Asplenium montanum

Mountain Spleenwort
Conservation status
Secure  (NatureServe)
Scientific classification
Kingdom:	Plantae
Division:	Pteridophyta
Class:	Polypodiopsida
Order:	Polypodiales
(unranked):	Eupolypods II
Family:	Aspleniaceae
Genus:	Asplenium
Species:	A. montanum
Binomial name
Asplenium montanum Willd.
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. First described in 1810, it grows in small crevices in sandstone cliffs with highly acid soil. It can be recognized by its dark blue-green, highly divided leaves. Asplenium montanum is the ancestor, through hybridization, of several other ferns of similar habitat.

Description

Asplenium montanum is a small, evergreen fern which grows in tufts.^[1] The leaves are bluish-green and highly divided, proceeding from a long and often drooping stalk. It is monomorphic, with no difference in form between sterile and fertile fronds.^[1]

The horizontal rhizomes, which are about 1 millimeter across,^[2] 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 (0.08 to 0.2 in) long and from 0.2 to 0.4 millimeters across, with untoothed edges.^[3] They are strongly clathrate (bearing a lattice-like pattern).^[2]

The stipe (the portion of the stem below the leaf 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 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 at the very base of the stipe only;^[3] it is slender and fragile,^[1] and not winged.^[2]

The leaf blade is thick and hairless,^[3] and of a dark blue-green color;^[4] the rachis (leaf axis), like the stipe, is a dull green, with occasional hairs.^[3] 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 in) long and from 1 to 7 centimeters (0.4 to 3 in) wide, occasionally as much as 10 centimeters (4 in).^[3][5] 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,^[3] which cut them into pinnules or deep lobes,^[1] and a rounded to angled base.^[3] The pinnules are indented, but not further cut.^[1] The longest pinnae are those nearest the base of the leaf, which range from 6 to 35 millimeters (0.2 to 1 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.^[3]

Asplenium montanum sori beneath a fertile frond

On fertile fronds, from 1 to 15 elliptical or narrow sori can be found on each pinna.^[1][3] They are 0.5 to 1.5 millimeters long, covered by translucent, pale tan indusia with somewhat jagged edges.^[6] Each sporangium holds 64 spores. The species has a chromosome number of 2n = 72 in the sporophyte; it is a diploid.^[3]

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.^[7] Wall-rue (A. ruta-muraria) has a green stipe,^[1] and its pinnae have longer stalks and are broadest near the tip.^[7] 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.^[8]

Taxonomy

This fern was at first identified by André Michaux, in 1803, as black spleenwort (Asplenium adiantum-nigrum).^[9] Carl Ludwig Willdenow recognized and described it as a separate species, which he named Asplenium montanum, in 1810.^[10] In 1901, John A. Shafer attempted to transfer it to the genus Athyrium as Athyrium montanum,^[11] 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.^[12] The change was not widely accepted and current authorities do not recognize this segregate genus.^[3] 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.^[4]

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),^[13] and in 1936 concluded that Trudell's spleenwort was a hybrid between the first two.^[14] 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).^[15]

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.^[16] 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.^[17]

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.^[18] 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.^[19] 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.^[20]

In addition to the A. pinnatifidum backcross with A. montanum, A. × trudellii, A. bradleyi is also capable of backcrossing with A. montanum. Wherry collected specimens of this hybrid from a cliff near Blairstown, New Jersey in 1935.^[21] The species received no further attention until 1961, when it was described and named in Wherry's honor as Wherry's spleenwort (A. × wherryi).^[8] The sterile diploid A. montanum × platyneuron, precursor to A. bradleyi, was collected in 1972 at Crowder's Mountain, Georgia;^[22] 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

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.^[23] Arkansas populations were discovered in Garland County and Stone County in 2002 and 2008, respectively.^[24] One outlying population in Missouri, collected in 1960, is considered historical;^[23] 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.^[3][24] 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.^[25]

Asplenium montanum grows on acid rocks, in crevices^[3][6] into which moisture seeps from within the rock strata. It usually is under overhangs that protect it from sun and direct rain. It has been found at altitudes from 0 to 2,000 meters (0 to 6,562 ft).^[3] 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.^[26][27]

Ecology and conservation

Asplenium montanum is considered threatened in Connecticut, New York, and Vermont, and endangered in Indiana, Massachusetts, and Rhode Island. The principal threat to New York populations is rock climbing.^[28]

Cultivation

Asplenium montanum may be cultivated outdoors or in a terrarium. In either case, soil should be enriched with chips of acidic rock.^[6]

Citations

1. Cobb, Farnsworth & Lowe 2005, p. 66.
2. Lellinger 1985, p. 242.
3. Wagner, Moran & Werth 1993.
4. Reeves 1974.
5. Lellinger 1985, pp. 242–243.
6. Lellinger 1985, p. 243.
7. Lellinger 1985, p. 233.
8. Smith, Bryant & Tate 1961.
9. Michaux 1803, pp. 265–266.
10. Willdenow 1810, p. 342.
11. Shafer 1901, p. 124.
12. Farwell 1931, p. 273.
13. Wherry 1925.
14. Wherry & Gray 1936.
15. Wagner 1951.
16. Wagner 1953.
17. Wagner 1954.
18. Smith & Levin 1963.
19. Smith & Harborne 1971.
20. Harborne, Williams & Smith 1973.
21. Wherry 1935.
22. Wagner et al. 1973.
23. Kartesz 2014.
24. Peck 2011.
25. Drife & Drife 1990.
26. Wherry 1920.
27. Wherry 1920b.
28. Mountain Spleenwort, NYNHP.

