# Asplenium resiliens

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***Asplenium resiliens***, the **blackstem spleenwort** or **little ebony spleenwort**, is a species of [fern](/source/Fern) native to the Western Hemisphere, ranging from the southern United States south to Uruguay, including parts of the Caribbean. Found on [limestone](/source/Limestone) substrates, it is named for its distinctive purplish-black [stipe](/source/Stipe_(botany)) and [rachis](/source/Rachis). A [triploid](/source/Triploid), it is incapable of sexual reproduction and produces spores [apogamously](/source/Apomixis). First described by [Martens](/source/Martin_Martens) and [Galeotti](/source/Henri_Guillaume_Galeotti) in 1842 under the previously used name *Asplenium parvulum*, the species was given its current, valid name by [Kunze](/source/Gustav_Kunze) in 1844. Several similar species are known from the tropics; *A. resiliens* may have arisen from these species by [reticulate evolution](/source/Reticulate_evolution), but precise relationships among the group are not yet certain.

## Description

It is a small fern with [pinnate](/source/Pinnate) [fronds](/source/Frond), growing in erect tufts, with a shiny black [stipe](/source/Stipe_(botany)) and [rachis](/source/Rachis) (stem and leaf axis). Sterile and fertile fronds are similar in appearance.[1]

The roots are thin and wiry and do not proliferate to form new plants.[2][3] The [rhizome](/source/Rhizome) is short and erect,[4][5] about 2 millimeters (0.08 in) in diameter.[6] It has variously been described as sometimes branching[6] or unbranched.[2] It bears stiff[1] filamentous, linear, or lance-shaped scales, which are blackish in color[2][4] and obscurely clathrate (bearing a lattice-like pattern)[4] 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,[2] with untoothed, often brown, margins[2][4] and long, drawn-out tips.[5]

Leaves are erect and borne in dense clumps,[5] varying in size from 3 to 45 centimeters (1.2 to 18 in) long[3] and from 0.5 to 2.5 centimeters (0.20 to 0.98 in) wide.[4][5] The stipe (the stalk of the leaf, below the blade) is straight and stiff and a glossy black[2] to purplish-black in color.[3][4][5][6] 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).[2][5] The stipe measures 1.5 to 3 centimeters (0.6 to 1 in)[2][3] long (rarely as long as 5 centimeters (2 in)),[4] and comprises one-tenth to one-quarter[2] or one-third of the length of the blade.[3] It is round in cross-section but slightly flattened [adaxially](/source/Adaxial) and has indistinct [wings](/source/Glossary_of_botanical_terms#W) 0.1 to 0.3 millimeters (0.0039 to 0.012 in) on either side,[3][6] or lacks them entirely.[7]

The leaf blade is linear in shape, sometimes slightly wider below the tip or just above the base.[2][3] It measures from 4 to 35 centimeters (1.6 to 14 in) long[4][7] 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.[2][4] Unlike the related Palmer's spleenwort (*[A. palmeri](/source/Asplenium_palmeri)*), it does not form proliferating buds at the tip;[4] however, the pinnatifid is often deciduous, leaving behind a naked rachis.[7] The blade is hairless[2] or bears scattered club-shaped hairs, 0.1 millimeters (0.004 in) long, beneath.[3] The leaf tissue is often bluish-green[8] and thick in texture,[2] not quite leathery.[4] 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.[2][3] 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,[4][7] or a whitish to tan wing similar in dimensions to that of the stipe.[3]

