Asplenium montanum

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Mountain spleenwort
Asplenium montanum specimen.jpg
Status TNC G5.svg
Secure  (NatureServe) [1]
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Aspleniineae
Family: Aspleniaceae
Genus: Asplenium
Species:
A. montanum
Binomial name
Asplenium montanum
Synonyms
  • Chamaefilix montana(Willd.) Farw.
  • Athyrium montanum(Willd.) Shafer , nom. illeg. hom.

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.

Contents

Description

Asplenium montanum is a small, evergreen fern which grows in tufts. [2] The leaves are bluish-green and highly divided, proceeding from a long and often drooping stalk. A. montanum is monomorphic, with no difference in form between sterile and fertile fronds. [2]

The horizontal rhizomes, which are about 1 millimeter across, [3] may curve upward. They are not branched, but as new plants can form at root tips, a tightly packed cluster of stems may give the appearance of branching. The rhizomes are covered in dark brown, narrowly triangular scales, from 2 to 4 millimeters (0.1 to 0.2 in) long and from 0.2 to 0.4 millimeters across, with untoothed edges. [4] They are strongly clathrate (bearing a lattice-like pattern). [3]

The stipe (the stalk of the leaf, below the blade) is dark brown to purplish-black and shiny at the base, gradually turning dull green as it ascends to the leaf blade. The stipe is from 2 to 11 centimeters (0.8 to 4.3 in) long, and may be from 0.5 to 1.5 times the length of the blade. Dark, narrowly lance-shaped scales and tiny hairs are present only at the very base of the stipe, [4] which is slender and fragile, [2] and lacks wings. [3]

The leaf blade is thick and hairless, [4] and of a dark blue-green color; [5] the rachis (leaf axis), like the stipe, is a dull green, with occasional hairs. [4] The blade is triangular or lance-shaped, with a squared-off or slightly rounded base and a pointed tip. It ranges from 2 to 11 centimeters (0.8 to 4.3 in) long and from 1 to 7 centimeters (0.4 to 2.8 in) wide, occasionally as wide as 10 centimeters (3.9 in). [4] [6] The blade varies from pinnate-pinnatifid to bipinnate-pinnatifid; that is, it is cut into lobed pinnae, and sometimes the pinnae themselves are cut into lobed pinnules. There are four to ten pairs of widely spaced pinnae per leaf, each of which is triangular to lance-shaped, with coarse incisions in the edges, [4] which cut them into pinnules or deep lobes, [2] and a rounded to angled base. [4] The pinnules are indented, but not further cut. [2] The longest pinnae are those nearest the base of the leaf, which range from 6 to 35 millimeters (0.2 to 1.4 in) long and from 4 to 20 millimeters (0.2 to 0.8 in) across. The veins in the leaf do not form a meshwork, and are obscure. [4]

Asplenium montanum sori beneath a fertile frond Asplenium montanum sori.JPG
Asplenium montanum sori beneath a fertile frond

On fertile fronds, from 1 to 15 elliptical or narrow sori can be found on the underside of each pinna. [2] [4] They are 0.5 to 1.5 millimeters long, covered by translucent, pale tan indusia with somewhat jagged edges. [7] Each sporangium holds 64 spores. The species has a chromosome number of 2n = 72 in the sporophyte; it is a diploid. [4]

Identification

The dark bluish-green color and the widely spaced, deeply cut and indented pinnae differentiate A. montanum from most related species. The pinnae of Bradley's spleenwort (A. bradleyi) are toothed and less deeply cut, and the dark color of the stipe continues partway up the rachis in that species. [8] Wall-rue (A. ruta-muraria) has a green stipe, [2] and its pinnae have longer stalks and are broadest near the tip. [8] Wherry's spleenwort (A. × wherryi), a hybrid between Bradley's spleenwort and mountain spleenwort, is intermediate between its parents. When compared with mountain spleenwort, the blade of Wherry's spleenwort is lance-shaped, rather than triangular; the upper parts of the blade are not as deeply cut; and the dark color of the stipe extends to the beginning of the rachis. [9]

Taxonomy

This fern was at first identified by André Michaux, in 1803, as black spleenwort ( Asplenium adiantum-nigrum ). [10] Carl Ludwig Willdenow recognized and described it as a separate species, which he named Asplenium montanum, in 1810. [11] In 1901, John A. Shafer attempted to transfer it to the genus Athyrium as Athyrium montanum, [12] but this name is illegitimate as a later homonym of Athyrium montanum (Lam.) Röhl. ex Spreng. The species was segregated from Asplenium as Chamaefilix montana by Oliver Atkins Farwell in 1931. [13] The change was not widely accepted and current authorities do not recognize this segregate genus. [4]

