Common Names in English:
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 compact
to long-creeping, ascending
to horizontal, usually branched; scales
brown to black or often bicolored
with dark central stripe and lighter margins
, linear-subulate to ovate-lanceolate, margins entire or denticulate
. Leaves monomorphic
, clustered to widely scattered
, 4--60 cm. Petiole
brown to black or straw-colored, rounded
, flattened, or with single longitudinal
, or glabrous
, with a single vascular bundle. Blade
linear-oblong to lanceolate, ovate
, or elongate-pentagonal, pinnate-pinnatifid to 4-pinnate at base
, leathery or rarely somewhat herbaceous, abaxially pubescent and/or scaly, rarely glabrous, adaxially pubescent to glabrous, dull
, not striate
; rachis straight. Ultimate
segments of blade stalked or sessile, usually free
from costae, round
, usually less than 4 mm wide, base rounded, truncate
, or cuneate; stalks
(when present) often lustrous
and dark colored
; segment margins usually recurved to form confluent
, poorly defined false indusia, extending entire length of segment or discontinuous on apical or lateral
. Veins of ultimate segments free or rarely anastomosing, pinnately branched and divergent distally. False indusia greenish to whitish, usually narrow, clearly marginal
or rarely inframarginal, often concealing sporangia. Sporangia confined to submarginal
or scattered along veins near segment margins, containing 64 or 32 spores, not intermixed with farina-producing glands
. Spores brown to black or gray, rarely yellowish, tetrahedral-globose, rugose
, lacking prominent
. Gametophytes glabrous. x
= 30 (29 in Cheilanthes alabamensis complex
Species ca. 150: mostly Western Hemisphere but a few in Europe, Asia, Africa, Pacific Islands, and Australia.
Cheilanthes is by far the largest and most diverse genus of xeric-adapted ferns. In its classic circumscription, the genus has been notoriously difficult to distinguish from other cheilanthoid genera, especially Notholaena and Pellaea (R. M. Tryon and A. F. Tryon 1982). This has led some authors (e.g. , J. T. Mickel 1979b) to abandon several of the segregate genera and greatly expand the number of species assigned to Cheilanthes. Taxonomic problems in this group have motivated an ongoing series of biosystematic studies that offer hope for a stable classification through the identification of natural, monophyletic groups. The circumscription of Cheilanthes has been clarified recently by a redefinition of Notholaena and the transfer of several species (e.g., N. parryi, N. newberryi, and N. aurea ) to Cheilanthes (R. M. Tryon and A. F. Tryon 1982). The boundaries of the genus have been further sharpened by the recognition of Aspidotis (A. R. Smith 1975), Argyrochosma (M. D. Windham 1987), and Astrolepis (D. M. Benham and M. D. Windham 1992) as distinct genera. Despite these efforts , Cheilanthes remains a very heterogeneous (and probably polyphyletic) genus in need of further critical study.
The circumscription of Cheilanthes used here closely parallels that proposed by T. Reeves (1979), who recognized four New World subgenera and a small group of species of uncertain placement. Among the North American species, C. pringlei and C. wrightii belong to the latter group, and C. arizonica is the sole representative of subgenus Othonoloma Link ex C. Christensen. The subgenus that Reeves called the Cheilanthes alabamensis group is represented in North America by C. aemula, C. alabamensis, C. microphylla, and C. horridula. It differs from other members of the genus in a number of critical features (e.g., blade indument, sporangial distribution, and chromosome base number ) that suggest a relationship to the genus Pellaea. Although the group is somewhat anomalous in Cheilanthes, inclusion of the C. alabamensis complex in Pellaea (e.g., R. Cranfill 1980) would make that genus polyphyletic. The group is maintained here in Cheilanthes with the recognition that it may constitute a natural group worthy of consideration as a distinct genus.
The remaining 21 North American species of Cheilanthes are almost evenly divided between subgenus Physapteris (C. Presl) Baker in Hooker & Baker and subgenus Cheilanthes. The former subgenus is characterized by T. Reeves (1979) as having noncircinate vernation , usually scaly blades with small, beadlike ultimate segments, and hairs with cell walls that fit together in "tongue and groove" fashion. Species of subgenus Cheilanthes typically have circinate vernation, nonscaly blades with larger ultimate segments, and hairs with straight cross-walls. Despite the consistency of these differences, the two groups are closely related and linked by occasional intersubgeneric hybridization between C. covillei (subg. Physapteris ) and C. parryi and C. newberryi (subg. Cheilanthes ).
Flowers: Flower Color: inconspicuous, none
Typically found at an altitude of 0 to 4,057 meters (0 to 13,310 feet).
- Whittaker & Margulis,1978
- Haeckel, 1866
- Cavalier-Smith, 1981
- Sinnott, 1935 ex Cavalier-Smith, 1998
- Vascular Plants
- Kenrick & Crane, 1997, Nom. Nud.
- Cronquist et al.
