Lobed spleenwort | |
---|---|
Lobed spleenwort growing in a crevice in schist | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Division: | Polypodiophyta |
Class: | Polypodiopsida |
Order: | Polypodiales |
Suborder: | Aspleniineae |
Family: | Aspleniaceae |
Genus: | Asplenium |
Species: | A. pinnatifidum |
Binomial name | |
Asplenium pinnatifidum | |
Synonyms | |
Asplenium rhizophyllum var. pinnatifidum Muhl. nom. nud. Contents |
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 ( Asplenium rhizophyllum ), 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 ( Asplenium montanum ), 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.
Asplenium pinnatifidum is a small fern with bright green, wrinkled, pinnatifid (lobed) fronds. [2] [3] These form evergreen, perennial tufts. [3] Notable characteristics are the shiny stem, dark only at the base, and the long-tapering, variably lobed leaf blades. [2] The fronds are monomorphic, the sterile and fertile fronds appearing the same size and shape. [3]
The roots of A. pinnatifidum are not proliferous, so it appears as clusters of leaves springing from a single rhizome. The leaves are closely spaced on the rhizome, which is frequently branched. [2] The rhizome is about 1 millimeter (0.04 in) in diameter, covered with narrowly triangular scales which are dark reddish-brown or blackish in color, and strongly clathrate (bearing a lattice-like pattern). [2] [4] The scales are 3 to 5 millimeters (0.1 to 0.2 in) long and 0.3 to 0.5 millimeters wide, with untoothed edges. The stipe (the stalk of the leaf, below the blade) is shiny and dark reddish brown at the base. The color fades to green in the upper one-third to one-half of the stipe. It is covered in narrowly triangular, dark reddish-brown scales at the base, which diminish into hairs in the upper part of the stipe. [2] It may show narrow wings from the base of the leaf to near the base of the stipe. [4] The stipe is 1 to 10 centimeters (0.4 to 4 in) long, [2] and may be from one-tenth to one and one-half times the length of the blade. [3]
The overall shape of the blade is narrowly triangular or lance-shaped, sometimes with an irregular outline. The blade tapers to a long point, the length of the taper being variable among specimens. [2] [4] The blades are generally curled with downward-pointing tips. [3] The tip of the blade sometimes develops a swelling which may differentiate into a proliferous bud and, very rarely, into a plantlet, as in walking ferns. [5] Adventitious sporangia may appear around the buds when they form, even, unusually, on the upper surface of the leaf. [6] The base of the blade may be squared off or notched to a varying extent along the rachis (central axis of the leaf). The blade ranges from 2 to 17 centimeters (0.8 to 7 in) long, rarely to 20 centimeters (7.9 in), and 1 to 4 centimeters (0.4 to 2 in) in width, rarely to 13 centimeters (5.1 in), and is thick [2] and somewhat leathery. [3] Blades are either entirely pinnatifid (lobed but not completely cut), or cut to form a single pair of pinnae at the base. When they exist, the pinnae are roughly oval or triangle-shaped, sometimes narrow, and are from 5 to 20 millimeters (0.2 to 0.8 in) long, rarely to 90 millimeters (3.5 in), and 0.4 to 1 millimeters in width (rarely to 1.2 millimeters). The base of the pinnae may be squared off or taper to a point, while the edges are wrinkled to toothed. The tip can vary from rounded to pointed. [2] The lobes of the blade gradually diminish towards the tip, sometimes becoming simply wavy. [3] The rachis is green, sometimes turning tan when dry. The underside of the rachis and blade have a few scattered, small hairs. [2] Overall, the blades are quite morphologically variable; [3] in younger blades, the edges may be not at all lobed or may be wavy. [4] The veins are free and forking, only rarely anastomosing (rejoining one another to form nets). [2]
Each segment (pinna or lobe) of a fertile frond has one to six sori, sometimes more than forty in extreme cases. The sori usually fuse with one another as they age. [2] These are 1 to 2 millimeters (0.04 to 0.08 in) long They are covered with thin, whitish indusia with untoothed edges, [4] which are persistent. [3] Each sporangium holds 64 spores. The species has a chromosome number of 144 in the sporophyte, indicating an allotetraploid origin. [2]
While no named varieties or forms of A. pinnatifidum have been described, an unusual population was described from Giant City State Park in southern Illinois in 1956. In it, the leaf blade was highly reduced, barely exceeding the rachis, except for a series of stubby projections under which the sori were borne. [7] Individual plants have also been known on occasion to develop forked leaves, which appears to be a developmental accident rather than a stable genetically-controlled trait. [8]