References

• Cobb, Boughton; Farnsworth, Elizabeth; Lowe, Cheryl (2005). A Field Guide to Ferns of Northeastern and Central North America. Peterson Field Guides. New York City: Houghton Mifflin. ISBN 0-618-39406-0.
• Drife, Donald C.; Drife, Joyce E. (1990). "Oliver A. Farwell's early pteridophyte records from the Keweenaw Peninsula". Michigan Botanist 29: 89–96.
• Farwell, Oliver Atkins (1931). "Fern Notes II. Ferns in the Herbarium of Parke, Davis & co". American Midland Naturalist 12: 233–311. doi:10.2307/2420088.
• Harborne, Jeffrey B.; Williams, Christine A.; Smith, Dale M. (1973). "Species-specific kaempferol derivatives in ferns of the Appalachian Asplenium complex". Biochemical Systematics and Ecology 1 (1): 51–54. doi:10.1016/0305-1978(73)90035-5.
• Kartesz, John T. (2014). "Asplenium". Biota of North America Program.
• Lellinger, David B. (1985). A Field Manual of the Ferns & Fern-Allies of the United States & Canada. Washington, DC: Smithsonian Institution Press. ISBN 0874746035.
• Michaux, André (1803). Flora boreali-americana 2. Paris: Charles Crapelet.
• "Mountain Spleenwort". New York Natural Heritage Program. 2011. Retrieved July 5, 2012.
• Peck, James H. (2011). "New and noteworthy additions to the Arkansas fern flora" (PDF). Phytoneuron. 2011-30: 1–33.
• Reeves, Timothy (1974). "A new form of Asplenium montanum from New York". American Fern Journal 64 (4): 105–106. doi:10.2307/1546827.
• Shafer, John A. (1901). "A preliminary list of the vascular flora of Allegheny County, Pennsylvania". Annals of the Carnegie Museum I: 14–127.
• Smith, Dale M.; Bryant, Truman R.; Tate, Donald E. (1961). "Another Asplenium hybrid from Kentucky". American Fern Journal 51 (2): 70–72. doi:10.2307/1546943.
• Smith, Dale M.; Harborne, Jeffrey B. (1971). "Xanthones in the Appalachian Asplenium complex". Phytochemistry 10 (9): 2117–2119. doi:10.1016/S0031-9422(00)97205-4.
• Smith, Dale M.; Levin, Donald A. (1963). "A chromatographic study of reticulate evolution in the Appalachian Asplenium complex". American Journal of Botany 50 (9): 952–958. doi:10.2307/2439783.
• Wagner, Warren H., Jr. (1951). "Recent Fern Literature". American Fern Journal 41 (3): 90–93. doi:10.2307/1545050.
• Wagner, Warren H., Jr. (1953). "A cytological study of the Appalachian spleenworts". American Fern Journal 43 (3): 109–114. doi:10.2307/1545766.
• Wagner, Warren H., Jr. (1954). "Reticulate evolution in the Appalachian Aspleniums". Evolution 8: 103–118. doi:10.2307/2405636.
• Wagner, Warren H., Jr.; Moran, Robbin C.; Werth, Charles R. (1993). "Asplenium montanum". In Flora of North America Editorial Committee. Flora of North America. Flora of North America North of Mexico 2 (New York and Oxford: Oxford University Press). Retrieved July 4, 2012.
• Wagner, Warren H., Jr.; Wagner, Florence S.; Lankalis, Joseph A.; Matthews, James F. (1973). "Asplenium montanum × platyneuron. A new primary member of the Appalachian spleenwort complex from Crowder's Mountain, North Carolina USA.". Journal of the Elisha Mitchell Scientific Society 89: 218–223.
• Wherry, Edgar T. (1920). "The soil reactions of certain rock ferns—II". American Fern Journal 10 (2): 45–52. doi:10.2307/1543831.
• Wherry, Edgar T. (1920b). "Soil acidity—its nature, measurement, and relation to plant distribution". Annual Report of the Board of Regents of the Smithsonian Institution: 247–268.
• Wherry, Edgar T. (1925). "The Appalachian Aspleniums". American Fern Journal 15 (2): 48–54. doi:10.2307/1544001. JSTOR 1544001.
• Wherry, Edgar T. (1935). "Fern field notes, 1935". American Fern Journal 25 (4): 123–126. doi:10.2307/1544060.
• Wherry, Edgar T.; Gray, William D. (1936). "Variety of some Appalachian Aspleniums". American Fern Journal 26 (3): 77–86. doi:10.2307/1543680.
• Willdenow, Carl Ludwig (1810). Linnaei species plantarum. 5 pt. 1. Berlin: G. C. Nauk.

External links

External identifiers for Asplenium montanum
Encyclopedia of Life	597648
GBIF	2650651
ITIS	17351
NCBI	243998
Also found in: Wikispecies

• NatureServe Explorer
• United States Department of Agriculture Plants Profile - Asplenium montanum