Polystichum californicum |
Dryopteridaceae |
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California sword-fern |
wood fern family |
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Habit | Plants perennial, terrestrial or on rock, occasionally hemiepiphytic or epiphytic. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Stems | erect or ascending. |
creeping to erect, rarely arborescent, sometimes climbing, branched or unbranched, dictyostelic, bearing scales. |
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Leaves | monomorphic, arching or erect, 2–8 dm; bulblets absent. |
circinate in bud, monomorphic or dimorphic. |
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Petiole | 1/5–1/3 length of leaf; scales light brown, abruptly diminishing in size distally, falling off early distally. |
usually not articulate to stem, scales usually persistent at base, in cross section with 2–many roundish bundles, or bundles 2 and lunate. |
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Blade | lanceolate to linear-lanceolate, 1-pinnate-pinnatifid, base slightly narrowed. |
simple to commonly 1–5-pinnate or more divided, leaf buds absent or present. |
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Pinnae | oblong to lanceolate to falcate, shallowly to deeply divided, pinnae overlapping or not, in 1 plane, 2–10 cm; base oblique, acroscopic auricle lobed; margins not incised to costae, serrulate-spiny with teeth ascending; apex acute-attenuate, subapical and apical teeth same size (southern form) or obtuse and cuspidate with subapical teeth smaller than apical teeth (northern form); microscales filiform, dense abaxially, sparse adaxially. |
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Veins | pinnate or parallel in ultimate segments, simple or forked, free or anastomosing, areoles sometimes with included free veinlets. |
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Indusia | ciliate. |
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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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Sporangia | with stalk of 2–3 rows of cells; annulus vertical, interrupted by stalk. |
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Spores | brown. |
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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Gametophytes | green, aboveground, cordate, glabrous or often bearing glands or hairs; archegonia and antheridia borne on lower surface, antheridia 3-celled. |
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Indument | on blade commonly of glands, hairs, and/or scales, especially on rachis and costae abaxially. |
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2n | = 164. |
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Polystichum californicum |
Dryopteridaceae |
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Habitat | On forest floor in southern part of range and in rock crevices at cliff bottoms (most commonly andesite) to north | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Elevation | 100–850 m (300–2800 ft) | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Distribution |
CA; OR; WA; BC
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Worldwide |
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Discussion | Polystichum californicum is restricted to the Coast Ranges and the Sierra-Cascade axis. It is most abundant in the Coast Range north of San Francisco. Polystichum californicum is an allopolyploid, the evolutionary roots of which include P. dudleyi as the 2-pinnate ancestor. Morphologic and ecological data indicate P. imbricans is ancestor to the northern forms and P. munitum is ancestor to southern forms, suggesting P. californicum is an amalgam of interfertile tetraploids with polyphyletic origins (D. H. Wagner 1979). Cytological analysis corroborates this (A. D. Callan 1972; W. H. Wagner Jr. 1973), but chloroplast DNA studies have detected only the involvement of P. imbricans in the ancestry of P. californicum (P. S. Soltis et al. 1991). The more xeric, rock-inhabiting members of the complex (showing the parental influence of P. imbricans) occupy the northern half of the range whereas plants of more mesic habitats are found to the south. Hybrids with both P. dudleyi and P. munitum are found frequently, because these three species are often sympatric (W. H. Wagner 1973). The hybrid with P. dudleyi (a triploid) will key to that species. The hybrid with P. munitum resembles a less-incised form of P. californicum with aborted sporangia. Polystichum californicum × imbricans has been found only once, in Oregon (A. D. Callan 1972). Another hybrid that will key here, based on its overall appearance, is P. munitum × scopulinum. It lacks filiform microscales and also has malformed sporangia. Such a specimen was the basis of the report of Polystichum californicum in eastern Washington (C. L. Hitchcock et al. 1955–1969, vol. 1). The sterile diploid hybrid between P. dudleyi and P. munitum is indistinguishable from P. californicum except for aborted sporangia and chromosome number (W. H. Wagner Jr. 1973). (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.) |
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Key |
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Source | FNA vol. 2. | FNA vol. 2, p. 246. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Parent taxa | Dryopteridaceae > Polystichum | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Sibling taxa | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Synonyms | Aspidium californicum, P. aculeatum var. californicum | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Name authority | (D. C. Eaton) Diels: in Engler & Prantl, Nat. Pflanzenfam. 1(4): 191. (1899) | Herter | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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