Myriopteris clevelandii | |
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Frond of Myriopteris clevelandii showing the bead-like appearance of its segments | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Order: | Polypodiales |
Family: | Pteridaceae |
Genus: | Myriopteris |
Species: | M. clevelandii |
Binomial name | |
Myriopteris clevelandii | |
Synonyms | |
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Myriopteris clevelandii, formerly known as Cheilanthes clevelandii, is a species of lip fern known by the common name Cleveland's lip fern. It is native to southern California and Baja California in Mexico. The leaf is divided into small, bead-like segments densely covered with scales beneath. In M. clevelandii, some of these scales are reduced to hairlike structures, which help distinguish it from the closely related M. covillei . It is usually found growing on exposed rock, particularly igneous rock.
The rhizomes are horizontal and range from 1 to 3 millimeters (0.04 to 0.1 in) in diameter. The leaves are closely [2] [3] or broadly spaced along them. [4] The rhizome bears persistent linear-lanceolate scales, which are dark brown [2] [3] or brown to red-brown [4] [5] in color and shiny. The scales may be of a uniform brown color, or bear a dark central stripe with paler edges. [2] [3] [4] The margins of the scales are entire [2] or erose to slightly toothed with teeth well-spaced. [5] The scales are straight or slightly twisted and strongly appressed (pressed against the rhizome). [3]
The fronds arise from the rhizome in clusters or as somewhat scattered individual leaves. Unlike many ferns, they do not emerge as coiled fiddleheads (noncircinate vernation). [3] When mature, they are 8 to 40 centimeters (3 to 20 in) long. [2] [3] The stipe (the stalk of the leaf below the blade) makes up about half the length of the frond, [2] measuring 5 to 31 centimeters (2.0 to 12 in) long. [5] The stipe is shiny, rounded, and dark to light brown, [2] [3] covered with 1-to-2-millimeter (0.04 to 0.08 in)-long hairs and filiform (threadlike) scales [2] that are gray to red-brown in color. [4] The covering is lost as the frond ages. [2] The stipe is typically less than 2 millimeters (0.08 in) wide, sometimes up to 3 millimeters (0.1 in). [4]
The leaf blades are oblong-lanceolate to ovate and tetrapinnate (cut into pinnae, pinnules, pinnulets, and divisions of pinnulets) at the base. They are typically 6 to 23 centimeters (2.4 to 9.1 in) long [5] and 2 to 8 centimeters (0.8 to 3 in) broad. [2] [3] [5] The rachis (leaf axis) is rounded, rather than grooved, on its upper surface, and there is no distinct joint where the pinnae attach to the rachis, the dark color of the latter continuing into the base of the costa (pinna axis). [3] 10 to 12 pairs of pinnae are present in Mexican specimens, somewhat more in some Californian material. [2] Each pinna is equilateral in shape, [2] [3] and the lowest pair of pinnae is not significantly enlarged compared to the others. [3] The upper surface of the costae is green along much of their length. [3] The lower surface of the costae is covered in conspicuous broad scales. These are ovate-lanceolate to broadly deltate in shape, and deeply cordate (notched at the base to appear heart-shaped). They are about 1 millimeter (0.04 in) long and 0.4 to 1 millimeter (0.02 to 0.04 in) wide, overlapping, and sometimes conceal the final subdivisions of the leaf from below. Those closer to the base of the costa are ciliate. [2] [3] The smallest divisions of the leaf are round or slightly heart-shaped, beadlike in appearance, [3] [4] not exceeding 1 to 2 millimeters (0.04 to 0.08 in) across [3] and concave below. [4] The upper surface of the leaf is glabrous (free of hairs). [2] [3] [4] [5] The lower surface of the leaf is covered in ciliate scales, similar to those of the costa but reduced in width so as to appear like branched hairs in some cases, [2] [3] more or less concealing the surface. [4]
On fertile fronds, the sori are protected by false indusia formed by the edge of the leaf curling strongly back over the underside, often concealing the sori. [4] [5] The recurved edges are only a little modified in comparison to the rest of the leaf tissue. They are 0.05 to 0.25 millimeters (0.0020 to 0.0098 in) wide, with entire margins. The sori contain brown spores, with 64 spores in each sporangium. [2] [3]
Specimens from some of the northern Channel Islands are larger, with more dissected scales, and have been referred to as "var. clokeyi", but this name has never been formally published. [2] [4] M. clevelandii is quite similar to M. covillei , usually found more inland. In the latter, the reduced, hairlike scales are not present on the abaxial surface of the leaf tissue, while the scales on the abaxial surface of the costa are larger and lack cilia except on their basal lobes. [2]
The species was first described in 1875 by Daniel Cady Eaton as Cheilanthes clevelandii. He named it for Daniel Cleveland, the collector of the type specimen, which came from "a mountain about forty miles from San Diego, California". [6] By a strict application of the principle of priority, Oliver Atkins Farwell transferred the species to the genus Allosorus as Allosorus myriophyllus var. clevelandii in 1931, that genus having been published before Cheilanthes . [7] Farwell's name was rendered unnecessary when Cheilanthes was conserved over Allosorus in the Paris Code published in 1956.
