Asplenium resiliens

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Asplenium resiliens
Asplenium resiliens habit.jpg
Leaves of A. resiliens, showing bluish-green color and opposite pinnae
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. resiliens
Binomial name
Asplenium resiliens
Synonyms

Asplenium ebeneum Ait. var. minus Hook.
Asplenium lealii Alston
Chamaefilix resiliens(Kunze) Farw.

Contents

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.

Description

It is a small fern with pinnate fronds, growing in erect tufts, with a shiny black stipe and rachis (stem and leaf axis). Sterile and fertile fronds are similar in appearance. [2]

The roots are thin and wiry and do not proliferate to form new plants. [3] [4] The rhizome is short and erect, [5] [6] about 2 millimeters (0.08 in) in diameter. [7] It has variously been described as sometimes branching [7] or unbranched. [3] It bears stiff [2] filamentous, linear, or lance-shaped scales, which are blackish in color [3] [5] and obscurely clathrate (bearing a lattice-like pattern) [5] or entirely black. The scales are 3 to 5 millimeters (0.1 to 0.2 in) long and 0.2 to 0.6 millimeters (0.008 to 0.02 in) wide, [3] with untoothed, often brown, margins [3] [5] and long, drawn-out tips. [6]

Leaves are erect and borne in dense clumps, [6] varying in size from 3 to 45 centimeters (1.2 to 18 in) long [4] and from 0.5 to 2.5 centimeters (0.20 to 0.98 in) wide. [5] [6] The stipe (the stalk of the leaf, below the blade) is straight and stiff and a glossy black [3] to purplish-black in color. [4] [5] [6] [7] It may be smooth or bear scattered blackish-brown, threadlike scales 1 to 1.5 millimeters (0.039 to 0.059 in) long and tan, club-shaped hairs 0.1 to 0.2 millimeters (0.004 to 0.008 in) long which are appressed (lie flat against the stipe). [3] [6] The stipe measures 1.5 to 3 centimeters (0.6 to 1 in) [3] [4] long (rarely as long as 5 centimeters (2 in)), [5] and comprises one-tenth to one-quarter [3] or one-third of the length of the blade. [4] It is round in cross-section but slightly flattened adaxially and has indistinct wings 0.1 to 0.3 millimeters (0.0039 to 0.012 in) on either side, [4] [7] or lacks them entirely. [8]

The leaf blade is linear in shape, sometimes slightly wider below the tip or just above the base. [3] [4] It measures from 4 to 35 centimeters (1.6 to 14 in) long [5] [8] and from 1 to 2 centimeters (0.4 to 0.8 in) wide, sometimes as wide as 2.5 centimeters (0.98 in). It abruptly converges to a lobed, pointed tip and gradually tapers at its base. [3] [5] Unlike the related Palmer's spleenwort ( A. palmeri ), it does not form proliferating buds at the tip; [5] however, the pinnatifid is often deciduous, leaving behind a naked rachis. [8] The blade is hairless [3] or bears scattered club-shaped hairs, 0.1 millimeters (0.004 in) long, beneath. [4] The leaf tissue is often bluish-green [9] and thick in texture, [3] not quite leathery. [5] The rachis, like the stipe, is rounded, blackish and shiny; it may be smooth or have a few of the tan hairs found on the stipe. [3] [4] The winging of the stipe extends up the rachis; it is variously described as taking the form of parallel, cartilaginous ribs, with a narrow, green, leafy wing, the ribs fusing into a wing towards the tip of the leaf, [5] [8] or a whitish to tan wing similar in dimensions to that of the stipe. [4]

Sori on the underside of an Asplenium resiliens leaf Asplenium resiliens sori.jpg
Sori on the underside of an Asplenium resiliens leaf

The blade is cut into pinnae throughout its length, from 20 to 40 pairs per leaf. [3] The pinnae are sessile (stalkless) or have minute stalks [8] and are rectangular in shape, [3] tapering slightly toward the tip. [7] In North American and Mexican material, those in the middle of the leaf blade measure from 10 to 20 millimeters (0.4 to 0.8 in) in length (rarely as small as 4 millimeters (0.2 in)) and from 2 to 5 millimeters (0.08 to 0.2 in) in width. [3] [4] In Guatemalan material, the pinnae typically measured from 1.5 to 8 millimeters (0.059 to 0.31 in) in length and from 1 to 3 millimeters (0.04 to 0.1 in) in width, the ratio of length to breadth being typically 1.5 to 2.5. [8] Each pinna usually has an auricle at its base, pointing towards the tip of the blade; occasionally auricles pointing towards the base of the blade are also present. The edges of the pinnae are untoothed or have shallowly rounded teeth (or deep, rounded teeth in exceptional shade-grown specimens), and are often rolled under. [3] [4] [10] The tips of the pinnae are blunt. [3] [4] The lower pinnae are widely spaced on the rachis, and reflexed downwards. [2] [4] Leaf veins are free (they do not rejoin one another) and are difficult to see; [3] fertile veins are once-forking and do not terminate in hydathodes (prominent swellings). [8] Fertile pinnae bear 2 to 6 pairs of sori (rarely as few as 1 pair or as many as 10), [3] [4] about 1 millimeter (0.04 in) in length, [7] on both sides of the midrib; the sori are crowded at the edges and often merge as they age. [3] [5] The indusia covering them are from 0.8 to 1.5 millimeters (0.031 to 0.059 in) long (rarely to 2 millimeters (0.079 in)) and from 0.3 to 0.5 millimeters (0.01 to 0.02 in) wide, [4] greenish or pale yellowish to whitish in color and opaque, with straight or slightly jagged edges. [7] [8] They are persistent after the spores mature, but may be hidden by the full sporangia. [5] A. resiliens has a chromosome number of n = 2n = 108 and produces 32 unreduced, [lower-alpha 1] round or egg-shaped spores per sporangium. [3] [4]

