Asplenium septentrionale | |
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Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Order: | Polypodiales |
Suborder: | Aspleniineae |
Family: | Aspleniaceae |
Genus: | Asplenium |
Species: | A. septentrionale |
Binomial name | |
Asplenium septentrionale | |
Subspecies | |
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Synonyms | |
Asplenium septentrionale is a species of fern known by the common names northern spleenwort and forked spleenwort. It is native to Europe, Asia and western North America, where it grows on rocks. Its long, slender leaves give it a distinctive appearance. Three subspecies exist, corresponding to a tetraploid and a diploid cytotype and their triploid hybrid.
Asplenium septentrionale is a small fern which grows in dense clusters superficially resembling tufts of grass. The long, dark stems support narrow, leathery leaf blades, which may appear slightly forked at the tip. The fronds are monomorphic, with no difference in shape or size between fertile and sterile fronds. [2]
The rhizome from which the many leaves of each plant spring is about 1 millimetre in diameter, and covered with scales. The scales are narrowly triangular, and range in color from black to a dark reddish brown. They are 2 to 4 millimetres (0.08 to 0.2 in) long, 0.3 to 0.6 millimetres wide, and entire (untoothed) at the edges. The stipes (the stalks of the leaf, below the blade) are reddish brown at the base, fading to green above. They range from 2 to 13 centimetres (0.8 to 5 in) in length, and are about 2 to 5 times the length of the leaf blade itself. [2]
The leaf blades are narrow, with parallel edges following the stem, ranging from 0.5 to 4 centimetres (0.2 to 2 in) in length and 0.1 to 0.4 centimetres (0.04 to 0.2 in) in width. They come to a point at both base and tip, and have a leathery texture. They are often divided into pinnae (leaflets) near the tip, usually two (but sometimes four). These pinnae are sharply angled towards the tip of the blade, giving it a forked appearance. They also come to a point at the tip, and have a few widely spaced, irregular teeth. The rachis (central axis of the leaf) is green and shiny, and the leaves, including the rachis, are free of hairs or scales. Fertile and sterile fronds are the same in appearance; in fertile fronds, the sori are linear, parallel to the edges of the pinna, usually two or more per pinna. [2] The sori are covered by thin, pale tan indusia, with entire edges. [3] It contains 64 spores per sporangium, and most sporophytes have a chromosome number of 2n=144 (a tetraploid). [2]
Individual plants have an abundant number of leaves, forming dense tufts from a rhizome of about 1 millimetre in diameter, and sometimes mats on flat rocks. [2] [3]
A. septentrionale is easily distinguished from other related ferns by its narrow blades, often forked at the tip. The presence of sori on fertile leaves distinguishes it from the vegetative material of a grass or sedge.
Originally named Acrostichum septentrionale by Linnaeus in 1753, the species was placed in genus Asplenium by Georg Franz Hoffmann in 1796. [2] The species has twice been placed in genera segregated from Asplenium : as Chamaefilix septentrionalis by Farwell in 1931, [4] and as Tarachia septentrionalis by Momose in 1960. [5] Neither combination was widely accepted and current authorities do not recognize these segregate genera. [2] A global phylogeny of Asplenium published in 2020 divided the genus into eleven clades, [6] which were given informal names pending further taxonomic study. A. septentrionale belongs to the "A. septentrionale subclade" of the "Schaffneria clade". [7] The Schaffneria clade has a worldwide distribution, and members vary widely in form and habitat. [8] Most members of the A. septentrionale subclade have small leaves with a long green stalk, entire or divided into two or three segments, and always grow on rocks. [9] Other members of the subclade include the European A. seelosii and A. celtibericum and an undescribed species from Venezuela. [7]
In 1980, the discovery of diploid specimens prompted the division of the species into A. septentrionale subsp. septentrionale, the tetraploid population, and A. septentrionale subsp. caucasicum, the diploid population. The diploids are limited to southwest Asia, while the tetraploid population has a circumboreal distribution. The tetraploid subspecies is believed to have arisen from the diploid subspecies by autopolyploidy. The two subspecies are very similar in form, but can be distinguished by the narrower blades of A. septentrionale subsp. caucasicum (to 0.12 centimetres (0.05 in) wide) and its smaller spores (averaging 34 μm in diameter). In 2003, Ronald Viane and Tadeus Reichstein elevated this subspecies to a species, as A. caucasicum, and described a sterile triploid formed by hybridization of the tetraploid and diploid, identifiable by its abortive spores. They designated the hybrid A. × dirense. [10] However, in his taxonomic revision of Indian pteridophytes, Christopher Fraser-Jenkins reduced all of these taxa to subspecific rank, with the hybrid A. × dirense becoming the nothosubspecies A. septentrionale subsp. rehmanii.
Asplenium septentrionale can hybridize with maidenhair spleenwort (A. trichomanes) to form the hybrid A. × alternifolium, with A. foreziense to form A. × costei, with black spleenwort (A. adiantum-nigrum) to form A. × contrei, with A. obovatum subsp. lanceolatum to form A. × souchei, and with wall-rue (A. ruta-muraria) to form A. × murbeckii and A. × tavelio. A. × murbeckii can back-cross with A. septentrionale to form A. × hungaricum.