The blade is cut into pinnae throughout its length, from 20 to 40 pairs per leaf.[2] The pinnae are [sessile](/source/Sessile_(botany)) (stalkless) or have minute stalks[7] and are rectangular in shape,[2] tapering slightly toward the tip.[6] 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.[2][3] 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.[7] Each pinna usually has an [auricle](/source/Auricle_(botany)) 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](/source/Crenate) (or deep, rounded teeth in exceptional shade-grown specimens), and are often rolled under.[2][3][9] The tips of the pinnae are blunt.[2][3] The lower pinnae are widely spaced on the rachis, and reflexed downwards.[1][3] Leaf veins are free (they do not rejoin one another) and are difficult to see;[2] fertile veins are once-forking and do not terminate in [hydathodes](/source/Hydathode) (prominent swellings).[7] Fertile pinnae bear 2 to 6 pairs of [sori](/source/Sorus) (rarely as few as 1 pair or as many as 10),[2][3] about 1 millimeter (0.04 in) in length,[6] on both sides of the midrib; the sori are crowded at the edges and often merge as they age.[2][4] The [indusia](/source/Indusium) 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,[3] greenish or pale yellowish to whitish in color and opaque, with straight or slightly jagged edges.[6][7] They are persistent after the spores mature, but may be hidden by the full [sporangia](/source/Sporangium).[4] *A. resiliens* has a [chromosome number](/source/Chromosome_number) of *n* = 2*n* = 108 and produces 32 unreduced,[a] round or egg-shaped spores per sporangium.[2][3]

### Similar species

*A. resiliens* resembles several of its congeners; in particular, it belongs to a group comprising varicolored spleenwort (*[A. heterochroum](/source/Asplenium_heterochroum)*), *[A. nesioticum](/source/Asplenium_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,[10] the latter characteristic distinguishing them from single-sorus spleenwort (*[A. monanthes](/source/Asplenium_monanthes)*) and others.[11] 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.[12][13] 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.[14]

In the more temperate parts of its range, *A. resiliens* may be confused with ebony spleenwort (*[A. platyneuron](/source/Asplenium_platyneuron)*) and maidenhair spleenwort (*[A. trichomanes](/source/Asplenium_trichomanes)*). Its stipe and rachis are darker[1] and its pinnae smaller[8] and more rounded than that of *A. platyneuron*, which also displays [frond dimorphism](/source/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,[1] the texture of the leaf tissue is more leathery,[12] and the stipe darker.[2]

A hybrid between *A. resiliens* and *A. heterochroum*, Morzenti's spleenwort (*[A. heteroresiliens](/source/Asplenium_heteroresiliens)*), is found in Florida and the Carolinas. As *A. resiliens* is [triploid](/source/Triploid) and *A. heterochroum* is [tetraploid](/source/Tetraploid), the hybrid is [pentaploid](/source/Pentaploid) and reproduces [apogamously](/source/Apomixis).[2] 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.[2][15] 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.[16][b]

## Taxonomy

The species was described from Mexico in 1842 by [Martin Martens](/source/Martin_Martens) and [Henri Guillaume Galeotti](/source/Henri_Guillaume_Galeotti).[17] 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](/source/Gustav_Kunze) in 1844.[18] He did not discuss the reason for his choice of epithet, although it may refer to the springy character of the stems.[19] 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](/source/Andr%C3%A9_Michaux); in fact, Michaux's material was ebony spleenwort, and in any case the name *A. trichomanoides* was preoccupied and invalid.[20][21] [W. J. Hooker](/source/William_Jackson_Hooker) considered it a variety of ebony spleenwort, giving it the name *A. ebeneum* var. *minus* in 1860,[22] but others continued to maintain it as a separate species; [Eaton](/source/Daniel_Cady_Eaton) refers to it as "little ebony spleenwort".[21] [Oliver Atkins Farwell](/source/Oliver_Atkins_Farwell) transferred it to a segregate genus as *Chamaefilix resiliens* in 1931,[23] but this name was never widely accepted.

In 1940, [A.H.G. Alston](/source/Arthur_Hugh_Garfit_Alston) described material from [Argentina](/source/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](/source/Hyaline) cells on the scale margins.[24] 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,[25] 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".[26] 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.[27] Within this group, a study based on [nuclear](/source/Cell_nucleus) and [plastid](/source/Plastid) [genetic markers](/source/Genetic_marker), using material from the United States, [Mexico](/source/Mexico), and [Costa Rica](/source/Costa_Rica), showed that *A. resiliens* forms a [clade](/source/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.[28]