A. pinnatifidum is somewhat similar to its parent species A. rhizophyllum. In comparison, however, A. pinnatifidum is distinctly lobed when mature, tends to have longer stipes in proportion to its leaf size, and has a more upright habit. [3] It might be confused with Countess Dalhousie's spleenwort (A. dalhousiae), of Asia and the American Southeast, but the latter has short, dull stipes with larger, toothed scales. [2] A. pinnatifidum closely resembles the hybrid Scott's spleenwort (A. × ebenoides) (including the fertile Tutwiler's spleenwort, A. tutwilerae), but those species have a wholly dark stipe, with the dark color extending into the rachis, [2] [3] and longer lobes on the blade. [4]
Among the hybrid species of which it is a parent, A. pinnatifidum is most similar to Graves' spleenwort (A. × gravesii), a hybrid with Bradley's spleenwort (A. bradleyi), and to a lesser extent, to Trudell's spleenwort (A. × trudellii) and Kentucky spleenwort (A. × kentuckiense). In A. × gravesii, the dark color of the stipe extends to the base of the leaf blade, the blades often have more than one pair of pinnae, and their edges are shallowly wrinkled or toothed. In addition, the basal pinnae, which may themselves be pinnatifid, lack a stalk, the leaf blade is pointed at the tip but not drawn out at length, and there are generally fewer fronds. Its sori are dark brown, rather than cinnamon brown. [9] A. trudellii is fully pinnate in the lower half of the blade, and its pinnae are toothed. [10] A. × kentuckiense is also fully pinnate towards the base of the blade, with four to six pairs of pinnae, and the brown color of its stipe extends up into the basal part of the rachis. [11]
Lobed spleenwort was first recognized by Henry Muhlenberg in 1813, who considered it a variety of Asplenium rhizophyllum, although he did not provide a description distinguishing the variety. [12] In 1818, Thomas Nuttall observed that it was always distinguishable from A. rhizophyllum, and described it as a species under the name of Asplenium pinnatifidum. [13] Alphonso Wood used the name Camptosorus pinnatifidus for the species in 1870, [14] but this was not widely accepted.
Oliver A. Farwell, observing an unusual specimen of A. pinnatifidum, was led to suggest that the species might be a hybrid between American walking fern, Camptosorus rhizophyllus (now A. rhizophyllum), and ebony spleenwort (A. platyneuron). Such a hybrid, Scott's spleenwort (A. × ebenoides) was already known, but Farwell thought it bore a greater affinity to A. platyneuron while A. pinnatifidum had a greater affinity to A. rhizophyllum. [15] He was correct in viewing A. pinnatifidum as a hybrid descendant of A. rhizophyllum, but incorrect in identifying the other parent, and his suggestion was not widely taken up in the literature. Nor did his later attempt at subdividing Asplenium , moving A. pinnatifidum to a new genus as Chamaefilix pinnatifida in 1931, [16] meet with much favor.
As a member of the "Appalachian Asplenium complex", A. pinnatifidum readily acts as the progenitor of hybrids, as well. A. × gravesii was recognized as a hybrid of A. pinnatifidum and A. bradleyi by W. R. Maxon in 1918. [9] Edgar T. Wherry noted the similarities between A. montanum, A. pinnatifidum, and A. × trudellii in 1925, [10] and in 1936 concluded that Trudell's spleenwort was a hybrid between the first two. [17] That same year, A. kentuckiense was described by Thomas McCoy; Wherry identified it as a hybrid between A. pinnatifidum and A. platyneuron. [17] 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 A. rhizophyllum. [18]
In 1953, he 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. [19] His further experiments, published the following year, strongly suggested that A. pinnatifidum is an allotetraploid, the product of hybridization between A. montanum and A. rhizophyllum to form a sterile diploid, followed by chromosome doubling that restored fertility. [20] However, the hypothesized sterile diploid has never been found. [2] [a] Partial pairing of homologous chromosomes in A. × gravesii and A. × trudellii confirmed A. montanum parentage for A. pinnatifidum, [21] while an artificial hybrid between A. pinnatifidum and Tutwiler's spleenwort (A. tutwilerae) helped confirm their shared A. rhizophyllum parentage. [22]
Wagner's conclusions as to the parentage of A. pinnatifidum were supported by later chromatographic analyses, in which the chromatograms of A. pinnatifidum contained all the compounds detected in the chromatograms of both parents. [23] In 1985, an allozyme analysis confirmed the hybrid parentage of the species, [24] and revealed that A. pinnatifidum had probably originated independently through chromosome doubling at more than one locality. [25]
