Common Names in English:
Coast Polypody, Leathery Polypody
, on rock, or often epiphytic, erect
, or occasionally pendent. Stems long- to short-creeping, branched or not, bearing scales
and few to numerous
, usually dictyostelic. Leaves monomorphic
in bud. Petiole
, as in Loxogramme ], lacking scales or sometimes scaly
, with usually 3 vascular bundles
. Blade simple
to often pinnatifid
, or pinnate, infrequently more divided
; rachis grooved
or not adaxially. Veins free
(and simple to several times forked
) to often anastomosing in complex
, areoles with or without included
. Indument on blade
absent, or petiole, rachis, costae, and sometimes blade tissue usually bearing hairs
(these often septate
and with reddish crosswalls) and/or scales. Sori borne abaxially on veins, round
, occasionally elongate
, rarely marginal
, rarely covering surface; paraphyses present or absent; sporangia with stalk
of 2 or 3 rows
; indusia absent. Spores usually transparent or yellowish (rarely greenish), all 1 kind, bilateral
, monolete [rarely trilete, as in some Loxogramme], surface most often smooth
, or granulate
, occasionally spiny
, 64 per sporangium (spores globose
and 32 per sporangium in apogamous spp.
) . Gametophytes green, aboveground, cordate or elliptic
or sometimes glandular
; archegonia and antheridia borne on lower surface, antheridia 3-celled.
Genera ca. 40, species perhaps 500 (7 genera, 25 species in the flora ) : worldwide, especially tropics and subtropics.
Genera in this family are variously circumscribed, and the New World species historically were placed in the single genus Polypodium . Many of the segregates recognized here are still placed in Polypodium in recent floristic accounts. Limits of genera in both Old World and New World are controversial and are currently under study by several workers.
on rock, occasionally terrestrial
or epiphytic. Stems creeping
, usually branched, 3--15 mm diam., sometimes whitish pruinose
concolored to bicolored
, lanceolate to ovate-acuminate, not clathrate to strongly clathrate, glabrous
entire to denticulate
. Leaves monomorphic
, closely spaced to distant
, not conspicuously narrowed at tip
, to 90 cm. Petiole
to stem, straw-colored, somewhat flattened or grooved
to nearly terete
to deltate, pinnatifid
to 1-pinnate at base
, not pectinate
, usually with fewer than 25 pairs of pinnae, not glaucous or conspicuously scaly
; rachis sparsely scaly to glabrescent
to glabrous adaxially; scales ovate-lanceolate to linear
, not peltate or clathrate. Segments linear to oblong
; margins entire to serrate; apex rounded
to attenuate. Venation
to anastomosing, if strongly anastomosing, then never with more than 1 included
areoles. Sori often confined to distal region of leaf, discrete, circular to oval
, borne at tips of single veins, in 1--3 rows
on either side of midrib
absent or of modified sporangia (sporangiasters), often bearing glandular
on bulbous head
. Spores monolete, rugose
Species ca. 100: worldwide.
Some species traditionally included in Polypodium are treated here in other genera, for example, Pleopeltis and Pecluma.
Except for the tropical species Polypodium triseriale, North American Polypodium is a complex assemblage of interactive species. The North American species have ties to European taxa (e.g. , P . vulgare sensu stricto , which probably originated by allopolyploidy between P . glycyrrhiza and P . sibiricum ) but are quite distinct from them. Morphologic comparisons and continuing biochemical and molecular studies indicate that two groups of diploid species occur within the North American P . vulgare complex. One group includes P . glycyrrhiza and P . californicum ; the second, P . amorphum, P . appalachianum, and P . sibiricum . Allopolyploid species have originated following hybridizations within a species group (i.e. , P . calirhiza from P . glycyrrhiza × californicum, P . saximontanum from P . amorphum × sibiricum, and P . virginianum from P . appalachianum × sibiricum ) as well as between members of the two groups (i.e., P . hesperium from P . amorphum × glycyrrhiza ) . These reticulate relationships are summarized in the reticulogram. We consider P . scouleri to be peripheral to the "core" diploids even though hybrids have been reported.
Species Polypodium scouleri
Stems conspicuously whitish pruinose
, stout, 3--12 mm diam., bland
to slightly sweet-tasting; scales
concolored to weakly bicolored
uniformly dark brown or with pale
, lanceolate, symmetric
. Leaves to 85 cm. Petiole
stout, to 3 mm diam.
, usually widest just above base,
to 27 cm wide, stiff and leathery; rachis sparsely scaly
adaxially; scales bicolored, ovate-lanceolate,
much more than 6 cells
wide. Segments oblong
, usually more
than 12 mm wide; margins entire to crenulate
; apex rounded
broadly acute; midrib
glabrous adaxially. Venation
usually forming 1 row
of areoles. Sori crowded against midrib, usually
more than 3 mm diam., circular when immature
. Sporangiasters absent.