A global phylogeny of Asplenium published in 2020 divided the genus into eleven clades, [14] which were given informal names pending further taxonomic study. A. montanum belongs to the "Onopteris subclade" of the "Pleurosorus clade". [15] The Pleurosorus clade has a worldwide distribution; members are generally small and occur on hillsides, often sheltering among rocks in exposed habitats. The Onopteris subclade has Aspidium-type gametophytes. [16] The closest relatives of A. montanum within the subclade are A. onopteris and its allopolyploid descendant, A. adiantum-nigrum. [15]

Varieties

In 1974, Timothy Reeves described an unusual population of A. montanum from the Shawangunk Mountains. Having used chromatography to show that it was not a hybrid, he interpreted it as a new form, Asplenium montanum forma shawangunkense. In this form, as contrasted with the usual forma montanum, the leaf blade is yellow-green, the fronds continue highly dissected to the apex and do not come to a pointed tip, the fronds are shorter and more highly dissected than usual, and all fronds are sterile. [5]

Hybrids

Asplenium montanum readily forms hybrids with a number of other species in the "Appalachian Asplenium complex". In 1925, Edgar T. Wherry noted the similarities between A. montanum, lobed spleenwort (A. pinnatifidum), and Trudell's spleenwort (A. × trudellii), [17] and in 1936 concluded that Trudell's spleenwort was a hybrid between the first two. [18] In 1951, Herb Wagner, while reviewing Irene Manton's Problems of Cytology and Evolution in the Pteridophyta, suggested in passing that A. pinnatifidum itself might represent a hybrid between A. montanum and the American walking fern, Camptosorus rhizophyllus (now A. rhizophyllum). [19]

In 1953, he reported preliminary cytological studies on the Aspleniums and suggested that A. montanum had crossed with ebony spleenwort (A. platyneuron) to yield Bradley's spleenwort (A. bradleyi), noting that D. C. Eaton and W. N. Clute had already made tentative suggestions along those lines. He also made chromosome counts of A. × trudellii, which had been classified by some simply as a variety of A. pinnatifidum. As A. pinnatifidum proved to be a tetraploid while A. montanum was a diploid, a hybrid between them would be a triploid, and Wagner showed that this was in fact the case for A. × trudellii. [20] His further experiments, published the following year, strongly suggested that both A. bradleyi and A. pinnatifidum were allotetraploids, the product of hybridization between A. montanum and another Asplenium to form a sterile diploid, followed by chromosome doubling that restored fertility. [21]

These cytotaxonomic findings were supported by subsequent chromatographic studies. A. montanum was shown to produce a pattern of seven substances chromatographically distinct from those produced by the other diploid members of the Appalachian Asplenium complex. These substances were present in the chromatograms of all tested hybrids believed to descend from A. montanum at one or more removes: A. bradleyi, A. × gravesii, A. × kentuckiense, A. pinnatifidum, A. × trudellii, and A. × wherryi. [22] Four of the compounds present in the chromatograms of A. montanum and its descendants, fluorescing gold-orange under ultraviolet light, were subsequently identified as the xanthonoids mangiferin, isomangiferin, and their O-glucosides. [23] The other two were identified as kaempferol derivatives, but could not be more precisely determined due to lack of material; the last was a trace compound which could not be studied. [24] A chloroplast phylogeny has suggested that A. montanum is the maternal ancestor of A. bradleyi. [16]

The allotetraploid hybrid species derived from A. montanum can backcross with A. montanum to form triploid hybrids. The backcross hybrid between A. montanum and A. pinnatifidum is A. × trudellii, as suggested by Wherry. He also collected specimens of the backcross hybrid between A. montanum and A. bradleyi from a cliff near Blairstown, New Jersey in 1935. [25] The hybrid received no further attention until 1961, when it was described and named in Wherry's honor as Wherry's spleenwort (A. × wherryi). [9] The sterile diploid A. montanum × platyneuron, precursor to A. bradleyi, was collected in 1972 at Crowder's Mountain, Georgia; [26] A. montanum × rhizophyllum, precursor to A. pinnatifidum, has never been found.