## Distribution and habitat

*Asplenium resiliens* is found in the southern United States, Mexico, [Hispaniola](/source/Hispaniola), [Jamaica](/source/Jamaica), [Guatemala](/source/Guatemala), and South America[2] from [Colombia](/source/Colombia) and [Venezuela](/source/Venezuela) south to northern Argentina, [Uruguay](/source/Uruguay), and [Brazil](/source/Brazil).[4][3] Within the United States, it is found from [Florida](/source/Florida) west to [Arizona](/source/Arizona) and southern [Nevada](/source/Nevada) and north to south-central [Pennsylvania](/source/Pennsylvania).[6][29][c] It is found throughout most of Mexico, but is most abundant in the northeast, in [Nuevo León](/source/Nuevo_Le%C3%B3n) and [Coahuila](/source/Coahuila).[3] In Guatemala, it is known from the western departments of [Huehuetenango](/source/Huehuetenango_Department), [El Quiché](/source/Quich%C3%A9_Department), and [Sololá](/source/Solol%C3%A1_Department).[5] In Argentina, its distribution follows the arc of the mountains from [Jujuy Province](/source/Jujuy_Province) southward to the [Sierra de la Ventana](/source/Sierra_de_la_Ventana_(mountains)) in [Buenos Aires Province](/source/Buenos_Aires_Province); it is also known from [Mendoza Province](/source/Mendoza_Province), in the west of the country, from the departments of [Las Heras](/source/Las_Heras_Department) and [Luján de Cuyo](/source/Luj%C3%A1n_de_Cuyo_Department).[30]

It is found on or at the base of cliffs or [sinkholes](/source/Sinkhole), on [limestone](/source/Limestone) or other [alkaline](/source/Alkaline) rocks,[2][3][6] although specimens have also been reported from crevices in [granite](/source/Granite)[31] and [sandstone](/source/Sandstone).[32][33] Growth on soil is rare.[4] It may be found in forests or on [boulders](/source/Boulder), ledges, and in crevices of cliffs.[5][4] Plants grow at altitudes ranging from 100 meters (300 ft) (in North America) to 3,900 meters (13,000 ft) (in Guatemala).[2][5] 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](/source/Peru) material) are believed to be due to environmental factors rather than any taxonomic distinction.[7]

## Ecology and conservation

*Asplenium resiliens* is a [triploid](/source/Triploid) and reproduces apogamously,[6] producing 32 spores per sporangium.[3] Specimens with 64 well-formed spores per sporangium, believed to be sexually reproducing, were collected from Green Gulch in the [Chisos Mountains](/source/Chisos_Mountains) in 1937, although other specimens since collected in the area have the typical 32 spores per sporangium.[34]

While globally secure (G5), it is considered an [endangered species](/source/Endangered_species) in many of the [states](/source/U.S._state) at the northern edge of its North American range. [NatureServe](/source/NatureServe) considers it to be critically imperiled (S1) in Colorado, Illinois, Maryland, Mississippi, Nevada, Ohio, Pennsylvania, South Carolina, and Utah, imperiled (S2) in Indiana, Kansas and North Carolina, and vulnerable (S3) in West Virginia.[35] It has become extinct in [Louisiana](/source/Louisiana) since the limestone caprock of a [salt dome](/source/Salt_dome) at [Winnfield](/source/Winnfield,_Louisiana), the only location for the fern in the state, was quarried away.[36] It is also believed to be extinct in [Ohio](/source/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.[35]

## Cultivation

*Asplenium resiliens* is tolerant of cold to [USDA](/source/United_States_Department_of_Agriculture) [hardiness zone](/source/Hardiness_zone) 6. It prefers moist conditions, a [basic](/source/Alkaline) potting soil, and a medium amount of light.[8]

## Notes and references

### Notes

1. Normally, ferns have a [diploid](/source/Diploid) [sporophyte](/source/Sporophyte) which produces [haploid](/source/Haploid) spores through [meiosis](/source/Meiosis); these grow into [gametophytes](/source/Gametophyte), whose haploid [gametes](/source/Gamete) 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.

1. 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.

