Asplenium × gravesii

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Graves' spleenwort
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. ×gravesii
Binomial name
Asplenium ×gravesii
Synonyms

×Asplenosorus gravesii(Maxon) Mickel

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.

Description

Asplenium × gravesii is a small fern, whose fronds grow in loosely bundled tufts. Its stem below the leaf blade is a shiny purplish-brown, while the green, narrowly triangular blades are cut into pinnae near the base, which diminish into lobes in the upper part. [1] The fronds are monomorphic, showing little or no difference between sterile and fertile fronds. [lower-alpha 1] [2]

The fronds of A. × gravesii, which are 10 to 13 centimeters (3.9 to 5.1 in) long, are closely spaced along an upward-curving, densely scale-covered rhizome 1 to 1.5 centimeters (0.4 to 0.6 in) in length. The scales are long and narrow, about 5 millimeters (0.2 in) long and 0.5 millimeters wide at the base, coming to a pointed tip. They are dark reddish-brown in color and strongly clathrate (bearing a lattice-like pattern). The stipe (the stalk of the leaf, below the blade) is typically 2 to 7 centimeters (0.8 to 3 in) long, shiny and purplish-brown, sometimes becoming green near the base of the leaf blade. It is rounded below and flat or grooved above. [1]

The overall shape of the blade is narrowly triangular, from 3 to 8 centimeters (1 to 3 in) long and 1.5 to 2.8 centimeters (0.59 to 1.1 in) wide. The leaf tissue is of a medium texture (neither delicate nor leathery), with clavate (club-shaped) hairs on the underside becoming gland-tipped, narrow scales on the veins. The lower part of the blade is cut into pinnae, diminishing to lobes in the upper part. The basal pinnae are triangular in shape, roughly equilateral, and nearly heart-shaped at their base; [1] they are borne on short stalks. [3] [lower-alpha 2] They are shallowly rounded or toothed around the edges. In larger specimens, the basal pinnae bear a pair of rounded lobes at their base. Successive pinnae above the base are narrower and less deeply cut, gradually diminishing into fused lobes or overlapping. The lobes continue to the pointed tip of the blade. The rachis (central axis of the blade) is typically green and flat, with narrow wings, and a groove on the upper side near the base of the blade, where it becomes the stipe. Larger individuals may have a narrower, glossy-brown rachis. A few scales are present on stipe and rachis, becoming narrower and more twisted as they ascend from the rhizome. [1]

The abundant sori are dark brown and variable in shape, fusing with one another as they grow. The sori are covered with firm, white indusia. [1] In wild specimens, they are found beneath the costa (midrib of the pinnae). As a sterile tetraploid hybrid, the spores are seen to be misshapen and abortive under microscopic examination. The species has a chromosome number of 144 (2 × 72) in the sporophyte. [3]

Asplenium × gravesii can potentially be confused with its parental species or with a number of the other Asplenium hybrids in the Appalachian Asplenium complex. Smaller specimens are most similar to A. pinnatifidum, but can be distinguished from that species by their brown stipes (and often rachides), a smaller number of fronds in each tuft, and by the tip of their leaves (which is pointed, but not drawn out at length as in A. pinnatifidum). [1] A. × gravesii also has a slightly finer leaf texture, slightly sharper teeth on its leaves, and darker brown sori than A. pinnatifidum. [4] When compared to A. bradleyi, larger A. × gravesii individuals are most similar, but the blades are not so deeply cut (A. bradleyi being wholly pinnate) nor the pinnae toothed around the edges, the rachis shows at least some traces of winging, [1] and more of the rachis tends to be green. The leaf texture is somewhat more delicate than the leathery blades of A. bradleyi. [4]

Asplenium × gravesii differs from Trudell's spleenwort (A. × trudellii), another descendant of A. pinnatifidum, by having a blade broadest at the middle or between the middle and the base, rather than at the base itself, and by the presence of brown color throughout the stipe and sometimes into the rachis. In contrast to Boydston's spleenwort (A. × boydstoniae), A. × gravesii has fewer than fifteen pairs of pinnae, which are not sessile, and when dark color is present in the rachis, it covers less than seven-eights of that structure. The most similar hybrid to A. × gravesii is probably Kentucky spleenwort (A. × kentuckiense), a hybrid of A. pinnatifidum and ebony spleenwort (A. platyneuron). In A. × kentuckiense, the blade tapers at the base, the second and third pairs of pinnae being shorter than the fourth and fifth; in A. × gravesii, all these pairs are approximately equal in size. A. × kentuckiense takes on a somewhat papery textures when dried, while A. × gravesii is more leathery. Finally, the guard cells of the latter average 49 micrometers, slightly larger than the 46 micrometers of the former. [lower-alpha 3] As this character can only be examined by microscope, and the ranges of individual guard cell size overlap, [3] some care is required in its use; 30 measurements from a single pinna were used to obtain an average length in previous studies. It is particularly useful in determining the identity of dried material. [5]

