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Eaton's holly fern, mountain holly fern, polystic des rochers, rock sword fern

California sword-fern

Stems

ascending.

erect or ascending.

Leaves

erect, 1–3(–5) dm;

bulblets absent.

monomorphic, arching or erect, 2–8 dm;

bulblets absent.

Petiole

1/5–1/3 length of leaf, densely scaly but scales falling off distally;

scales light brown, abruptly diminishing in size distally.

1/5–1/3 length of leaf;

scales light brown, abruptly diminishing in size distally, falling off early distally.

Blade

narrowly lanceolate, 1-pinnate-pinnatifid, base narrowed.

lanceolate to linear-lanceolate, 1-pinnate-pinnatifid, base slightly narrowed.

Pinnae

oblong-lanceolate, overlapping, folded inward and twisted horizontally, 1–3 cm;

base oblique;

margins serrulate with teeth curved inward;

apex obtuse to cuspidate with subapical teeth smaller than apical tooth;

microscales narrowly lanceolate, with stout projections, sparse, on abaxial surface only.

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.

Indusia

entire-ciliate.

ciliate.

Spores

brown.

brown.

2n

= 164.

= 164.

Polystichum scopulinum

Polystichum californicum

Habitat Rock crevices and at base of boulders, serpentine to acidic substrates, usually exposed to full sun On forest floor in southern part of range and in rock crevices at cliff bottoms (most commonly andesite) to north
Elevation 0–3500 m (0–11500 ft) 100–850 m (300–2800 ft)
Distribution
from FNA
AZ; CA; CO; ID; MT; NV; OR; UT; WA; WY; BC; NF; QC
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from FNA
CA; OR; WA; BC
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[BONAP county map]
Discussion

Polystichum scopulinum is widely distributed in the United States west of the 110th meridian, where it occurs in sporadic, usually small populations. The species is abundant only on montane serpentine outcrops. The populations in Newfoundland and Quebec are dramatically disjunct.

Polystichum scopulinum is an allopolyploid, believed on morphologic grounds to be derived from P. imbricans × lemmonii (D. H. Wagner 1979). Based on putative hybridization between P. scopulinum and P. munitum (P. S. Soltis et al. 1989; W. H. Wagner Jr. 1973), however, P. munitum may also be involved. This hybrid is discussed under P. californicum.

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

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.)

Source FNA vol. 2. FNA vol. 2.
Parent taxa Dryopteridaceae > Polystichum Dryopteridaceae > Polystichum
Sibling taxa
P. acrostichoides, P. aleuticum, P. andersonii, P. braunii, P. californicum, P. dudleyi, P. imbricans, P. kruckebergii, P. kwakiutlii, P. lemmonii, P. lonchitis, P. microchlamys, P. munitum, P. setigerum
P. acrostichoides, P. aleuticum, P. andersonii, P. braunii, P. dudleyi, P. imbricans, P. kruckebergii, P. kwakiutlii, P. lemmonii, P. lonchitis, P. microchlamys, P. munitum, P. scopulinum, P. setigerum
Synonyms Aspidium aculeatum var. scopulinum, P. mohrioides var. scopulinum Aspidium californicum, P. aculeatum var. californicum
Name authority (D. C. Eaton) Maxon: Fern Bull. 8: 29. (1900) (D. C. Eaton) Diels: in Engler & Prantl, Nat. Pflanzenfam. 1(4): 191. (1899)
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