Woodsia oregana |
Woodsia cochisensis |
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cliff fern, Oregon cliff-fern, Oregon woodsia, western cliff fern, woodsie de l'Oregon |
Cochise cliff fern |
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Stems | compact, erect to ascending, with few to many 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. |
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. |
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Leaves | 4–25 × 1–4 cm. |
5–25 × 1.5–6 cm. |
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Petiole | reddish brown to dark purple proximally when mature, not articulate above base, somewhat pliable and resistant to shattering. |
light brown or straw-colored throughout when mature, occasionally darker at very base, not articulate above base, relatively brittle and easily shattered. |
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Blade | linear-lanceolate to narrowly ovate, pinnate-pinnatifid or 2-pinnate proximally, sparsely to moderately glandular, never viscid; glandular hairs with thin stalks and slightly expanded tips; rachis with scattered glandular hairs and occasional hairlike scales. |
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. |
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Pinnae | ovate-deltate to elliptic, longer than wide, abruptly tapered to a rounded or broadly acute apex; largest pinnae with 3–9 pairs of pinnules; abaxial and adaxial surfaces glabrescent to moderately glandular, lacking nonglandular hairs or scales. |
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. |
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Indusia | of narrow, usually filamentous segments, these uniseriate for most of length, 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. |
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Spores | averaging 39–50 µm. |
averaging 43–49 µm. 2n = 152. |
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Pinnules | dentate, often shallowly lobed; margins nonlustrous, thin, with occasional glands, lacking cilia, rarely with 1–2-celled translucent projections. |
dentate, often shallowly lobed; margins lustrous adaxially, usually thickened, lacking cilia but sparsely glandular, with occasional 1–2-celled translucent projections. |
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Vein | tips slightly (if at all) enlarged, barely visible adaxially. |
tips enlarged to form whitish hydathodes visible adaxially. |
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Woodsia oregana |
Woodsia cochisensis |
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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 |
AZ; CA; CO; IA; ID; KS; MI; MN; MT; ND; NE; NM; NV; NY; OK; OR; SD; UT; WA; WI; WY; AB; BC; MB; ON; QC; SK; only in the flora
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AZ; NM; n Mexico
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Discussion | The variability and promiscuity of Woodsia oregana have been major sources of taxonomic difficulties in Woodsia, and more work will be necessary before relationships in this complex are fully resolved. As defined here, W. oregana comprises two subspecies that are chromosomally and biochemically distinct. In addition, the two taxa are nearly allopatric, with the diploid (subsp. oregana) confined to the Pacific Northwest and the tetraploid (subsp. cathcartiana) extending from the southwestern United States to eastern Canada. Isozyme studies indicate that subsp. cathcartiana is not an autotetraploid derived from known diploid populations of subsp. oregana, as was hypothesized by D. F. M. Brown (1964), and it may be more appropriate to recognize these taxa as distinct species. The morphologic features that distinguish these subspecies are very subtle, however, and they are associated primarily with differences in chromosome number. Until further systematic analyses are undertaken, these cytotypes should be maintained as subspecies of W. oregana. Subspecies 2. (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.) |
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Key |
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Source | FNA vol. 2. | FNA vol. 2. | ||||
Parent taxa | ||||||
Sibling taxa | ||||||
Subordinate taxa | ||||||
Name authority | D. C. Eaton: Canad. Naturalist & Quart. J. Sci. n. s. 2: 90. (1865) | Windham: Contr. Univ. Michigan Herb. 19: 54. (1993) | ||||
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