Myriopteris aurea | |
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Myriopteris aurea growing in Peru, showing pinnate-pinnatifid leaves and hairs on both surfaces | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Order: | Polypodiales |
Family: | Pteridaceae |
Genus: | Myriopteris |
Species: | M. aurea |
Binomial name | |
Myriopteris aurea | |
Synonyms | |
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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.
Leaf bases are closely spaced along the horizontal rhizome, which is variously described as being 3 millimeters (0.1 in) [2] or 4 to 8 millimeters (0.2 to 0.3 in) in diameter. [3] The rhizome bears scales, which are linear to lanceolate, with untoothed [4] [5] or very slightly toothed margins. [6] They are bi-colored, with a shiny central stripe red-brown [7] or shiny chestnut-brown [6] to black in color [2] and narrow light-brown margins, [3] and measure 3 millimeters (0.1 in) long. [2] They are slightly twisted and strongly pressed against the rhizome. [3]
The fronds spring up in clusters; [2] unlike many ferns, they do not unfold as fiddleheads (noncircinate vernation). [3] [8] They range from 10 to 60 centimeters (3.9 to 24 in) long, [3] [5] sometimes up to 75 centimeters (30 in), [2] and are 0.5 to 3.5 centimeters (0.20 to 1.4 in) broad. [6] The stipe (the stalk of the leaf, below the blade) represents from one-sixth to one-third of the total length of the leaf, [2] [5] and is 3 to 15 centimeters (1.2 to 5.9 in) long. [6] The upper surface of the stipe is rounded and not grooved. [2] [3] [6] The stipe is shiny, dark chestnut brown to black [2] [6] or purplish-black in color. [6] [7] It is covered in straight, white to tan hairs, 2 millimeters (0.08 in) long and more or less pressed against the stipe. [2] [6]
The leaf blades are linear to lanceolate [2] or elliptical, [5] and pinnate-pinnatifid (cut into deeply lobed pinnae), [2] [6] much less dissected than most Myriopteris species. [9] It is from 1 to 4 centimeters (0.4 to 2 in) wide. [3] The rachis (leaf axis) is densely covered in pubescent hairs, but lacks scales. [3] [6] From 15 to 44 pairs of pinnae are present, [2] attached directly to the rachis or with a short stalk. [6] Each pinna is approximately equilateral in shape, [3] has from 3 to 8 pairs of lobes, which may be cut as shallowly as one-fourth or as deeply as three-fourths of the distance to the costa (pinna axis). The number of lobes and deepness of cutting can vary a great deal between individuals. [2] They are joined to the rachis by a distinct stalk; the dark color of the rachis extends into the stalk but terminates abruptly in a swollen node covered with hairs. [3] The lowest pair of pinnae is slightly smaller than the one above. [3] At the other end of the frond, the pinnae gradually taper to an acute or obtuse apex. [7] The leaf tissue is parchment-like to almost leathery. [6] The upper surface of the pinnae is covered with scattered to abundant hairs, stiff, flattened against the surface, one-celled, and about 2 millimeters (0.08 in) long. [2] They are pale golden-tan in color. [7] [6] The lower surface of the pinnae is also covered in hairs, matted so thickly as to hide the leaf tissue, which vary from white, particularly when young, [5] to rusty red in color. [2] [6] They do not curl up when dried out. [7]
On fertile fronds, the sori are located at the ends of veins near the margin of the leaf, [2] [6] forming a more or less continuous zone adjacent to the margin, [3] which curls back slightly but does not form a distinct false indusium to protect them. [2] The recurved margin is 0.05 to 0.25 millimeters (0.0020 to 0.0098 in) wide. [3] It is somewhat thinner and more delicate than the rest of the leaf tissue, though not quite hyaline. [5] [6] Each sporangium contains 32 dark brown to black spores. [2] [3] The vast majority of M. aurea individuals thus far examined are apogamous triploids, with a chromosome number of 90 present in both sporophyte and gametophyte. [2] [3] A few populations forming 64 spores per sporangium have reportedly been found, and are presumed to be sexual diploids. [10]
Among its congeners M. aurea is most similar to M. yatskievychiana , known only from Sonora which is smaller and has dense white (rather than rusty) hairs on the underside of the leaf. [11] The pinnate-pinnatifid leaf blades distinguish these two species from the rest of the genus, the other species being more highly dissected. [9] M. aurea superficially resembles some species of Astrolepis , such as A. sinuata , which have short, deeply lobed pinnae, but they bear the stellate scales that give their genus its name rather than the hairs seen on M. aurea. [2] In the southern part of its range, it might be confused with Cheilanthes fraseri , but the latter has leaves widely spaced along the rhizome, pinnae more or less triangular (rather than equilateral), and rhizome scales light brown at the apex, rather than dark brown. [12]
The species was first described as Pteris aurea by Jean Louis Marie Poiret in Lamarck's Encyclopédie Méthodique, Botanique in 1804. He based his description on a specimen collected in Peru by Joseph de Jussieu. [13] The specific epithet aurea, meaning "golden", [14] evidently refers to the "yellow, almost golden" hairs covering the upper surface of the leaves. [13]
