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Interesting Facts
Description
Family Dryopteridaceae
Plants
perennial
, terrestrial
or on rock, occasionally hemiepiphytic
or epiphytic. Stems creeping
to erect
, rarely arborescent
, sometimes climbing
, branched or unbranched, dictyostelic, bearing scales
. Leaves circinate
in bud, monomorphic
or dimorphic
. Petiole
usually not articulate
to stem, scales usually persistent
at base
, in cross
section
with 2--many roundish bundles, or bundles 2 and lunate
. Blade
simple
to commonly 1--5-pinnate or more divided
, leaf buds
absent or present. Veins pinnate or parallel in ultimate
segments, simple or forked
, free
or anastomosing, areoles sometimes with included
free veinlets. Indument
on blade commonly of glands
, hairs
, and/or scales, especially on rachis and costae abaxially. 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. Sporangia with stalk
of 2--3 rows
of cells
; annulus vertical
, interrupted
by stalk. Spores 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.
) . Gametophytes green, aboveground, cordate, glabrous
or often bearing glands or hairs; archegonia and antheridia borne on lower surface, antheridia 3-celled.
Genera ca.
60, species perhaps exceeding 3000 (18 genera, 79 species in the flora
) : worldwide.
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.[1]
Genus Polystichum
Plants
terrestrial
. Stems decumbent
to erect
, stolons absent. Leaves monomorphic
(dimorphic
in P. acrostichoides ), evergreen
. Petiole
1/9--1 times length
of blade
, bases
swollen or not; vascular bundles
more than 3, arranged in an arc, ± round
in cross
section
. Blade linear-lanceolate to broadly lanceolate, 1--3-pinnate, gradually reduced distally to pinnatifid
apex, somewhat leathery to leathery. Pinnae not articulate
to rachis, segment or pinna margins
spinulose-toothed (except P. lemmonii ) ; proximal
pinnae (several pairs) usually gradually reduced, sessile to short-petiolulate, bases usually inequilateral
with acroscopic
lobe
; costae adaxially grooved
, grooves
continuous from rachis to costae; indument
of linear
to lanceolate scales on costae and sometimes between veins abaxially (microscales), ± glabrous
or similarly scaly
adaxially (scales
forming loosely tangled network
over blade and sori in P. dudleyi ). Veins free, forked
, rarely ( P. imbricans ) anastomosing. Sori in 1 row
(to several) between midrib
and margins, round (confluent
, covering abaxial
surface in P. acrostichoides ) ; indusia peltate, persistent
or caducous
[absent]. Spores yellow or brownish to black, with inflated
folds. x
= 41.
Species ca.
180: worldwide.
The mating systems
of Polystichum seem to be highly outcrossing (P. S. Soltis and D. E. Soltis 1987; P. S. Soltis et al.
1989) ; hybrids are frequent where two or more species occur. Sterile
hybrids are discussed under one of their putative parents.
Sterile hybrids are best recognized by their misshapen sporangia, which produce
little black dots at the end of the season
instead of forming the fuzzy brown bump
typical of sori after spores have been expelled. In many cases the intermediacy and robustness
of hybrids make them stand out as odd. At least one or two hybrid plants are to be expected in large, mixed populations. The allopolyploids, having hybrid origins
, present particular problems. They exhibit
the Vavilov effect: allopolyploids tend to resemble one of their parental species when they grow with, or in the habitat
typical of, that species (D. S. Barrington et al. 1989).
In the flora
there are six diploids, five tetraploids
, one hexaploid
, and three species whose chomosome number is unknown. Relationships
among the diploids are generally not very close; that is, each is probably more closely related to a species outside the flora than to one of the other species in the flora. The exception to this is the group composed of Polystichum acrostichoides, P. imbricans, and P. munitum. Polystichum acrostichoides appears to share a Tertiary common ancestor
with P. munitum, and P. imbricans is more recently derived from P. munitum. All of the polyploid species are fertile
allopolyploids. One of these species ( P. braunii ) is also involved in the formation of the hexaploid P. setigerum (see below).