1. Reports that the range includes [Delaware](/source/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.

### References

1. Cobb, Farnsworth & Lowe 2005, p. 72.

1. Wagner, Moran & Werth 1993.

1. Mickel & Smith 2004, p. 117.

1. Tryon & Stolze 1993, p. 15.

1. Stolze 1981, p. 87.

1. Lellinger 1985, p. 238.

1. Stolze 1981, p. 88.

1. Hoshizaki & Moran 2001, p. 203.

1. Maxon 1913, p. 144.

1. Mickel & Smith 2004, p. 100.

1. Mickel & Smith 2004, p. 106.

1. Mickel & Smith 2004, p. 75.

1. Lellinger 1985, pp. 231–232.

1. Stolze 1981, p. 74.

1. Lellinger 1985, p. 232.

1. Wagner 1966.

1. Martens & Galeotti 1842, p. 60.

1. Kunze 1844, pp. 331–332.

1. Snyder & Bruce 1986, p. 78.

1. Kunze 1848, pp. 85–86.

1. Eaton 1879, p. 279.

1. Hooker 1860, p. 139.

1. Farwell 1931, p. 269.

1. Alston 1940.

1. Xu et al. 2020, p. 27.

1. Xu et al. 2020, p. 30.

1. Xu et al. 2020, p. 44.

1. Dyer, Savolainen & Schneider 2012.

1. Kartesz 2014.

1. Arana et al. 2011.

1. Correll 1949, p. 256.

1. Lathrop 1958, p. 142.

1. Welsh et al. 1987, p. 19.

1. Wendt 1979.

1. NatureServe 2024.

1. Moore 2013.

1. Cusick & Schneider 1993.

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- Arana, Marcelo; Bianco, César; Carretero, Eduardo Martínez; Oggero, Antonia (2011). ["Flora de Mendoza: Licofitas y Helechos de Mendoza"](http://www.scielo.org.ar/scielo.php?pid=S1852-73292011000200005&script=sci_arttext) (in Spanish). *Multequina*. **20** (2)
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- [*A Utah Flora*](https://www.biodiversitylibrary.org/page/7934110). Vol. 9. Great Basin Naturalist Memoirs. Brigham Young University. 1987. ISBN 0-8425-2260-3.
- Wendt, Tom (1979). ["Sexuality in Asplenium resiliens"](https://www.biodiversitylibrary.org/page/31876969). *[American Fern Journal](/source/American_Fern_Journal)*. **69** (3): 96. [doi:10.2307/1546391](https://doi.org/10.2307/1546391). [JSTOR 1546391](https://www.jstor.org/stable/1546391)
- Xu, Ke-Wang; Zhang, Liang; Rothfels, Carl J.; Smith, Alan R.; Viane, Ronald; Lorence, David; Wood, Kenneth R.; Cheng, Cheng-Wei; Knapp, Ralf; Zhou, Lin; Lu, Ngan Thi; Zhou, Xin-Mao; Wei, Hong-Jin; Fan, Qiang; Chen, Su-Fang; Cicuzza, Daniele; Gao, Xin-Fen; Li, Wen-Bo; Zhang, Li-Bing (2020). "A global plastid phylogeny of the fern genus Asplenium (Aspleniaceae)". *[Cladistics](/source/Cladistics_(journal))*. **36** (1): 22–71. [doi:10.1111/cla.12384](https://doi.org/10.1111/cla.12384). [PMID 34618950](https://pubmed.ncbi.nlm.nih.gov/34618950). [S2CID 201197385](https://api.semanticscholar.org/CorpusID:201197385)

## External links

- [Illinois conservation assessment](http://hdl.handle.net/2142/10069)
- [Comparison to other Arizona ferns](http://www.mineralarts.com/ferns/DesertFernsGuide.html)
- [Good illustrations of the plant](http://wnmu.edu/academic/nspages/gilaflora/asplenium_resiliens.html)

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Adapted from the Wikipedia article [Asplenium resiliens](https://en.wikipedia.org/wiki/Asplenium_resiliens) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Asplenium_resiliens?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.