Spores usually less than 52 µm, rugose
, surface projections
less than 3 µm tall. 2 n = 74, 111. Sporulating late fall--spring.
The distinctive Polypodium scouleri has occasionally been assigned to the genus Goniophlebium because of its anastomosing venation and conspicuous areoles. Its venation pattern can be quite variable, however, and cannot be used as the sole feature distinguishing P. scouleri from P. californicum. Combining venation characteristics with others provided in the key distinguishes it clearly from its congeners in Polypodium. Some evidence suggests that P. scouleri hybridizes with P. californicum (S. A. Whitmore, unpubl.). I. Manton (1951) reported diploid and triploid cytotypes for P. scouleri, and variation in spore size suggests that the species may also include tetraploid populations. [source]
Flowers: Flower Color: inconspicuous, none
Size: 6-12" tall.
Typically found at an altitude of 0 to 1,265 meters (0 to 4,150 feet).
Sunlight: Sun Exposure: Partial shade to full shade.
Temperature: Cold Hardiness: 8a, 8b, 9a, 9b. (map)
- Whittaker & Margulis,1978
- Haeckel, 1866
- Cavalier-Smith, 1981
- Sinnott, 1935 ex Cavalier-Smith, 1998
- Vascular Plants
- Kenrick & Crane, 1997, Nom. Nud.
- Cronquist et al.
- Berchtold & J. Presl, 1820
- common ferns, licorice ferns
- Subfamily: Polypodioideae ()
- Family: Polypodiaceae () - Berchtold & J. Presl, 1820 - common ferns, licorice ferns
- Order: Polypodiales () - Link
- Class: Polypodiopsida () - Cronquist et al.
- Infraphylum: Moniliformopses () - Kenrick & Crane, 1997, Nom. Nud.
- Subphylum: Euphyllophytina ()
- Phylum: Tracheophyta () - Sinnott, 1935 ex Cavalier-Smith, 1998 - Vascular Plants
- Subkingdom: Viridaeplantae () - Cavalier-Smith, 1981
- Kingdom: Plantae () - Haeckel, 1866 - Plants
Status: Accepted Name
Last scrutiny: 15-Mar-2000
Members of the genus Polypodium
ZipcodeZoo has pages for 37 species, subspecies, varieties, forms, and cultivars in this genus:
P. amorphum (Irregular Polypody) · P. appalachianum (Appalachian Polypody) · P. attenuatum (Tapered Polypody) · P. bulbiferum (Bulblet Bladderfern) · P. californicum (California Polypody) · P. calirhiza (Nested Polypody) · P. cambricum (Welsh Polypody) · P. carthusianum (Spinulose Wood Fern) · P. dissimile (Forkvein Polypody) · P. dulce (Treetrunk Polypody) · P. exaltatum (Sword Fern) · P. falax (Crested Poly) · P. formosanum (Catepillar Fern) · P. glycyrrhiza (Licorice Fern) · P. hesperium (Western Licorice Fern) · P. hexagonopterum (Broad Beech Fern) · P. ilvense (Rusty Woodsia) · P. interjectum (Polypody) · P. lanceolatum (Lanceleaf Polypody) · P. lonchitis (Northern Holly Fern) · P. loriceum (Clambering Polypody) · P. musifolium (Polypodium) · P. pellucidum (Ae) · P. pellucidum Kaulf. var. vulcanicum Skottsb. (Dotted Polypody) · P. pellucidum var. pellucidum (Dotted Polypody) · P. pellucidum var. vulcanicum (Dotted Polypody) · P. punctatum 'Grandiceps Cobra' (Climbing Bird's Nest Fern) · P. robertianum (Limestone Oak Fern) · P. saximontanum (Rocky Mountain Polypody) · P. scouleri (Coast Polypody) · P. sibiricum (Siberian Polypody) · P. subauriculatum var. knightii (Polypodium) · P. submarginale (Brownhair Lacefern) · P. triseriale (Angle-Vein Fern) · P. virginianum (Common Polypody) · P. virginianum var. peraferens (American Wall Fern) · P. 'Green Wave' (Polypodium)
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- Bulletin of the Southern California Academy of Sciences. Los Angeles, Calif.: The Academy, 1902-1971. url p. 47.
- Contributions from the United States National Herbarium 11 1906 Washington, D.C.: Smithsonian Institution Press, 1890- url p. 46, p. 76, p. 77.
- Ferns and fern allies of Canada / [Ottawa]: Research Branch, Agriculture Canada, 1989. url p. 295, p. 296.
- Flora of the Queen Charlotte Islands. [Ottawa: Queen's Printer], 1968-. url p. 105, p. 15, p. 152, p. 153.
