Asplenium rhizophyllum

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Asplenium rhizophyllum
Asplenium rhizophyllum.jpg
Asplenium rhizophyllum on rocks in the Red River Gorge, Daniel Boone National Forest, Kentucky, US
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. rhizophyllum
Binomial name
Asplenium rhizophyllum
L.
Synonyms

Antigramma rhizophylla(L.) J.Sm.
Camptosorus rhizophyllus(L.) Link

Contents

Asplenium rhizophyllum, the (American) walking fern, is a frequently-occurring fern native to North America. It is a close relative of Asplenium ruprechtii [2] (syn: Camptosorus sibiricus) which is found in East Asia and also goes by the common name of "walking fern". [3]

Description

Asplenium rhizophyllum is a small fern whose undivided, evergreen leaves and long, narrow leaf tips, sometimes curving back and rooting, give it a highly distinctive appearance. It grows in tufts, often surrounded by child plants formed from the leaf tips. The leaves of younger plants tend to lie flat to the ground, while older plants have leaves more erect or arching. [4]

Roots and rhizomes

It does not spread and form new plants via the roots. Its rhizomes (underground stems) are upright or nearly so, [2] [5] short, [4] about 1 millimetre (0.04 in) in diameter, [5] and generally unbranched. [2] [5] They bear dark brown [2] or blackish, [5] narrowly triangular [2] or lance-shaped [5] scales which are strongly clathrate (bearing a lattice-like pattern). [5] The scales are 2 to 3 millimetres (0.08 to 0.1 in) long and 0.5 to 1 millimetre (0.02 to 0.04 in) wide (occasionally as narrow as 0.2 millimetres (0.008 in)) with untoothed margins. [2]

Leaves

Asplenium rhizophyllum plantlet sprouting from the leaf apex of its parent plant Asplenium rhizophyllum walking.JPG
Asplenium rhizophyllum plantlet sprouting from the leaf apex of its parent plant

The stipe (the stalk of the leaf, below the blade) is 0.5 to 12 centimetres (0.20 to 4.7 in) long [2] (occasionally up to 15 centimetres (5.9 in) long), and ranges from one-tenth to one and one-half times the length of the blade. The stipe is reddish-brown and sometimes shiny at the base, becoming green above, [2] and narrowly winged. [5] Scales like those of the rhizome are present at the stipe base, changing to tiny club-shaped hairs above. [2]

The leaf blades are not subdivided, as in most other ferns, but are narrowly triangular [5] to linear or lance-shaped. Their shape can be quite variable, even on the same plant. [2] They measure from 1 to 30 centimetres (0.4 to 10 in) long and from 0.5 to 5 centimetres (0.2 to 2 in) across and have a leathery texture with sparse hairs, more abundant below than above. The rachis (leaf axis) is dull green in color and almost devoid of hairs. On the underside of the blade, the veins are difficult to see and anastomose (split and rejoin each other), forming a series of areoles (the small areas enclosed by the veins) near the rachis. Fertile fronds are usually larger than sterile fronds, but their shape is otherwise the same. The base of the blade is typically heart-shaped (with the stipe protruding from the cleft); [2] [4] the bulges on either side of the cleft are frequently enlarged into auricles (rounded lobes), or occasionally into sharply-pointed, tapering lobes. [2] [5] The leaf tips may be rounded but are typically very long and attenuate (drawn out); the attenuate tips are capable of sprouting roots and growing into a new plant when the tip touches a surface suitable for growth. On rare occasions, the auricles at the leaf base will also take on an attenuate shape and form roots at the tip. [2] The ability of the leaf tips to root and form a new plant at some distance from the parent gives the species its common name. [6] The young leaves forming from a bud at the leaf tip are round to pointed at their apex, not yet having developed the long-attenuate shape. [5]

Specimens of A. rhizophyllum with forked blades have been found in Arkansas and Missouri. The fork usually occurs in the tip, perhaps due to growth after insect damage, but one specimen was found forking from the upper part of the stipe. [7]

Sori and spores

Fertile fronds bear a large number of sori underneath, 1 to 4 millimetres (0.04 to 0.2 in) long, [5] which are not arranged in any particular order. [2] The sori are often fused where veins join, [2] and may curve to follow the vein to which they are attached. [8] The sori are covered by inconspicuous [4] thin, white indusia with untoothed edges. [5] Each sporangium in a sorus carries 64 spores. The diploid sporophyte has a chromosome number of 72. [2]