In 1810, Carl Ludwig Willdenow recognized Pteris aurea in the 5th edition of Species Plantarum , [15] but also described a new species, Acrostichum bonariense, based on material collected near Buenos Aires (Latinized as "Bonaria"), an illustration of which was published by Johann Amman in 1738. [16] [17] Willdenow evidently did not have access to any of Amman's material, and so failed to recognize the similarity between Amman's fern and Pteris aurea. [18] Confusion was compounded in 1822, when Heinrich Friedrich Link described the species yet again, based on a Humboldt and Bonpland specimen which Willdenow had labeled Cheilanthes ferruginea but had omitted from Species Plantarum. [19] The specific epithet ferruginea means "rusty-reddish", [20] perhaps referring to the color of the hairs below, so described by Willdenow for A. bonariense. [21]
In his Reliquae Haenkeanae in 1825, Carl Borivoj Presl recognized that A. bonariensis and C. ferruginea were the same species, but (illegitimately) invented the new name of Notholaena rufa to encompass both. [22] Placement in Notholaena reflected the placement of the sori at or near the margin of the leaf, combined with an absence of a distinct false indusium formed by that margin. [23] The specific epithet rufa, meaning "reddish", [24] corresponds to his description of the color of hairs on both sides of the leaf. Unfortunately, Presl also included the farinose N. ferruginea (Desv.) Desv. and N. tomentosa Desv. (now considered N. trichomanoides and C. hypoleuca , respectively) within his concept of the species. He distinguished between var. minor of the species, with dense hairs on the upper surface, and var. major, with sparse hairs on the upper surface. [22] Meanwhile, Nicaise Auguste Desvaux also recognized Notholaena, and transferred P. aurea there as N. aurea in 1827. [25] Christian Nees von Esenbeck transferred C. ferruginea to Pellaea as P. ferruginea in 1847, [26] but this was not widely followed.
William Jackson Hooker transferred C. ferruginea to Notholaena in his Species Filicum in 1864, and recognized the distinctness of N. trichomanoides, but his use of the name N. ferruginea was illegitimate as it had already been used by Desvaux, and he failed to recognize N. tomentosa as a distinct species. [27] Hooker's re-use of N. ferruginea for this species led to continued confusion with N. trichomanoides. [28] In 1906 Carl Christensen transferred the more senior A. bonariensis to Notholaena as Notholaena bonariensis, but recognized both N. ferruginea (Desv.) Desv. and N. tomentosa as synonyms. [29] This confusion appears to have led José N. Rovirosa to found another name for the species, Notholaena chiapensis, named for one of his collections in Chiapas; he noted that it was non-farinose, and thought that Hooker's name applied to the farinose species (i.e., N. trichomanoides). [30] It was not until 1939 that Charles Alfred Weatherby, who was looking for type specimens in Desvaux's collections at the Paris herbarium, identified Notholaena aurea as the most senior name for the species in that genus and unraveled the erroneous synonymies. [28]
In 1953, George R. Proctor transferred the species to Cheilanthes as Cheilanthes bonariensis, since the name Cheilanthes aurea was preoccupied. [31] However, most authors continued to place it in Notholaena. [18] One of these was Rolla M. Tryon Jr. who, in 1956, published a revision of American Notholaena incorporating material from Weatherby, who had died in 1949. Tryon accepted the species in Notholaena as N. aurea, and circumscribed it in the fashion presently accepted (including A. bonariensis, C. ferruginea, and N. chiapensis, and excluding N. ferruginea (Desv.) Desv. and N. tomentosa Desv.). He accepted as the type of A. bonariensis a specimen seen by Weatherby in Willdenow's herbarium in Berlin. [32]
Tryon noted that the generic delimitation of Notholaena and Cheilanthes, based on the traditional criterion that the latter had leaf edges curled under and modified into distinct false indusia while the former did not, was unsatisfactory, given the existence of many intermediate forms. In 1982, he and his wife Alice published an extended survey of the ferns in which they narrowed the circumscription of Notholaena, treating N. aurea as C. bonariensis. [33] This classification was widely taken up by other workers. [18] Research at the Berlin herbarium subsequently revealed that the specimen in Willdenow's herbarium came from Mexico, and did not match his original description, registering it ineligible as a type. In 2011, M. Mónica Ponce and Brigitte Zimmer designated Amman's illustration as the lectotype of A. bonariense; as there was some question as to whether the drawing represented C. bonariensis in the conventional sense or Cheilanthes buchtienii , they further designated Willdenow's Mexican specimen as an epitype. [18]
Meanwhile, 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. bonariensis; since the epithet aurea was not preoccupied, they transferred P. aurea to the genus to become Myriopteris aurea. [34] In 2018, Maarten J. M. Christenhusz transferred the species to Hemionitis as H. bonariensis (H. aurea being preoccupied), as part of a program to consolidate the cheilanthoid ferns into that genus. [35]