Relationships among Polystichum Species
Allopolyploid Presumed Originating Crosses: andersonii kwakiutlii × munitum californicum dudleyi × imbricans or dudleyi × munitum kruckebergii lemmonii × lonchitis scopulinum lemmonii × imbricans or lemmonii × munitum setigerum braunii × munitum
The morphological similarity
among Polystichum species may make identification difficult, particularly among the species with more divided
leaves. The keys
presented here are designed for mature
, typical individuals. Some of the characters mentioned in the keys and descriptions
require the use of a microscope. The microscales (small trichomes
that occur on the abaxial leaf surface of all species and adaxially in some) are best observed by peeling them off with cellophane tape and mounting the tape on a slide
, sticky side up, under a coverslip. The tape can also be used to lift
off the components
of the sori. Polystichum acrostichoides, P. andersonii, P. lemmonii, and P. munitum are known to have sclereid
clusters
in their pith
. Polystichum imbricans lacks such clusters, and data are not available for the other species.[2]
Habitat
Biome: Terrestrial [3].
Ecology:
Occurs in low Andean forest
(1,500–2,000 m
).[3].
List of Habitats
:
- 1 Forest
- 1.6 Forest - Subtropical/Tropical Moist Lowland
- 1.9 Forest - Subtropical/Tropical Moist Montane [more info]
Taxonomy
- Domain:
Eukaryota
(
)
- Whittaker & Margulis,1978
- eukaryotes
- Kingdom:
Plantae
(
)
- Haeckel, 1866
- Plants
- Subkingdom:
Viridaeplantae
(
)
- Cavalier-Smith, 1981
- Phylum:
Tracheophyta
(
)
- Sinnott, 1935 ex Cavalier-Smith, 1998
- Vascular Plants
- Subphylum:
Euphyllophytina
(
)
- Infraphylum:
Moniliformopses
(
)
- Kenrick & Crane, 1997, Nom. Nud.
- Class:
Polypodiopsida
(
)
- Cronquist et al.
- Order:
Polypodiales
(
)
- Link
- Family:
Dryopteridaceae
(
)
- Ching, 1965, nom. cons.
- Wood Fern Family
- Genus:
Polystichum
(
)
- A.W. Roth, 1799, nom. cons.
- [Sword fern, Christmas fern, holly fern [Greek poly, many, and stichos, row, presumably in reference to the rows of sori on each pinna]
- Specific epithet:
bulbiferum
- Barrington
- Botanical name: - Polystichum bulbiferum Barrington
- Specific epithet:
bulbiferum
- Barrington
- Genus:
Polystichum
(
- Family:
Dryopteridaceae
(
- Order:
Polypodiales
(
- Class:
Polypodiopsida
(
- Infraphylum:
Moniliformopses
(
- Subphylum:
Euphyllophytina
(
- Phylum:
Tracheophyta
(
- Subkingdom:
Viridaeplantae
(
- Kingdom:
Plantae
(
Similar Species
Members of the genus Polystichum
ZipcodeZoo has pages for 61 species, subspecies, varieties, forms, and cultivars in this genus:
P. acrostichoides (Christmas Fern) · P. acrostichoides f. crispum (Christmas Fern) · P. acrostichoides f. demittens (Christmas Fern) · P. acrostichoides f. gymnosorum (Christmas Fern) · P. acrostichoides f. incisum (Christmas Fern) · P. acrostichoides f. ligulatum (Christmas Fern) · P. acrostichoides f. multifida (Christmas Fern) · P. acrostichoides f. recurvatum (Christmas Fern) · P. acrostichoides f. spathiforme (Christmas Fern) · P. acrostichoides f. ventroperaferens (Christmas Fern) · P. acrostichoides var. acrostichoides (Christmas Fern) · P. acrostichoides var. crispum (Christmas Fern) · P. acrostichoides var. incisum (Christmas Fern) · P. acrostichoides var. lonchitoides (Christmas Fern) · P. aculeatum (Hard Shield Fern) · P. aleuticum (Aleutian Holly Fern) · P. andersonii (Anderson Holly-Fern) · P. bonseyi (Bonsey's Hollyfern) · P. braunii (Braun's Holly-Fern) · P. braunii f. alaskense (Braun´s Holly Fern) · P. braunii var. andersonii (Braun´s Holly Fern) · P. calderonense (Monte Guilarte Hollyfern) · P. californicum (California Sword-Fern) · P. dudleyi (Dudley's Sword Fern) · P. echinatum (Rusty Swordfern) · P. falcatum var. falcatum (Japanese Holly Fern) · P. hagenahii (Hagenah's