Distribution and habitat

Illustration from How to Know the Ferns by Frances Theodora Parsons (1899); illustration by Marion Satterlee Asplenium montanum parsons.png
Illustration from How to Know the Ferns by Frances Theodora Parsons (1899); illustration by Marion Satterlee

One of the "Appalachian spleenworts", A. montanum is found in the Appalachian Mountains from Vermont and Massachusetts southwestward to Tennessee, Alabama, and Georgia, and to a lesser extent in the Ohio Valley in Ohio, Indiana, and Kentucky. [27] Arkansas populations were discovered in Garland County and Stone County in 2002 and 2008, respectively. [28] One outlying population in Missouri, collected in 1960, is considered historical; [27] it is represented by one specimen collected near Graham Cave and has never been relocated. The site is thought to have been destroyed by road construction. [4] [28] A collection by Farwell from the Keweenaw Peninsula of Michigan was considered valid by M. L. Fernald, but is of questionable authenticity; the population has never been relocated. [29]

Asplenium montanum grows on acidic rocks such as sandstone, in crevices [4] [7] into which moisture seeps from within the rock strata. It has been found at altitudes up to 2,000 meters (7,000 ft). [4] Like the closely related A. bradleyi, A. montanum requires that the thin soil in its favored crevices be subacid (pH 4.5–5.0) to mediacid (pH 3.5–4.0), and it is intolerant of calcium. [30] [31] This habitat is unfavorable to most other plants, but its allotetraploid descendants and their backcross hybrids may occur alongside it. [4]

Ecology and conservation

Asplenium montanum is globally secure, but threatened at the edges of its range. It is known only historically from Missouri, and NatureServe considers it critically imperiled in Indiana, Massachusetts, Rhode Island, and Vermont, imperiled in New Jersey, and imperiled to vulnerable in Connecticut and New York. [1] The principal threat to New York populations is rock climbing. [32]

Cultivation

Asplenium montanum may be cultivated outdoors or in a terrarium. In either case, the soil used should be amended with chips of acidic rock. [7]

Notes and references

Related Research Articles

<i>Asplenium</i> Genus of ferns in the family Aspleniaceae

Asplenium is a genus of about 700 species of ferns, often treated as the only genus in the family Aspleniaceae, though other authors consider Hymenasplenium separate, based on molecular phylogenetic analysis of DNA sequences, a different chromosome count, and structural differences in the rhizomes. The type species for the genus is Asplenium marinum.

<i>Asplenium platyneuron</i> Species of fern

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.

<i>Asplenium rhizophyllum</i> Species of fern in the family Aspleniaceae

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".

<i>Asplenium septentrionale</i> Species of fern in the family Aspleniaceae

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.

<i>Myriopteris cooperae</i> Species of 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.

The fern genus Asplenium is well known for its hybridization capacity, especially in temperate zones.

<i>Asplenium pinnatifidum</i> Species of fern in the family Aspleniaceae

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.

<i>Asplenium bradleyi</i> Species of fern in the family Aspleniaceae

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.

<i>Asplenium <span style="font-style:normal;">×</span> ebenoides</i> Hybrid fern in the family Aspleniaceae

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.

<i>Asplenium resiliens</i> Species of fern in the family Aspleniaceae

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.

<i>Asplenium tutwilerae</i> Species of fern in the family Aspleniaceae

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 × wherryi, known as Wherry's spleenwort, is a rare hybrid fern of the Appalachian Mountains. The sterile triploid offspring of mountain spleenwort (A. montanum) and Bradley's spleenwort (A. bradleyi), it is known from a few sites where those species grow together. First collected by Edgar T. Wherry in 1935, it was largely ignored until a new colony was found in 1961, and the species was named in his honor.

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.

<i>Asplenium <span style="font-style:normal;">×</span> trudellii</i> Species of fern

Asplenium × trudellii, commonly known as Trudell's spleenwort, is a rare hybrid fern of the eastern United States, first described in 1925. It is formed by the crossing of mountain spleenwort (A. montanum) with lobed spleenwort (A. pinnatifidum). Trudell's spleenwort is intermediate in form between its two parents, and is generally found near them, growing on exposed outcrops of acidic rock. While A. × trudellii is triploid and sterile, there is some evidence that it can occasionally reproduce apogamously.

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 × boydstoniae, commonly known as Boydston's spleenwort, is a rare, sterile, hybrid fern. It is formed by the crossing of Tutwiler's spleenwort (A. tutwilerae) with ebony spleenwort (A. platyneuron). The hybrid was produced in culture in 1954. It was not discovered in the wild until 1971, when it was found by Kerry S. Walter at Havana Glen, Alabama, the only known wild site for Tutwiler's spleenwort. Walter named it for Kathryn E. Boydston, an expert in fern culture. Except for the tip of its leaf blade, it largely resembles its ebony spleenwort parent.

Asplenium arcanum is a fern known from western Mexico and Nicaragua.

<i>Myriopteris lanosa</i> Species of fern

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.

<i>Myriopteris aurea</i> Species of fern

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.

<i>Myriopteris rufa</i> Species of fern

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.

References

Works cited