Common Names in English:
California Cloak Fern
40, species ca. 1000 (13 genera, 90 sp: worldwide.
Considerable disagreement exists concerning the circumscription and proper name of this family . The taxa comprising the Pteridaceae in this treatment were assigned to the Sinopteridaceae and Pteridaceae by D. B . Lellinger (1985) and were included in five families by R. E. G. Pichi-Sermolli (1977). The broad concept followed here is similar (except for the exclusion of Ceratopteris ) to that espoused by R. M. Tryon and A. F. Tryon (1982), who applied the name Pteridaceae to the group. Until very recently, the newer name Adiantaceae was more commonly used.
As represented in North America, Pteridaceae comprise three major evolutionary lines (the adiantoids, the pteroids, and the cheilanthoids). Characteristics holding the family together include abaxial (usually submarginal ) sori that lack indusia or are protected by a reflexed or revolute leaf margin , spores that are usually globose-tetrahedral and trilete, and chromosome base numbers of 30 or 29 (rarely 27). The xeric-adapted members of the family (particularly the cheilanthoids) have undergone extensive parallel and convergent evolution, and they have frustrated attempts to produce a natural generic classification based on macromorphologic characteristics alone. Although some workers have aggregated species into a few large genera (e.g. , J. T. Mickel 1979b), most tend to recognize smaller segregate genera based on a combination of morphologic, chromosomal, and biochemical data. The latter approach seems to provide a more useful, evolutionarily informative classification and is the one adopted here. Aspidotis and Notholaena are maintained here as distinct from Cheilanthes, and three recently described genera ( Argyrochosma, Astrolepis, and Pentagramma ) have been incorporated into the treatment. The reasons for these changes in generic circumscription are discussed under the individual genera.
usually on rock. Stems short-creeping to compact
to horizontal, usually branched; scales
black or often bicolored
with dark central stripe and lighter margins
, linear-subulate to lanceolate; margins ciliate
, or entire. Leaves monomorphic
, clustered, 4--35 cm. Petiole
brown or black, usually rounded
, flattened, or with single longitudinal
adaxially, often bearing scales, hairs
, or farinose
, with single vascular bundle. Blade
, deltate, or pentagonal
, pinnate-pinnatifid to 4-pinnate, leathery, abaxially covered by yellowish or whitish farina
(completely obscured by stellate
scales in Notholaena aschenborniana ), adaxially often sparsely glandular
, not striate
; rachis straight. Ultimate
segments of blade sessile to subsessile
, often adnate
to costae, narrowly elliptic
to oblong-ovate or deltate, usually less than 4 mm wide; base
rounded to truncate
or cuneate; stalks
(when present) usually lustrous
and dark-colored; segment margins recurved to form confluent
, poorly defined false indusia extending entire length of segment. Veins of ultimate segments free
, pinnately branched and divergent distally. False indusia greenish, narrow, clearly marginal
, occasionally concealing the sporangia. Sporangia confined to submarginal
, containing 64, 32, or 16 spores, intermixed with farina-producing glands. Spores black to dark brown, globose
or tetrahedral-globose, granulate
, lacking prominent
. Gametophytes glandular-farinose (gametophytes of all other Pteridaceae genera lack farina). x
Species ca. 25: North America, Mexico, West Indies, Central America, South America.
As pointed out by R. M. Tryon (1956), J. T. Mickel (1979), and many others, Notholaena in the broad sense is poorly defined and difficult to distinguish from either Cheilanthes or Pellaea. North American taxa traditionally assigned to Notholaena represent at least four distinct evolutionary lineages (M. D. Windham 1986). In order to clarify species relationships and generic boundaries among cheilanthoid ferns, Notholaena is defined here in a very restricted sense. The pubescent , nonfarinose species (such as N. newberryi ) have been placed in Cheilanthes following R. M. Tryon and A. F. Tryon (1982). The scaly , nonfarinose taxa of the N. sinuata complex have been transferred to Astrolepis for reasons discussed by D. M. Benham and M. D. Windham (1992). The glabrous and farinose species related to Pellaea (e.g. , N. jonesii and N. dealbata ) have been placed in Argyrochosma (M. D. Windham 1987). The species retained in Notholaena following this reorganization (i.e. , members of the farinose N. grayi -- N. standleyi alliance ) form a coherent, monophyletic group found only in the Western Hemisphere. The correct generic name for this group is in dispute because Notholaena has been lectotypified by several authors citing three different type species. The rules of priority favor the first typification (by J. Smith in 1875) based on N. trichomanoides (Linnaeus) Desvaux, which is definitely a member of the group here called Notholaena. R. E. G. Pichi-Sermolli (1989), however, urged acceptance of the typification by C. Christensen ([1905--]1906). Both the second and third lectotypifications of Notholaena were based on species unrelated to the North American taxa discussed here. If Smith's typification is overturned, the correct generic name for our species will be Chrysochosma (J. Smith) Kümmerle.
