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cliff-fern, mountain cliff fern, Rocky Mountain woodsia, woodsia, woodsie des rochers

Cochise cliff fern

Stems

compact, erect to ascending, with few to many persistent petiole bases of unequal lengths;

scales uniformly brown or bicolored with dark central stripe and pale brown margins, ovate to narrowly lanceolate.

compact, erect to ascending, with a few persistent petiole bases of unequal lengths;

scales often uniformly brown but at least some bicolored with dark central stripe and pale brown margins, narrowly lanceolate.

Leaves

9–35 × 1–8 cm.

5–25 × 1.5–6 cm.

Petiole

usually reddish brown to dark purple proximally when mature, not articulate above base, relatively brittle and easily shattered.

light brown or straw-colored throughout when mature, occasionally darker at very base, not articulate above base, relatively brittle and easily shattered.

Blade

lanceolate to linear-lanceolate, 2-pinnate proximally, moderately glandular, rarely somewhat viscid;

most glandular hairs with thick stalks and distinctly bulbous tips;

rachis usually with abundant glandular and nonglandular hairs.

narrowly lanceolate to ovate, pinnate-pinnatifid to 2-pinnate proximally, sparsely to moderately glandular, never viscid;

glandular hairs with thin stalks and slightly expanded tips;

rachis with glandular hairs and occasional hairlike scales.

Pinnae

lanceolate-deltate to ovate, longer than wide, abruptly tapered to a rounded or broadly acute apex, occasionally attenuate;

largest pinnae with 5–14 pairs of pinnules;

abaxial and adaxial surfaces glandular and sparsely villous, with flattened, multicellular hairs concentrated along midribs.

ovate-deltate to elliptic, longer than wide, abruptly tapered to a rounded or broadly acute apex, occasionally attenuate;

largest pinnae with 4–9 pairs of pinnules;

abaxial and adaxial surfaces glandular, lacking nonglandular hairs or scales.

Indusia

of filamentous or nonfilamentous segments, these multiseriate proximally, often uniseriate distally, composed of ± isodiametric cells, concealed by or slightly surpassing mature sporangia.

of relatively broad segments;

segments multiseriate most of length, usually divided and uniseriate distally, composed of ± isodiametric cells, often surpassing mature sporangia.

Spores

averaging 39–57 µm.

averaging 43–49 µm. 2n = 152.

Pinnules

dentate, often shallowly lobed;

margins nonlustrous, thin, slightly glandular and occasionally ciliate with isolated, multicellular hairs, lacking translucent projections.

dentate, often shallowly lobed;

margins lustrous adaxially, usually thickened, lacking cilia but sparsely glandular, with occasional 1–2-celled translucent projections.

Vein

tips slightly (if at all) enlarged, barely visible adaxially.

tips enlarged to form whitish hydathodes visible adaxially.

Woodsia scopulina

Woodsia cochisensis

Phenology Sporulating late spring–fall.
Habitat Shaded ledges and alcoves near springs and seeps, usually on granitic or volcanic substrates
Elevation 1000–2200 m (3300–7200 ft)
Distribution
from FNA
AK; AR; AZ; CA; CO; ID; KY; MT; NC; NV; OR; SD; TN; UT; VA; WA; WV; WY; AB; BC; ON; QC; SK; YT; only in the flora
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[BONAP county map]
from FNA
AZ; NM; n Mexico
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[BONAP county map]
Discussion

Woodsia scopulina shows substantial variation in leaf size, shape, and dissection, and in the abundance of multicellular hairs on the pinnae. Although much of this variation seems to be environmentally induced, recent studies (M. D. Windham 1993) have identified three chromosomal/morphologic variants that are treated here as subspecies. Diploid populations of W. scopulina are divisible into two groups, one of which (subsp. scopulina) is scattered throughout the mountainous regions of western North America while the other (subsp. appalachiana) is confined to montane habitats in the southeastern United States. These taxa seem amply distinct (T. M. C. Taylor 1947) and might be considered separate species if not for the existence of populations in the Great Lakes region and western cordillera that tend to bridge the morphologic and geographic gap between them. These intermediate populations (subsp. laurentiana) appear to be uniformly tetraploid and may have arisen through ancient hybridization between subsp. scopulina and subsp. appalachiana. In regions where subsp. laurentiana is sympatric with subsp. scopulina, the two taxa are rarely found growing together, suggesting that they differ in their ecological tolerances and/or habitat requirements.

Subspecies 3.

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

Woodsia cochisensis traditionally has been identified as W. plummerae or (rarely) W. mexicana Fée. It is readily separated from W. plummerae by the characteristics given in the key, and from North American members of the mexicana group (W. phillipsii and W. neomexicana) by having indusial segments that are broad and nonfilamentous at the base. Woodsia cochisensis is less glandular than typical W. mexicana from northeastern Mexico and is further distinguished from that species by the thickened, lustrous pinnule margins and well-developed hydathodes. Isozyme and chromosome studies suggest that W. cochisensis is an allotetraploid that may have originated through hybridization between W. phillipsii and an undescribed Mexican diploid (M. D. Windham 1993). It crosses with the former species to produce sterile triploids of intermediate morphology.

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

Key
1. Scales of stems and petiole bases narrowly lanceolate, mostly bicolored with broad, usually continuous, dark central stripe; longest hairs on pinnae composed of 5-8 cells; indusial segments broad, not at all filamentous.
subsp. appalachiana
1. Scales of stem and petiole bases ovate-lanceolate, mostly concolored or weakly bicolored with narrow, often discontinuous, dark central stripe; longest hairs on pinnae composed of 2-5 cells; indusial segments narrow, often filamentous distally.
→ 2
2. Spores averaging 42-50 µm; stem and petiole base scales usually concolored or with a few isolated, dark, occluded cells.
subsp. scopulina
2. Spores averaging 50-57 µm; at least some stem and petiole base scales with clusters of dark, occluded cells near center forming narrow, usually discontinuous stripe.
subsp. laurentiana
Source FNA vol. 2. FNA vol. 2.
Parent taxa Dryopteridaceae > Woodsia Dryopteridaceae > Woodsia
Sibling taxa
W. alpina, W. cochisensis, W. glabella, W. ilvensis, W. neomexicana, W. obtusa, W. oregana, W. phillipsii, W. plummerae
W. alpina, W. glabella, W. ilvensis, W. neomexicana, W. obtusa, W. oregana, W. phillipsii, W. plummerae, W. scopulina
Subordinate taxa
W. scopulina subsp. appalachiana, W. scopulina subsp. laurentiana, W. scopulina subsp. scopulina
Synonyms W. obtusa var. lyallii, W. oregana var. lyallii
Name authority D. C. Eaton: Canad. Naturalist & Quart. J. Sci. 2: 91. (1865) Windham: Contr. Univ. Michigan Herb. 19: 54. (1993)
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