Polystichum) · P. haleakalense (Island Swordfern) · P. hillebrandii (Hillebrand Holly-Fern) · P. imbricans (Narrow-Leaf Swordfern) · P. imbricans curtum (Narrowleaf Swordfern) · P. imbricans imbricans (Narrowleaf Swordfern) · P. imbricans subsp. curtum (Narrowleaf Swordfern) · P. kruckebergii (Kruckeberg's Hollyfern) · P. kwakiutlii (Kwakiutl's Hollyfern) · P. lemmonii (Lemmon's Hollyfern) · P. lonchitis (Hollyfern) · P. makinoi (Makinoi's Holly Fern) · P. microchlamys (Attu Holly Fern) · P. munitum (Sword Fern) · P. munitum f. flabellatum (Western Sword Fern) · P. munitum f. inciso-serratum (Western Sword Fern) · P. munitum f. nudatum (Western Sword Fern) · P. munitum subsp. solitarium (Western Sword Fern) · P. muricatum (West Indian Hollyfern) · P. neolobatum (Long-Eared Holly Fern) · P. ovatopaleaceum (Polystichum) · P. platyphyllum (Flatleaf Hollyfern) · P. polyblepharum (Holly Fern) · P. polystichiforme (Antilles Hollyfern) · P. potteri (Potter's Polystichum) · P. proliferum (Mother Shield Fern) · P. rhizophyllum (Tailed Hollyfern) · P. scopulinum (Eaton's Hollyfern) · P. setiferum (Alaskan Fern) · P. setiferum Plumosomultilobum Group (Soft-Shield Fern) · P. setiferum 'Congestum' (Dwarf Soft Shield Fern) · P. setiferum 'Divisilobum' (Soft Shield Fern) · P. setigerum (Alaska Holly Fern) · P. tripteron (Trifid Holly Fern) · P. tsus-simense (Korean Rock Fern)
More Info
- Search for Pictures: images.google.com
- Search for Scholarly Articles: Google Scholar
- Search using Scientific Name and Vernacular Names: All the Web | AltaVista Canada | AltaVista | Excite | Google | HotBot | Lycos
- Search using Specialized Databases: GenBank | Medline | Scirus | CISTI/CAL | Agricola Periodicals | Agricola Books
Further Reading
- IUCN. 2003. 2003 IUCN Red List of Threatened Species. www.iucnredlist.org. Downloaded on 18 November 2003.
- Valencia, R., Pitman, N., León-Yánez, S. and Jørgensen, P.M. (eds). 2000. Libro Rojo de las Plantas Endmicas del Ecuador 2000. Publicaciones del Herbario QCA, Ponticicia Universidad Católica del Ecuador, Quito.
- Valencia, R., Pitman, N., Len-Ynez, S. and Jrgensen, P.M. (eds). 2000. Libro Rojo de las Plantas Endmicas del Ecuador 2000. Publicaciones del Herbario QCA, Ponticicia Universidad Catlica del Ecuador, Quito.
- Wagner, D. H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1--64.
- Wagner, W. H. Jr. 1973. Reticulation of holly ferns (Polystichum) in the western United States and adjacent Canada. Amer. Fern J. 63: 99--115.
- Welsh, S. L. 1974. Anderson's Flora of Alaska and Adjacent Parts of Canada. Provo.
- Yatskievych, G., D. B. Stein, and G. J. Gastony. 1988. Chloroplast DNA evolution and systematics of Phanerophlebia (Dryopteridaceae) and related fern genera. Proc. Natl. Acad. Sci. U.S.A. 85: 2589--2593.
Notes
Contributors
- Brands, S.J. (comp.) 1989-present. The Taxonomicon. Universal Taxonomic Services, Zwaag, The Netherlands. Accessed January 9, 2012.
- IUCN 2012. IUCN Red List of Threatened Species. Version 2011.2. . Downloaded on January 28, 2012.
- Navarrete, H. & Pitman, N. 2003. Polystichum bulbiferum. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2. <www.iucnredlist.org>. Downloadedon 04February2012.
Identifiers
- Biodiversity Heritage Library NamebankID: 7151279
- Global Biodiversity Information Facility Taxonkey: 15459851
- Globally Unique Identifier: urn:lsid:ipni.org:names:17529800-1
- IUCN ID: 234737
- Zipcode Zoo Species Identifier: 1027190
Footnotes
- Alan R. Smith "Dryopteridaceae". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- David H. Wagner "Polystichum". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- Navarrete, H. & Pitman, N. 2003. Polystichum bulbiferum. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2. <www.iucnredlist.org>. Downloaded on 04 February 2012. [back]