The development of molecular phylogenetic methods showed that the traditional circumscription of Cheilanthes is polyphyletic. Convergent evolution in arid environments is thought to be responsible for widespread homoplasy in the morphological characters traditionally used to classify it and the segregate genera that have sometimes been recognized. On the basis of molecular evidence, Amanda Grusz and Michael D. Windham revived the genus Myriopteris in 2013 for a group of species formerly placed in Cheilanthes. One of these was C. clevelandii, which thus became Myriopteris clevelandii. [8] In 2018, Maarten J. M. Christenhusz transferred the species to Hemionitis as H. clevelandii, as part of a program to consolidate the cheilanthoid ferns into that genus. [9]
Further molecular studies in Myriopteris demonstrated the existence of three well-supported clades within the genus. M. clevelandii belongs to what Grusz et al. informally named the covillei clade. Members of the "core covillei" clade, including M. clevelandii, have leaves finely divided into bead-like segments. Within this clade, M. clevelandii is sister to M. covillei . [10]
The fern is native to southern California, [11] specifically the Peninsular Ranges and several of the northern Channel Islands, [4] and to northern Baja California, Mexico. [2] [12]
It is found in a variety of rocky, exposed habitats, including chaparral, [4] on slopes and ledges, [2] [3] or at the bases of boulders and in crevices. [5] It usually prefers igneous rocks. It is found from 0 to 1,600 meters (0.0 to 5,200 ft) in elevation. [2]
Myriopteris clevelandii is classified as globally vulnerable (G3) by NatureServe. It faces few distinct threats, but its natural range is limited. [1]
Myriopteris covillei, formerly known as Cheilanthes covillei, is a species of cheilanthoid fern known by the common name Coville's lip fern. Coville's lip fern is native to the southwestern United States and Mexico.
Myriopteris cooperae, formerly Cheilanthes cooperae, is a species of lip fern known by the common name Mrs. Cooper's lip fern, or simply Cooper's lip fern. Its leaves grow in clusters and are highly dissected into oblong segments, rather than the beadlike segments found in some other members of the genus. The axes of the leaves are dark and covered in long, flattened hairs. It is endemic to California, where it grows in rocky habitats, usually over limestone. The species was named in honor of its collector, Sarah Paxson Cooper; according to Daniel Cady Eaton, who described it in 1875, it was the first fern species to be named for a female botanist.
Myriopteris gracillima, formerly known as Cheilanthes gracillima, is a species of lip fern known by the common name lace lip fern. It is native to western North America, where it grows in rocky habitat from British Columbia to California to Montana.
Myriopteris intertexta, formerly Cheilanthes intertexta, is a species of lip fern known by the common name coastal lip fern. It is native to montane California and western Nevada, Oregon east of the Cascades, and with a disjunct population in central Utah. It grows in dry rocky habitats in sun, typically in rock cracks with little or no soil.
Myriopteris tomentosa, formerly known as Cheilanthes tomentosa, is a perennial fern known as woolly lipfern. Woolly lipfern is native to the southern United States, from Virginia to Arizona and Georgia, and Mexico.
Myriopteris lanosa, the hairy lip fern, is a moderately-sized fern of the eastern United States, a member of the family Pteridaceae. Its leaves and stem are sparsely covered in hairs, but lack scales, hence its common name. One of the cheilanthoid ferns, it was usually classified in the genus Cheilanthes until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows in shallow, dry, soil, often in rocky habitats.
Myriopteris alabamensis, the Alabama lip fern, is a moderately-sized fern of the United States and Mexico, a member of the family Pteridaceae. Unlike many members of its genus, its leaves have a few hairs on upper and lower surfaces, or lack them entirely. One of the cheilanthoid ferns, it was usually classified in the genus Cheilanthes as Cheilanthes alabamensis until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows in shade on limestone outcrops.