Asplenium septentrionale subsp. septentrionale is found in Europe and Asia from the Macaronesian Islands east through Europe, western Asia, the former USSR, northern India, parts of western and central China (Xinjiang, Tibet, and Shaanxi), and Taiwan. [10] It is also found in North America, principally in the Rocky Mountains of the western United States, but also in the Black Hills of South Dakota–Wyoming. It also occurs in the mountains of the Cascade Range and the Sierra Nevada, from Oregon south through California into Baja California. Populations occur to the eastward in western Texas and near the tip of the Oklahoma Panhandle, and two very disjunct stations have been located on shale in Monroe County and Hardy County, West Virginia. [2] [3]
Asplenium septentrionale subsp. caucasicum has been reported from Georgia, Iran, Pakistan and Turkey. The type specimen of A. septentrionale subsp. rehmanii came from Pakistan, near the Lowari Pass. [10]
Asplenium septentrionale is epipetric and can be found in crevices of rocks, around boulders, and on cliffs. [2] [3] It can be found on a variety of substrates, [2] including granitic rocks [3] and limestone. [11] The West Virginia stations are on shale. [3] In the United States, it can be found growing at altitudes from 700 to 2,900 metres (2,300 to 9,500 ft). [2]
Asplenium septentrionale is hardy to USDA Zone 4. It is best grown in partial sun or medium light in moist to dry potting mix. [11] [3] It requires good drainage and little watering. Sensitive to root disturbance, it is difficult to transplant and establish, and is rarely sold by nurseries. [11]
Asplenium platyneuron, commonly known as ebony spleenwort or brownstem spleenwort, is a fern native to North America east of the Rocky Mountains. It takes its common name from its dark, reddish-brown, glossy stipe and rachis, which support a once-divided, pinnate leaf. The fertile fronds, which die off in the winter, are darker green and stand upright, while the sterile fronds are evergreen and lie flat on the ground. An auricle at the base of each pinna points towards the tip of the frond. The dimorphic fronds and alternate, rather than opposite, pinnae distinguish it from the similar black-stemmed spleenwort.
Asplenium scolopendrium, commonly known as the hart's-tongue fern, is an evergreen fern in the family Aspleniaceae native to the Northern Hemisphere.
Asplenium rhizophyllum, the (American) walking fern, is a frequently-occurring fern native to North America. It is a close relative of Asplenium ruprechtii which is found in East Asia and also goes by the common name of "walking fern".
Asplenium ruprechtii, which goes by the common name Asian walking fern, is a rare, hardy, low-lying fern native to East Asia. It is a close relative of Asplenium rhizophyllum which is found in North America and also goes by the common name of walking fern. The species should not be confused with Asplenium sibiricum which is a synonym of Diplazium sibiricum.
Asplenium ceterach, also known as the rustyback fern, is a fern species in the spleenwort family Aspleniaceae.
Asplenium trichomanes, the maidenhair spleenwort, is a small fern in the spleenwort genus Asplenium. It is a widespread and common species, occurring almost worldwide in a variety of rocky habitats. It is a variable fern with several subspecies.
Asplenium vespertinum is a species of fern known by the common name western spleenwort. It is native to southern California and Baja California, where it grows in moist, shady, rocky places, such as the shadows beneath cliff overhangs.
Asplenium montanum, commonly known as the mountain spleenwort, is a small fern endemic to the eastern United States. It is found primarily in the Appalachian Mountains from Vermont to Alabama, with a few isolated populations in the Ozarks and in the Ohio Valley. It grows in small crevices in sandstone cliffs with highly acid soil, where it is usually the only vascular plant occupying that ecological niche. It can be recognized by its tufts of dark blue-green, highly divided leaves. The species was first described in 1810 by the botanist Carl Ludwig Willdenow. No subspecies have been described, although a discolored and highly dissected form was reported from the Shawangunk Mountains in 1974. Asplenium montanum is a diploid member of the "Appalachian Asplenium complex," a group of spleenwort species and hybrids which have formed by reticulate evolution. Members of the complex descended from A. montanum are among the few other vascular plants that can tolerate its typical habitat.
Asplenium pinnatifidum, commonly known as the lobed spleenwort or pinnatifid spleenwort, is a small fern found principally in the Appalachian Mountains and the Shawnee Hills, growing in rock crevices in moderately acid to subacid strata. Originally identified as a variety of walking fern, it was classified as a separate species by Thomas Nuttall in 1818. It is believed to have originated by chromosome doubling in a hybrid between walking fern and mountain spleenwort, producing a fertile tetraploid, a phenomenon known as alloploidy; however, the hypothesized parental hybrid has never been located. It is intermediate in morphology between the parent species: while its leaf blades are long and tapering like that of walking fern, the influence of mountain spleenwort means that the blades are lobed, rather than whole. A. pinnatifidum can itself form sterile hybrids with several other spleenworts.