Similar species

A. resiliens resembles several of its congeners; in particular, it belongs to a group comprising varicolored spleenwort ( A. heterochroum ), A. nesioticum , and A. palmeri, as well as A. resiliens. The members of this group all share dark, lustrous stipes and lack prominent hydathodes on the surface of the blade, [11] the latter characteristic distinguishing them from single-sorus spleenwort ( A. monanthes ) and others. [12] Among the group, A. resiliens has a slightly more leathery leaf texture than the rest, lacks distinct teeth on the pinna margins, and tends to have once-forked, rather than simple, fertile veins. [13] [14] Stolze noted that specimens with leaf texture so thick as to obscure the veins may be identified as A. resiliens, as A. heterochroum and A. palmeri very rarely become this leathery. [15]

In the more temperate parts of its range, A. resiliens may be confused with ebony spleenwort ( A. platyneuron ) and maidenhair spleenwort ( A. trichomanes ). Its stipe and rachis are darker [2] and its pinnae smaller [9] and more rounded than that of A. platyneuron, which also displays frond dimorphism with prostrate sterile fronds. The pinnae of A. resiliens are more widely spaced than those of A. trichomanes, which also lack the upward-pointing auricle, [2] the texture of the leaf tissue is more leathery, [13] and the stipe darker. [3]

A hybrid between A. resiliens and A. heterochroum, Morzenti's spleenwort ( A. heteroresiliens ), is found in Florida and the Carolinas. As A. resiliens is triploid and A. heterochroum is tetraploid, the hybrid is pentaploid and reproduces apogamously. [3] The hybrid is difficult to distinguish from the parental species, being intermediate in morphology; its fertile veins are sometimes less forked than in A. resiliens, its leaves tend to have more toothed edges, and it bears misshapen sterile spores together with large, globose, unreduced spores. [3] [16] In A. resiliens, the sori are close to the pinna margin; in A. heteroresiliens, they are slightly closer to the margin than to the costa. [17] [lower-alpha 2]

Taxonomy

The species was described from Mexico in 1842 by Martin Martens and Henri Guillaume Galeotti. [18] However, the name they chose, Asplenium parvulum, had already been used, rendering their name a later illegitimate homonym. It was first described under its valid name, A. resiliens, by Gustav Kunze in 1844. [19] He did not discuss the reason for his choice of epithet, although it may refer to the springy character of the stems. [20] In 1848, Kunze assigned North American material of A. resiliens the name A. trichomanoides, believing it to be the plant described under that name by André Michaux; in fact, Michaux's material was ebony spleenwort, and in any case the name A. trichomanoides was preoccupied and invalid. [21] [22] W. J. Hooker considered it a variety of ebony spleenwort, giving it the name A. ebeneum var. minus in 1860, [23] but others continued to maintain it as a separate species; Eaton refers to it as "little ebony spleenwort". [22] Oliver Atkins Farwell transferred it to a segregate genus as Chamaefilix resiliens in 1931, [24] but this name was never widely accepted.

In 1940, A.H.G. Alston described material from Argentina, previously identified with A. trichomanes, as a new species, A. lealii, which he named for its collector, Adrián Ruiz Leal. He noted that this species resembled A. resiliens, but had fewer hyaline cells on the scale margins. [25] This species is now treated as a synonym of A. resiliens.