- Flora of the northwest coast, including the area west of the summit of the Cascade Mountains, from the forty-ninth parallel south to the Calapooia Mountains on the south border of Lane County, Oregon. Lancaster, Pa., Press of the New Era Printing Company, 1915. url , .
- Flora of the northwest coast: including the area west of the summit of the Cascade Mountains, from the forty-ninth parallel south to the Calapooia Mountains on the south border of Lane County, Oregon / by Charles V. Piper and R. Kent Beattie. Lancaster, Pa.: Press of the New era printing company, 1915. url p. 2, p. 3.
- Flora of the state of Washington. By Charles V. Piper. Washington, Govt. Print. Off., 1906. url p. 77.
- Memoirs of the California Academy of Sciences. San Francisco: The Academy, 1868- url p. 131.
- Muhlenbergia. Chico, Calif. [etc] url p. 21.
- Popular science monthly. New York: D. Appleton, 1872-1950. url p. 207.
- Postelsia: the year book of the Minnesota Seaside Station. St. Paul, Minn.: Pioneer Press, 1902-1906. url p. 33, p. 33.
- Principles of modern biology. New York, Holt, Rinehart and Winston url .
- Proceedings of the California Academy of Sciences, 4th series. San Francisco, California Academy of Sciences. url p. 605.
- Proceedings of the United States National Museum. Washington: Smithsonian Institution Press, [etc.] url p. 628.
- Shade gardens / by Oliver E. Allen and the editors of Time-Life Books. Alexandria, Va.: Time-Life Books, c1979. url p. 137.
- Syllogeus. Ottawa, National Museum of Natural Sciences, 1972-1995. url p. 117, p. 13, p. 23, p. 86.
- The Popular science monthly. New York, Popular Science Pub. Co., etc. url p. 207.
- The ferns of North America colored figures and descriptions, with synonymy and geographical distribution, of the ferns (including the ophioglassaceae) of the United States of America and the British North American possessions / by Daniel Cady Eaton; the drawings by J.H. Emerton and C.E. Faxon. Salem [Mass.]: S.E. Cassino, 1879. url .
- Zoe:a biological journal. 2 1891 San Diego, Calif. [etc.]Zoe Publishing Co. url p. 129, p. 382.
- Cranfill, R. and D. M. Britton. 1983. Typification within the Polypodium virginianum complex (Polypodiaceae). Taxon 32: 557--560.
- Evans, A. M. 1971. Polypodium. In: A. M. Evans, ed. 1971. A Review of Systematic Studies of the Pteridophytes of the Southern Appalachians. Blacksburg.
- Haufler, C. H. and M. D. Windham. 1991. New species of North American Cystopteris and Polypodium, with comments on their reticulate relationships. Amer. Fern J. 81: 7--23.
- Haufler, C. H. and Wang Z. R. 1991. Chromosomal analyses and the origin of allopolyploid Polypodium virginianum. Amer. J. Bot. 78: 624--629.
- Lang, F. A. 1971. The Polypodium vulgare complex in the Pacific Northwest. Madroño 21: 235--254.
- Whitmore, S. A. and A. R. Smith. 1991. Recognition of the tetraploid, Polypodium calirhiza (Polypodiaceae), in western North America. Madroño 38: 233--248.
- Brands, S.J. (comp.) 1989-present. The Taxonomicon. Universal Taxonomic Services, Zwaag, The Netherlands. Accessed March 27, 2012.
Accessed through GBIF Data Portal November 19, 2007:
- Berkeley Natural History Museums, University and Jepson Herbaria DiGIR provider
- Oregon State University, Vascular Plant Collection
- USDA PLANTS, USDA PLANTS Database
- University of Alaska Museum of the North, University of Alaska Museum of the North Herbarium
- University of Washington Burke Museum, Vascular Plant Collection - University of Washington Herbarium
- Utah State University, USU-UTC Specimen Database
- Biodiversity Heritage Library NamebankID: 2644469
- Catalogue of Life Accepted Name Code: ITS-17239
- Global Biodiversity Information Facility Taxonkey: 2481187
- Globally Unique Identifier: urn:lsid:ipni.org:names:17200880-1
- Integrated Taxonomic Information System (ITIS) Taxonomic Serial Number (TSN): 17239
- Natural Heritage Network Species Identifier: PPPOL020V0
- U.S.D.A. Plant Symbol: POSC4
- Zipcode Zoo Species Identifier: 57612
- Alan R. Smith "Polypodiaceae". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- Christopher H. Haufler, Michael D. Windham, Frank A. Lang, S. A. Whitmore "Polypodium". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- "Polypodium scouleri". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- Mean = 191.900 meters (629.593 feet), Standard Deviation = 294.730 based on 82 observations. Altitude information for each observation from British Oceanographic Data Centre. [back]