Asplenium bradleyi, commonly known as Bradley's spleenwort or cliff spleenwort, is a rare epipetric fern of east-central North America. Named after Professor Frank Howe Bradley, who first collected it in Tennessee, it may be found infrequently throughout much of the Appalachian Mountains, the Ozarks, and the Ouachita Mountains, growing in small crevices on exposed sandstone cliffs. The species originated as a hybrid between mountain spleenwort and ebony spleenwort ; A. bradleyi originated when that sterile diploid hybrid underwent chromosome doubling to become a fertile tetraploid, a phenomenon known as allopolyploidy. Studies indicate that the present population of Bradley's spleenwort arose from several independent doublings of sterile diploid hybrids. A. bradleyi can also form sterile hybrids with several other spleenworts.
Asplenium anceps is a diploid fern of family Aspleniaceae and one of the ancestors of the ferns that form the trichomanes complex. It lives exclusively in the three northernmost archipelagoes of the Macaronesian region, that is, is an endemic macaronesian fern. Its fronds are leathery and plastic and rachis is very thick, bright reddish brown and is traversed throughout its length of three wings, two on the upper surface to draw a groove and a third on the lower surface which is characteristic and unique to this species, since all other species of the trichomanes complex without. A typical feature of this fern, which he shares with all its hybrid offspring is the existence of a small atrium on the basis of medium and less pinnae directed toward the apex of the blade with one or two sori on its underside.
Asplenium viride is a species of fern known as the green spleenwort because of its green stipes and rachides. This feature easily distinguishes it from the very similar-looking maidenhair spleenwort, Asplenium trichomanes.
Asplenium onopteris, known as the Irish spleenwort or western black spleenwort, is a species of fern mostly found throughout the Mediterranean Basin but also around the Eastern Atlantic.
Asplenium × ebenoides is a hybrid fern native to eastern North America, part of the "Appalachian Asplenium complex" of related hybrids. The sterile offspring of the walking fern (A. rhizophyllum) and the ebony spleenwort (A. platyneuron), A. × ebenoides is intermediate in morphology between its two parents, combining the long, narrow blade of A. rhizophyllum with a dark stem and lobes or pinnae similar to those of A. platyneuron. While A. × ebenoides is generally sterile, fertile specimens with double the number of chromosomes are known from Havana Glen, Alabama. These fertile allotetraploids were reclassified as a separate species named A. tutwilerae in 2007, retaining the name A. × ebenoides for the sterile diploids only.
Asplenium resiliens, the blackstem spleenwort or little ebony spleenwort, is a species of fern native to the Western Hemisphere, ranging from the southern United States south to Uruguay, including parts of the Caribbean. Found on limestone substrates, it is named for its distinctive purplish-black stipe and rachis. A triploid, it is incapable of sexual reproduction and produces spores apogamously. First described by Martens and Galeotti in 1842 under the previously used name Asplenium parvulum, the species was given its current, valid name by Kunze in 1844. Several similar species are known from the tropics; A. resiliens may have arisen from these species by reticulate evolution, but precise relationships among the group are not yet certain.
Asplenium tutwilerae is a rare epipetric fern found only in Hale County, Alabama, United States. A. tutwilerae is a fertile allotetraploid, formed by the chromosomal doubling of a specimen of the sterile diploid A. × ebenoides, a hybrid of A. platyneuron and A. rhizophyllum. Except for its spores, which are fertile rather than malformed, A. tutwilerae is essentially identical to A. × ebenoides and was described as part of that species until 2007. It is named in honor of Julia Tutwiler, who discovered the only known wild population at Havana Glen in 1873.
Asplenium × gravesii, commonly known as Graves' spleenwort, is a rare, sterile, hybrid fern, named for Edward Willis Graves (1882–1936). It is formed by the crossing of Bradley's spleenwort (A. bradleyi) with lobed spleenwort (A. pinnatifidum). It is only found where its parent species are both present; in practice, this proves to be a few scattered sites in the Appalachian Mountains, Shawnee Hills, and Ozarks, reaching perhaps its greatest local abundance around Natural Bridge State Resort Park. Like its parents, it prefers to grow in acid soil in the crevices of sandstone cliffs.
Asplenium × kentuckiense, commonly known as Kentucky spleenwort, is a rare, sterile, hybrid fern. It is formed by the crossing of lobed spleenwort (A. pinnatifidum) with ebony spleenwort (A. platyneuron). Found intermittently where the parent species grow together in the eastern United States, it typically grows on sandstone cliffs, but is known from other substrates as well.
Asplenium appendiculatum, ground spleenwort, is a common native fern to Australia and New Zealand. It usually grows in cool damp conditions, among rocks, on logs or as an epiphyte.
Asplenium fontanum, commonly known as fountain spleenwort or smooth rock spleenwort, is a species of fern in the family Aspleniaceae, native to rocky areas in Western Europe.