Taxonomy

The first specimens of the fern to be recognized were collected by Edward Willis Graves in 1917 at Sand Mountain near Trenton, Georgia. While Graves at first thought they might be a variant form of A. pinnatifidum, further study in conjunction with William R. Maxon revealed them to be a hybrid between A. pinnatifidum and A. bradleyi, both of which occurred nearby. Maxon published a description of the new species in 1918. [1] Specimens in the herbarium of Harold W. Pretz, collected in 1913 along the lower Susquehanna River and labeled as "A. pinnatifidum", were subsequently identified as the new species. [4]

Thomas Darling Jr., successfully crossed A. bradleyi with A. pinnatifidum in cultivation from 1954 to 1955 to produce A. × gravesii. He supplied both the artificial crosses and live specimens collected at Sand Mountain to Herb Wagner for cytological studies. [6] Wagner had previously used the size of stomata in herbarium material to tentatively classify the species as a tetraploid. [5] Wagner and Darling were able to grow the wild and artificial specimens together. While they were clearly the same species, the cultured specimens had pinnae less broad, particularly at the base, more widely spaced, and less deeply toothed; the sori were also halfway between the costae and the edge of the leaf, rather than beneath the costa. As the specimens had been cultured under the same conditions, they attributed this to genetic variation within the two parent strains. They were also able to show that about half of the chromosomes typically paired for meiosis. Since A. bradleyi is a hybrid of mountain spleenwort (A. montanum) and A. platyneuron, while A. pinnatifidum is a hybrid of A. montanum and walking fern (A. rhizophyllum), half of the genetic material in A. × gravesii was ultimately contributed by A. montanum and should be able to pair, consistent with the results. [3] Chromatographic experiments reported in 1963 showed that, like A. × kentuckiense, chromatograms made from A. × gravesii contained all the compounds from the chromatograms of all three of its diploid ancestors: A. montanum, A. platyneuron, and A. rhizophyllum. [7]

In 1974, John Mickel published Asplenosorus gravesii as a new combination for the species to allow the continued recognition of the genus Camptosorus for the walking ferns. [8] Since then, phylogenetic studies have shown that Camptosorus nests within Asplenium, [9] [10] and current treatments do not recognize it as a separate genus. [11]

Distribution

In principle, A. × gravesii might be found anywhere the ranges of the parent species overlap: throughout the mid- to southern Appalachian Mountains and extending west through the Shawnee Hills into the Ozarks. In practice, its occurrences are highly scattered and rare, as the two parental species do not often occur adjacent to one another. It has been found in New Jersey, Pennsylvania, Maryland, Hocking County, Ohio, Illinois, Kentucky, Tennessee, Madison County, Missouri, Arkansas, Dade County, Georgia, and Alabama. [12] An occurrence in West Virginia reported in 1926 and 1938 was subsequently found to be A. × trudellii, and one from Virginia in 1944 to be A. × kentuckiense. [6] It has been noted as particularly abundant in the general vicinity of Natural Bridge State Resort Park. [13]

Ecology

Like the parental species, A. gravesii prefers acid soil; [4] in fact, it may tolerate only mediacid (pH 3.5–4.0) soils, while subacid (pH 4.5–5.0) soils are acceptable to both parents. [14] It usually grows, like the parents, in exposed sandstone cliff faces. [13]

Cultivation

A. gravesii was produced by artificial hybridization in 1954–1955 by Thomas Darling Jr., who provided a detailed account of the process. Spores of the two parental species were sown on damp peat moss and kept largely in the shade, except for a few hours of morning sun. Gametophytes developed, and the young sporophytes that grew from them were large enough to remove in autumn. They were transplanted into a well-drained, gravelly loam. They were raised in a humid terrarium exposed to morning sunlight, with a layer of tissue to shade them from direct sunlight. Of the A. gravesii in the mixed population, one reached maturity in August 1955; two others did not mature until June 1956. Difficulties were encountered due to aphid infestation and various diseases promoted by excess moisture. [6]

See also

Notes and references

Notes

  1. The "fertile" fronds are those bearing sori.
  2. Maxon describes the basal pinnae as sessile; Wagner and Darling provide drawings of the fronds of wild and artificial specimens, showing the stalks.
  3. This difference is associated with the difference in chromosome count; A. × kentuckiense is triploid and A. × gravesii tetraploid.

Related Research Articles

<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.

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

Asplenium trichomanes, the maidenhair spleenwort, is a small fern in the spleenwort genus Asplenium. It is a widespread and common species, occurring almost worldwide in a variety of rocky habitats. It is a variable fern with several subspecies.

<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 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.

<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.

Asplenium arcanum is a fern known from western Mexico and Nicaragua.

<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.

Myriopteris aemula, the Texas lip fern or rival lip fern, is a moderately-sized fern of Texas 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 aemula until 2013, when the genus Myriopteris was again recognized as separate from Cheilanthes. It typically grows on limestone rock.

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

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.

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

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".

References

Works cited

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