- 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
Publishing author : Maxon
Members of the genus Cheilanthes
ZipcodeZoo has pages for 47 species, subspecies, varieties, forms, and cultivars in this genus:
C. aemula (Rival Lipfern) · C. alabamensis (Alabama Lip Fern) · C. arizonica (Arizona Lip Fern) · C. austrotenuifolia (Green Rockfern) · C. bonariensis (Golden Lip Fern) · C. clevelandii (Cleveland Lipfern) · C. cooperae (Cooper's Lipfern) · C. covillei (Coville's Lip Fern) · C. decipiens (Triangle Digit Fern) · C. decora (Lance Fern) · C. distans (Bristly Cloak-Fern) · C. eatonii (Eaton Lipfern) · C. farinosa (Floury Cloak Fern) · C. feei (Fee Lipfern) · C. fendleri (Fendler Lipfern) · C. fibrillosa (Fibril Lipfern) · C. gracillima (Lace Lipfern) · C. horridula (Rough Lip Fern) · C. intertexta (Coastal Lipfern) · C. kaulfussii (Kaulfuss Lipfern) · C. lanosa (Hairy Lipfern) · C. lanosa var. album (Hairy Lipfern) · C. lasiophylla (Woolly Cloak-Fern) · C. lendigera (Nit-Bearing Lip Fern) · C. leucopoda (White-Foot Lipfern) · C. lindheimeri (Fairy Swords) · C. microphylla (Southern Lipfern) · C. newberryi (Newberry's Lipfern) · C. notholaenoides (Royal Lipfern) · C. parishii (Parish Cheilanthes) · C. parishii patellaria var. patellaria (Parish Cheilanthes) · C. parishii var. douglasii (Parish Cheilanthes) · C. parryi (Parry Cloakfern) · C. paupercula (Streambank Lipfern) · C. pringlei (Pringle's Lip Fern) · C. quadripinnata (Lesotho Lip Fern) · C. takeuchii (Takeuch's Lipfern) · C. ternifolia (Trans-Pecos Cliffbrake) · C. tomentosa (Woolly Lip Fern) · C. villosa (Villous Lip Fern) · C. viscida (Viscid Lipfern) · C. wootonii (Beaded Lip Fern) · C. wrightii (Wright Lipfern) · C. x (Fibril Lipfern) · C. x fibrillosa (Fibrillose Lip Fern) · C. x parishii (Parish Cheilanthes) · C. yavapensis (Graceful Lipfern)
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- Bulletin of the Natural History Museum. London: The Natural History Museum, c1993-2002. url p. 91.
- Contributions to the flora of Venezuela / Julian A. Steyermark and collaborators. 28 1957 Chicago, Ill.: Chicago Natural History Museum, 1957. url p. 750.
- Ferns and fern allies of Guatemala. Robert G. Stolze; the genus Elaphoglossum by: John T. Mickel; the genus Thelypteris by: Alan R. Smith. 6 1981 Chicago, Ill.: Field Museum of Natural History, 1981. url p. 136, p. 138.
- Phytologia memoirs. Plainfield, N.J.: H.N. Moldenke and A.L. Moldenke, 1980- url p. 17.
- Pteridophyta of Peru. Rolla M. Tryon, Robert G. Stolze. 22 1989 Chicago, Ill.: Field Museum of Natural History, c1989. url p. 25.
- Wrightia. 7 1981-1984 Richardson, Tex. [etc.]University of Texas at Dallas. url p. 196.
- 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.
- Benham, D. M. and M. D. Windham. 1992. Generic affinities of the star-scaled cloak ferns. Amer. Fern. J. 82: 47--58.
- Mickel, J. T. 1979b. The fern genus Cheilanthes in the continental United States. Phytologia 41: 431--437.
- Reeves, T. 1979. A Monograph of the Fern Genus Cheilanthes Subgenus Physapteris (Adiantaceae). Ph.D. dissertation. Arizona State University.
- Smith, A. R. 1975. The California species of Aspidotis. Madroño 23: 15--24.
- Tryon, R. M. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179: 1--106.
- Bisby, F.A., Y.R. Roskov, M.A. Ruggiero, T.M. Orrell, L.E. Paglinawan, P.W. Brewer, N. Bailly, J. van Hertum, eds (2007). Species 2000 & ITIS Catalogue of Life: 2007 Annual Checklist. Species 2000: Reading, U.K.
- Brands, S.J. (comp.) 1989-present. The Taxonomicon. Universal Taxonomic Services, Zwaag, The Netherlands. Accessed January 10, 2012.
- Global Biodiversity Information Facility. Accessed February 28, 2008. http://www.gbif.org Mediated distribution data from 3 providers.
- Ruggiero M., Gordon D., Bailly N., Kirk P., Nicolson D. (2011). The Catalogue of Life Taxonomic Classification, Edition 2, Part A. In: Species 2000 & ITIS Catalogue of Life: 2011 Annual Checklist (Bisby F.A., Roskov Y.R., Orrell T.M., Nicolson D., Paglinawan L.E., Bailly N., Kirk P.M., Bourgoin T., Baillargeon G., Ouvrard D., eds). DVD; Species 2000: Reading, UK.
- The International Plant Names Index. Accessed Dec 27, 2011.
- USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA.
Accessed through GBIF Data Portal February 28, 2008:
- Comisión nacional para el conocimiento y uso de la biodiversidad: Herbario del Instituto de Ecología, A.C., México (IE-BAJÍO)
- Comisión nacional para el conocimiento y uso de la biodiversidad: Herbario del Instituto de Ecología, A.C., México (IE-XAL)
- Herbarium of the University of Aarhus: The AAU Herbarium Database
- Missouri Botanical Garden
- Biodiversity Heritage Library NamebankID: 2666149
- Catalogue of Life Accepted Name Code: ITS-501476
- Globally Unique Identifier: urn:lsid:ipni.org:names:17375950-1
- Integrated Taxonomic Information System (ITIS) Taxonomic Serial Number (TSN): 501476
- International Plant Names Index (IPNI) ID: 1093919-1
- Natural Heritage Network Species Identifier: PPADI090L0
- U.S.D.A. Plant Symbol: CHNO2
- Zipcode Zoo Species Identifier: 29668
- Michael D. Windham "Pteridaceae". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- Michael D. Windham, Eric W. Rabe "Cheilanthes". in Flora of North America Vol. 2. Oxford University Press. Online at EFloras.org. [back]
- Mean = 1,519.370 meters (4,984.810 feet), Standard Deviation = 993.940 based on 84 observations. Altitude information for each observation from British Oceanographic Data Centre. [back]