In 1956, C. V. Morton pointed out that, as A. pinnatifidum had been shown to arise from hybridization between walking fern and mountain spleenwort, it would constitute an intergeneric hybrid if walking fern was placed in the genus Camptosorus (as Camptosorus rhizophyllus). The hybrid genus ×Asplenosorus had been published, but Morton noted that it lacked a Latin diagnosis and was therefore invalid under the International Code of Botanical Nomenclature; he preferred to continue recognizing Asplenium pinnatifidum in Asplenium. [26] The ICBN's rules were relaxed in 1972, and in 1974, John Mickel published Asplenosorus pinnatifidus as a new combination for the species to allow the continued recognition of Camptosorus. [27] Since then, phylogenetic studies have shown that Camptosorus nests within Asplenium, [28] [29] and current treatments do not recognize it as a separate genus. [2]
In addition to A. × gravesii, A. × trudellii, and A. × kentuckiense, A. pinnatifidum is known to be the parent of several other hybrid species. A sterile triploid hybrid, formed by the crossing of A. pinnatifidum with a diploid cytotype of maidenhair spleenwort (A. trichomanes ssp. trichomanes) was discovered in 1969 [30] and named A. × herb-wagneri in 1977. [31] [b] Finally, several unnamed hybrids have been grown in culture. The hybrid between A. pinnatifidum and A. tutwilerae showed a "nondescript" morphology intermediate between the two parents, with slightly narrower lobes and more dimorphic fronds than A. pinnatifidum, overall resembling A. × kentuckiense but with more regular lobes. [22] Another, between A. pinnatifidum and the tetraploid American hart's-tongue fern (A. scolopendrium var. americanum) yielded peculiar specimens with a long blade, similar in texture and doubled indusia to the hart's-tongue fern, but lengthened and tapering to a point, and not lobed except for two surprisingly large auricles at the base. [32]
Native to eastern North America, A. pinnatifidum occurs in the middle and southern Appalachian Mountains, from Pennsylvania and New Jersey southwest to Alabama and the northeastern corner of Mississippi. It is also found in the Shawnee Hills and to some extent in the Ozarks, with outlying occurrences in southeastern Oklahoma and in Iowa County, Wisconsin. [33] Early reports from New England proved to be variants of Scott's spleenwort, Asplenium × ebenoides. [34]
A. pinnatifidum can be found on acidic rocks, often in steep habitats, from altitudes of 0 to 1,000 meters (0 to 3,281 ft). [2] Sandstone is its usual substrate. [4] The soil formed when these rocks weather must be subacid (pH 4.5–5.0) to mediacid (pH 3.5–4.0) to support A. pinnatifidum. [35]
The species is considered apparently secure globally (G4), but is endangered in many parts of its range. NatureServe considers it to be critically imperiled (S1) in Illinois, Maryland, Mississippi, New Jersey, Oklahoma, South Carolina, and Wisconsin, imperiled (S2) in North Carolina, and vulnerable (S3) in Arkansas, Georgia, Pennsylvania, and Virginia. It is threatened by changes in land use, habitat fragmentation, and certain forest management practices. [1]
A. pinnatifidum can be cultivated in rock gardens and terraria. It prefers medium light and will grow on a moist soil or potting mixture. [36] Some authorities recommend adding sandstone chips to the soil. [4]
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.
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.
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".
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.
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.
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. Its leaves grow in clusters and are highly dissected into oblong segments, rather than the beadlike segments found in some other members of the genus. The axes of the leaves are dark and covered in long, flattened hairs. It is endemic to California, where it grows in rocky habitats, usually over limestone. The species was named in honor of its collector, Sarah Paxson Cooper; according to Daniel Cady Eaton, who described it in 1875, it was the first fern species to be named for a female botanist.
The fern genus Asplenium is well known for its hybridization capacity, especially in temperate zones.
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.
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.
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.
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.
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 × 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.
Argyrochosma peninsularis is a fern endemic to Baja California Sur. It grows in dry, rocky places. First described as a species in 1939, it was transferred to the new genus Argyrochosma in 1987, recognizing their distinctness from the "cloak ferns". A dusting of powdery material and the presence of occasional scales on the central axis of its leaves help distinguish it from related species.
Argyrochosma formosa is a fern known from eastern and central Mexico and Guatemala. It grows on rocky slopes, particularly on limestone. Unlike many members of the genus, it lacks white powder on the underside of its leaves. First described as a species in 1842, it was transferred to the new genus Argyrochosma in 1987, recognizing their distinctness from the "cloak ferns".