- Whittaker & Margulis,1978
- Haeckel, 1866
- Cavalier-Smith, 1981
- Sinnott, 1935 ex Cavalier-Smith, 1998
- Vascular Plants
- Kenrick & Crane, 1997, Nom. Nud.
- Cronquist et al.
- Maidenhair Fern Family Plants perennial [annual],
- Cloak fern [Greek notho, false, and chlaena, coat, in reference to the reflexed leaf segment margins that form false indusia]
- Specific epithet:
- D.C. Eat.
- Botanical name: - Notholaena californica californica D.C. Eat.
- Subspecies: californica
- Specific epithet: californica - D.C. Eat.
- Genus: Notholaena () - Cloak fern [Greek notho, false, and chlaena, coat, in reference to the reflexed leaf segment margins that form false indusia]
- Subfamily: Cheilanthoideae ()
- Family: Pteridaceae () - Kirchn. - Maidenhair Fern Family Plants perennial [annual],
- 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
Notholaena californica D. C. Eat.
Members of the genus Notholaena
ZipcodeZoo has pages for 19 species, subspecies, varieties, forms, and cultivars in this genus:
N. aliena (Foreign Cloak Fern) · N. aschenborniana (Scaled Cloak Fern) · N. atropurpurea (Purple-Stem Cliff-Brake) · N. bonariensis (Golden Cloak Fern) · N. californica (California Cloak Fern) · N. californica californica (California Cloak Fern) · N. californica leucophylla (California Cloak Fern) · N. californica subsp. leucophylla (California Cloak Fern) · N. copelandii (Copeland's Cloak Fern) · N. grayi (Gray's Cloak Fern) · N. grayi sonorensis (Gray's Cloak Fern) · N. grayi subsp. sonorensis (Sonoran Cloak Fern) · N. greggii (Gregg Cloak-Fern) · N. lemmonii (Lemmon's Cloak Fern) · N. nealleyi (Nealley's Cloak Fern) · N. neglecta (Maxon Cloak-Fern) · N. standleyi (Star Cloak Fern) · N. ternifolia (Trans-Pecos Cliffbrake) · N. trichomanoides (Rockslope Cloak Fern)
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- Lellinger, D. B. 1985. A Field Manual of the Ferns & Fern-allies of the United States & Canada. Washington.
- Mickel, J. T. 1979b. The fern genus Cheilanthes in the continental United States. Phytologia 41: 431--437.
- Pichi-Sermolli, R. E. G. 1977. Tentamen pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 31: 313--512.
- Tryon, R. M. and A. F. Tryon. 1982. Ferns and Allied Plants, with Special Reference to Tropical America. New York, Heidelberg, and Berlin.
- Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1--106.
- Windham, M. D. 1986. Reassessment of the phylogenetic relationships of Notholaena. Amer. J. Bot. 73: 742.
- Bisby FA, Roskov YR, Orrell TM, Nicolson D, Paglinawan LE, Bailly N, Kirk PM, Bourgoin T, van Hertum J, eds (2008). Species 2000 & ITIS Catalogue of Life: 2008 Annual Checklist Taxonomic Classification. CD-ROM; Species 2000: Reading, U.K.
- Brands, S.J. (comp.) 1989-present. The Taxonomicon. Universal Taxonomic Services, Zwaag, The Netherlands. Accessed January 10, 2012.
- Biodiversity Heritage Library NamebankID: 2644741
- Catalogue of Life Accepted Name Code: ITS-524335
- Global Biodiversity Information Facility Taxonkey: 13901882
- Natural Heritage Network Species Identifier: PPADI0G041
- Zipcode Zoo Species Identifier: 167531