs. Woodsia plummerae

Phenology

Sporulating late spring–fall.

Habitat

Cliffs and rocky slopes, usually on granite or volcanic substrates

Elevation

700–3100 m (2300–10200 ft)

Distribution

Mostly north temperate regions and higher elevations in the tropics

[BONAP county map]

AZ; CA; CO; NM; OK; TX; n Mexico

[WildflowerSearch map]

[BONAP county map]

Discussion

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 origin and phylogenetic affinities of the tetraploid Woodsia plummerae have not been established with certainty. The hypothesis that it arose as a hybrid between the Mexican species W. mollis (Kaulfuss) J. Smith and W. mexicana Fée (D. F. M. Brown 1964; D. B. Lellinger 1985) seems untenable in light of recent chromosome studies indicating that the latter species is also tetraploid (M. D. Windham 1993). On the basis of sporophyte morphology and spore ornamentation, W. plummerae appears most closely related to the W. mexicana complex and W. oregana. In fact, W. oregana can be difficult to separate from W. plummerae in western New Mexico and northern Arizona. Intermediate plants occurring in this region may represent stable allotetraploids resulting from hybridization between the diploid progenitors of W. plummerae and W. oregana subsp. cathcartiana. Considering the available evidence, populations of W. plummerae in the United States probably originated through autopolyploidy from a recently discovered, but as yet unnamed, Mexican diploid of similar morphology. Woodsia plummerae occasionally hybridizes with W. phillipsii to produce sterile, morphologically intermediate triploids.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key

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