Similar species

The leaf shape and proliferating tips easily distinguish A. rhizophyllum from most other ferns. Its hybrid descendants share the long-attenuate leaf tip, but are more deeply lobed. An artificial backcross between A. rhizophyllum and A. tutwilerae was closer to A. rhizophyllum in morphology, but still remained some lobes in the basal part of the blade, had a shallowly undulating, rather than smoothly curved, leaf edge in the apical part, showed a maroon color in the stipe up to the base of the leaf blade, and possessed the abortive spores of a sterile hybrid. [9] A. ruprechtii, the Asian walking fern, also possesses attenuate, proliferating tips, but has a lanceolate leaf blade, which tapers to a wedge at the base rather than forming a heart shape. [10] A. scolopendrium , the hart's-tongue fern, has larger, longer leaves that are glossy with a rounded tip. [11]

Taxonomy

This species is commonly known as North American walking fern [6] or simply walking fern, [4] because the growth of new plants at the leaf tip allows it to "walk" across surfaces over several generations. [6] The specific epithet "rhizophyllum", meaning "root leaf", also reflects this characteristic. [4]

Linnaeus first gave the walking fern the binomial Asplenium rhizophyllum in his Species Plantarum of 1753. [12] In 1833, J.H.F. Link placed the species in a segregate genus, Camptosorus, because of the irregular arrangement of its sori (in contrast to the rest of Asplenium, where the sori are confined to the edge of veins). [13] John Smith did not feel that this character was sufficient to segregate it from the rest of Asplenium, but placed it in the genus Antigramma, another Asplenium segregate, on the basis of its reticulate venation, to the convolutions of which he attributed the soral arrangement. [14] It was commonly placed either in Asplenium and Camptosorus by later authors, the latter genus including the similar Asian species A. ruprechtii [2] but phylogenetic studies have shown that Camptosorus is nested within Asplenium and its species should be treated as part of that genus. [15] [16]

The name Asplenium rhizophyllum has also been applied to two other species; in current botanical practice, these are illegitimate later homonyms of Linnaeus' name of 1753. The first of these homonyms was created by Linnaeus himself in 1763, when he accidentally used the name twice, applying it first to his original taxon and again to a species from the West Indies which also proliferates at the leaf tips. [17] He had referred to the West Indian species as A. radicans in 1759, [18] the name by which it is known today. [19] In 1834, Gustav Kunze transferred the species Caenopteris rhizophylla to Asplenium without changing the epithet; [20] George Proctor identified this species, based on a specimen from Dominica, with A. conquisitum, [21] now synonymized with A. rutaceum. [22]

A global phylogeny of Asplenium published in 2020 divided the genus into eleven clades, [23] which were given informal names pending further taxonomic study. A. rhizophyllum belongs to the "A. cordatum subclade" of the "Schaffneria clade". [24] The Schaffneria clade has a worldwide distribution, and members vary widely in form and habitat. [25] There is no clear morphological feature that unites the A. cordatum subclade; the sister species of A. rhizophyllum is A. ruprechtii, which shares an undivided leaf blade and a proliferating tip, while the other three species are scaly spleenworts of dry habitats in Africa and the Middle East. [26]

Hybrids

Walking fern is one of the three parental species of the "Appalachian Asplenium complex", a group of Asplenium hybrids and their progenitors known from eastern North America. Hybridization between walking fern and mountain spleenwort ( A. montanum ) has given rise through chromosome doubling to a new, fertile, species, lobed spleenwort ( A. pinnatifidum ). The sterile hybrid between walking fern and ebony spleenwort ( A. platyneuron ), known as Scott's spleenwort ( A. × ebenoides ), may be found where the two parents are in contact; at one locality, in Havana Glen, Alabama, A. × ebenoides has undergone chromosome doubling to produce a fertile species, Tutwiler's spleenwort ( A. tutwilerae ). [27]

Much more rarely, walking fern hybridizes with two other common spleenworts of eastern North America. The hybrid between walking fern and wall-rue ( A. ruta-muraria ), known as unexpected spleenwort ( A. × inexpectatum ), is known from a single specimen collected on dolomite in Adams County, Ohio. [28] The hybrid between walking fern and maidenhair spleenwort, ( A. trichomanes ssp. trichomanes), Shawnee spleenwort ( A. × shawneense ), is known from one collection on sandstone in the Shawnee Hills of Illinois. [29]

A triploid hybrid between walking fern and Tutwiler's spleenwort was accidentally produced in culture. A similar plant collected from limestone in Shepherdstown, West Virginia could have originated from the same parents, from an unreduced (diploid) gametophyte of Scott's spleenwort crossed with walking fern, or from an unreduced walking fern gametophyte crossed with ebony spleenwort. [9]

Infraspecific taxa

In 1813, Henry Muhlenberg listed lobed spleenwort as Asplenium rhizophyllum var. pinnatifidum, although he did not provide a description distinguishing the variety from the typical species. [30] It was described as the species A. pinnatifidum by Thomas Nuttall in 1818. [31]