Myriopteris maxoniana is a species of cheilanthoid fern endemic to the Mexican state of Tamaulipas. It is known only from one collection. It closely resembles Myriopteris longipila and was not described as a distinct taxon until 2004.
Myriopteris aemula, the Texas lip fern or rival lip fern, is a moderately-sized fern of Texas and Mexico, a member of the family Pteridaceae. Unlike many members of its genus, its leaves have a few hairs on upper and lower surfaces, or lack them entirely. One of the cheilanthoid ferns, it was usually classified in the genus Cheilanthes as Cheilanthes aemula until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows on limestone rock.
Myriopteris yatskievychiana is a small fern endemic to the Mexican state of Sonora, a member of the family Pteridaceae. It is known only from a single collection. While superficially similar to golden lip fern, a widespread species in Mexico, differences in its coating of hairs and its small size make it distinctive. One of the cheilanthoid ferns, it was usually classified in the genus Cheilanthes until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes.
Myriopteris allosuroides is a moderately-sized fern endemic to Mexico, a member of the family Pteridaceae. Unlike many members of its genus, its rachides are grooved on the upper surface and largely free of hairs or scales. One of the cheilanthoid ferns, it was usually classified in the genera Cheilanthes or Pellaea until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows on dry, rocky slopes over acidic, particularly basaltic, rock.
Myriopteris aurea, the golden lip fern or Bonaire lip fern, is a moderately-sized fern native to the Americas, a member of the family Pteridaceae. Unlike many members of its genus, its leaf is only modestly dissected into lobed leaflets (pinnae), which are hairy both above and below. One of the cheilanthoid ferns, until 2013 it was classified in the genus Cheilanthes as Cheilanthes bonariensis, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows on dry, rocky slopes, and ranges from Mexico, where it is common and widespread, and the southwestern United States south and east through Central and South America as far as Chile and Argentina.
Myriopteris rufa, commonly known as Eaton's lip fern, is a moderately-sized fern of Mexico and the southwestern United States, with outlying populations in Costa Rica and the Appalachian Mountains. One of the cheilanthoid ferns, it was usually classified in the genus Cheilanthes, as Cheilanthes eatonii, until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows in rocky habitats, most frequently on limestone but also sometimes on basalt or shale.
Myriopteris rawsonii, formerly known as Cheilanthes rawsonii, is a perennial fern native to Namaqualand in Southern Africa. Like many other cheilanthoid ferns, it is adapted to dry conditions, bearing a thick layer of pale hairs on the underside of its pinnate-pinnatifid leaves. It is the only African representative of its clade of cheilanthoids, the otherwise American genus Myriopteris. It spends much of the year in a dried-out, dormant state, rehydrating and putting out new growth during winter rains. Its name honors the botanist and civil servant Sir Rawson W. Rawson.
Myriopteris wootonii, formerly known as Cheilanthes wootonii, is a species of fern in the Pteridaceae family with the common name Wooton's lace fern.
Myriopteris fendleri, formerly known as Cheilanthes fendleri, is a species of fern in the Pteridaceae family with the common name Fendler's lip fern. It is native to the southwest United States and northern Mexico.
Myriopteris lindheimeri, formerly known as Cheilanthes lindheimeri, is a species of fern in the Pteridaceae family with the common name fairy swords.
Myriopteris chipinquensis is a fern endemic to Mexico, a member of the family Pteridaceae. One of the cheilanthoid ferns, it was classified in the genus Cheilanthes until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It grows in oak-pine forests of the Sierra Madre Oriental, often with the very similar and closely related Myriopteris tomentosa.
Myriopteris cinnamomea is a Central American fern. It is very similar to M. mickelii of southern Mexico and to other species in the "alabamensis clade" of Myriopteris. It is modestly sized, with leaves not more than 30 centimeters (12 in) long. They are divided into pinnae, which in turn are divided into lobed pinnules, and covered with pale brown hairs on the underside.
Myriopteris mickelii is a fern endemic to southern Mexico. It is very similar to M. cinnamomea of Central America, from which it was separated when described in 1980, and to other species in the "alabamensis clade" of Myriopteris. Named after the pteridologist John T. Mickel, it grows on dry, lightly shaded slopes and banks.