A global phylogeny of Asplenium published in 2020 divided the genus into eleven clades, [26] which were given informal names pending further taxonomic study. A. resiliens, A. heterochroum, and A. palmeri belong to the "A. monanthes subclade" of the "A. trichomanes clade". [27] The A. trichomanes clade has a worldwide distribution. Members of the clade grow on rocks and have once-pinnate leaf blades with slender, chestnut- to dark-brown stalks. The A. monanthes subclade principally occurs in the Neotropics. [28] Within this group, a study based on nuclear and plastid genetic markers, using material from the United States, Mexico, and Costa Rica, showed that A. resiliens forms a clade with A. palmeri and specimens resembling A. heterochroum. Within this clade, the latter two formed a sister clade to A. resiliens. Plastid markers indicated that the A. resiliens clade was divided into two separate groups, while nuclear markers from two distinct groups were distributed among the samples, with five specimens carrying markers from both groups, and one specimen each carrying markers from only one of the groups. This suggests that A. resiliens developed by reticulate evolution, i.e., as the descendant of a hybrid between two taxa, each parental taxon carrying one of the two groups of plastid and nuclear makers. In addition, of two specimens, one morphologically identified as A. resiliens, and one as A. palmeri, both were found to contain a nuclear marker from the A. resiliens clade and another from the A. palmeri clade. While apogamous specimens of A. palmeri are known, it is typically a sexually reproducing species, and the implication of these specimens is unclear. [29]

Distribution and habitat

Asplenium resiliens is found in the southern United States, Mexico, Hispaniola, Jamaica, Guatemala, and South America [3] from Colombia and Venezuela south to northern Argentina, Uruguay, and Brazil. [5] [4] Within the United States, it is found from Florida west to Arizona and southern Nevada and north to south-central Pennsylvania. [7] [30] [lower-alpha 3] It is found throughout most of Mexico, but is most abundant in the northeast, in Nuevo León and Coahuila. [4] In Guatemala, it is known from the western departments of Huehuetenango, El Quiché, and Sololá. [6] In Argentina, its distribution follows the arc of the mountains from Jujuy Province southward to the Sierra de la Ventana in Buenos Aires Province; it is also known from Mendoza Province, in the west of the country, from the departments of Las Heras and Luján de Cuyo. [31]

It is found on or at the base of cliffs or sinkholes, on limestone or other alkaline rocks, [3] [4] [7] although specimens have also been reported from crevices in granite [32] and sandstone. [33] [34] Growth on soil is rare. [5] It may be found in forests or on boulders, ledges, and in crevices of cliffs. [6] [5] Plants grow at altitudes ranging from 100 meters (300 ft) (in North America) to 3,900 meters (13,000 ft) (in Guatemala). [3] [6] Plants at the higher altitudes (3,000 meters (9,800 ft) and above) are stunted and more compact than usual, with leaves 4 to 10 centimeters (2 to 4 in) long and overlapping pinnae a few millimeters in length. These differences (observed in Guatemalan and Peruvian material) are believed to be due to environmental factors rather than any taxonomic distinction. [8]

Ecology and conservation

Asplenium resiliens is a triploid and reproduces apogamously, [7] producing 32 spores per sporangium. [4] Specimens with 64 well-formed spores per sporangium, believed to be sexually reproducing, were collected from Green Gulch in the Chisos Mountains in 1937, although other specimens since collected in the area have the typical 32 spores per sporangium. [35]

While globally secure, it is considered an endangered species in many of the states at the northern edge of its North American range. NatureServe considers it to be critically imperiled (S1) in Colorado, Illinois, Indiana, Maryland, Mississippi, Nevada, Pennsylvania, South Carolina, and Utah, imperiled (S2) in Kansas and North Carolina, and vulnerable (S3) in West Virginia. [1] It has become extinct in Louisiana since the limestone caprock of a salt dome at Winnfield, the only location for the fern in the state, was quarried away. [36] It is also believed to be extinct in Ohio, where it was last collected in 1900, although suitable habitat still exists in the state. [37]

Quarrying of the calcareous rocks on which it grows poses a "low-level" threat to the species. [1]

Cultivation

Asplenium resiliens is tolerant of cold to USDA hardiness zone 6. It prefers moist conditions, a basic potting soil, and a medium amount of light. [9]

Notes and references

Notes

  1. Normally, ferns have a diploid sporophyte which produces haploid spores through meiosis; these grow into gametophytes, whose haploid gametes fuse to form a diploid zygote which develops into a sporophyte. Since A. resiliens is triploid, it cannot form tetrads and undergo meiosis; the spores it forms are unreduced in chromosome number, and the sporophyte sprouts directly from the gametophyte without fertilization.
  2. Wagner's 1966 paper laid great stress on the extent to which the sorus-bearing veins of the basal auricle fork in distinguishing A. heteroresiliens from its parents, but this emphasis is not taken up in later keys.
  3. Reports that the range includes Delaware are an error derived from a misreading of Clyde F. Reed's Ferns and fern-allies of Maryland and Delaware. The state lacks suitable habitat for the plant.

Related Research Articles

<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>Myriopteris clevelandii</i> Species of fern

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.

<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 <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 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.

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 alabamensis</i> Species of fern

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 yatskievychiana is a small fern native to Mexico, a member of the family Pteridaceae. It is known only from a single collection in the Mexican state of Sonora. 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.

<i>Argyrochosma peninsularis</i> Species of fern in the family Pteridaceae

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

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<i>Argyrochosma microphylla</i> Species of fern in the family Pteridaceae

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".

<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