A number of forms have been described, of limited taxonomic value. In 1883, J. C. Arthur described walking ferns from limestone cliffs in Muscatine County, Iowa that lacked auricles at the leaf base, with the blade abruptly tapering at the base instead. In this respect, the plants closely resembled A. ruprechtii, but the leaf shape of the Iowa plants was lanceolate (widest near the base) rather than ovate (widest in the middle), and the wide point of the leaf in the Iowa plants appeared slightly lobed. He named these plants Camptosorus rhizophyllus var. intermedius; [32] the variety was subsequently given the status of a form by Willard N. Clute. [33] In 1922, Ralph Hoffmann gave the name C. rhizophyllus f. auriculatus to specimens with proliferating auricles, based on material on limestone from New Marlborough, Massachusetts. [34] In 1924, Frederick W. Gray described as C. rhizophyllus f. angustatus material from a sandstone boulder in Monroe County, West Virginia. These had a short stipe, less than 1 inch (3 cm) long, and narrow leaf blades, less than 0.375 inches (0.95 cm) wide, with the sori almost at the margins. He argued that as they were found along with normal material, they were not solely due to sun exposure. [35] Finally, in 1935, Carl L. Wilson described C. rhizophyllus f. boycei based on material collected from the base of a limestone boulder in Highgate Springs, Vermont by Guy Boyce. These plants had deeply lobed auricles, and erose (jagged or indented) leaf margins with rounded edges. [36]

Distribution

Range map of Asplenium rhizophyllum Asplenium rhizophyllum-range-untitled-green.png
Range map of Asplenium rhizophyllum

The principal range of A. rhizophyllum is in the Appalachian Mountains and the Ozarks. It can be found from southern Quebec and Ontario along the Appalachians and Piedmont southwestward to Mississippi and Alabama, along the Ohio Valley and into the Ozarks west to Nebraska and Oklahoma, and along the Mississippi Valley north to Wisconsin and Minnesota. It has become extinct in Maine and Delaware. [37] It is also presumably extinct in Texas, where one collection from the 19th century has been discovered. [38] The distribution typically follows area of limy soil; [4] sometimes said to be rare, it is better described as locally abundant where conditions favor it. [39]

Ecology and conservation

Walking fern grows on shaded boulders, ledges and in crevices, usually covered with moss. [2] On rare occasions, it is found on fallen tree trunks, [2] as an epiphyte, [40] or on the ground. [4] It is usually found on limestone or other alkaline rocks, rarely on sandstone or other acidic rocks. [4] [2]

While globally secure, it is endangered in some states and provinces at the edge of its range. NatureServe considers it critically imperiled (S1) in Mississippi, New Hampshire and Rhode Island, imperiled (S2) in Michigan and South Carolina, and vulnerable (S3) in North Carolina and Quebec. [1]

Cultivation

It was introduced into cultivation in England in 1680. [41] It prefers low to medium light levels, and a moist, basic potting mix, [6] [42] or soil with added lime chips. [42]

Notes and references

Related Research Articles

<span class="mw-page-title-main">Aspleniaceae</span> Family of ferns

The Aspleniaceae (spleenworts) are a family of ferns, included in the order Polypodiales. The composition and classification of the family have been subject to considerable changes. In particular, there is a narrow circumscription, Aspleniaceae s.s., in which the family contains only two genera, and a very broad one, Aspleniaceae s.l., in which the family includes 10 other families kept separate in the narrow circumscription, with the Aspleniaceae s.s. being reduced to the subfamily Asplenioideae. The family has a worldwide distribution, with many species in both temperate and tropical areas. Elongated unpaired sori are an important characteristic of most members of the family.

<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.

<span class="mw-page-title-main">Walking fern</span> Index of plants with the same common name

Walking fern may refer to two species of fern in the genus Asplenium, which are occasionally placed in a separate genus Camptosorus. The name "walking fern" derives from the fact that new plantlets grow wherever the arching leaves of the parent touch the ground, creating a walking effect. Both have evergreen, undivided, slightly leathery leaves that are triangular and taper to a thin point. On the bottom of the leaves, sori, or spore-bearing structures, cluster along the veins. These hardy plants can be found in shady spots of limestone ledges and limy forest places.

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

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.

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

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.

<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 viride</i> Species of fern in the family Aspleniaceae

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.

<i>Asplenium adiantum-nigrum</i> Species of ferns in the family Aspleniaceae

Asplenium adiantum-nigrum is a common species of fern known by the common name black spleenwort. It is found mostly in Africa, Europe, and Eurasia, but is also native to a few locales in Mexico and the United States.

<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.

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

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.

References

Works cited