Further molecular studies in Myriopteris demonstrated the existence of three well-supported clades within the genus. M. aurea belongs to what Grusz et al. informally named the covillei clade. M. aurea and the similar M. yatskievychiana are sister to the rest of the clade; all other members of the clade, except M. newberryi, belong to the "core covillei" clade, with leaves finely divided into bead-like segments, quite dissimilar to M. aurea. [36]
The common names golden lip fern and Bonaire lip fern [3] [37] refer to the epithets bestowed by Poiret and Willdenow, respectively. "Lip fern" comes from the position of the sporangia at the edge or lip of the leaf, typical of the genus. [3] Lellinger, who referred to the species as N. aurea, called it golden cloak fern; [7] "cloak fern" refers to the reflexed leaf margins of Notholaena, curled back on the sporangia. [38] It has also been called slender cloak fern. [37]
Myriopteris aurea grows throughout Mexico except for Tabasco and the Yucatan Peninsula, where it is the most widespread and abundant fern in the country. [2] [39] The range extends slightly north into the United States, into Arizona, New Mexico, and Trans-Pecos Texas. [40] In the east, it is found among the Greater Antilles and in Venezuela. To the south, it extends through Central America into South America and along the Andes Mountains, [2] where it has been described as "one of the characteristic ferns of the Altiplano", [5] as far south as Chile and Argentina, and east into Brazil, Paraguay and Uruguay. [2]
The species grows on dry, rocky slopes, [2] and cliffs, [6] [41] soil banks, and shrubby hillsides. [5] [6] It tolerates a variety of different rocks, although it is comparatively uncommon on limestone. [3] It is found at altitudes from 600 to 3,800 meters (2,000 to 12,000 ft). [2] [5] [6]
The species is globally secure (G5). NatureServe does not assign conservation rankings for the three states of the United States at the northern edge of its range. [1]
Myriopteris aurea can be grown on moist-dry to dry garden soil, supplemented with sand. The soil must be well-drained and it requires high levels of light. [37] The horticulturist George Schneider, writing in 1892, spoke of it as "an old inhabitant of our gardens". [42]
Edward Palmer collected a specimen at the market in Saltillo in 1898, where he reported that a decoction was prepared from it and drunk to treat "pain in the stomach" and "coughs". [43]
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 parryi, formerly known as Cheilanthes parryi, is a species of lip fern known by the common name Parry's lip fern.
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 newberryi, formerly Cheilanthes newberryi, is a species of lip fern known by the common name Newberry's lip fern. It is native to southern California and Baja California.
Argyrochosma nivea is an Andean fern species in the family Pteridaceae.
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 myriophylla, the Central American lace fern, is a species of lip fern. Despite its common name, this species is native as far south as Argentina. It is adapted to dry areas.
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, commonly known as the lip ferns, is a genus of cheilanthoid ferns. Like other cheilanthoids, they are ferns of dry habitats, reproducing both sexually and apogamously. Many species have leaves divided into a large number of small, bead-like segments, the probable inspiration for the generic name. Hairs and/or scales are often present on both the upper and lower surfaces of the leaf, and their presence and appearance are useful in distinguishing between species. The genus is most diverse in Mexico, but species are found from southwestern Canada south to southern Chile, and one species is endemic to southern Africa.
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 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.
Argyrochosma microphylla, the small-leaf false cloak fern, is a species of fern native to New Mexico, Texas and northern Mexico. It grows on limestone rocks and cliffs, and has finely-divided leaves with small leaf segments, often folded in half when dry, which lack the white powder present on the leaf underside of many related species. First described as a species in 1869, it was transferred to the new genus Argyrochosma in 1987, recognizing their distinctness from the "cloak ferns".
Myriopteris longipila, formerly known as Cheilanthes longipila, is a species of fern endemic to Mexico. It is characterized by a dense covering of long whitish hairs.
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 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.
Argyrochosma flavens is a South American fern. It has leathery, thrice-divided leaves with dark brown axes; the leaves are coated with yellow powder below. First described as a species in 1806, it was transferred to the new genus Argyrochosma in 1996, recognizing their distinctness from the "cloak ferns". Until recently, it has usually been treated as a variety of Argyrochosma nivea under the names A. nivea var. flava or Notholaena nivea var. flava.