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The fern genus Asplenium is well known for its hybridization capacity, especially in temperate zones.
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The fern genus Asplenium is well known for its hybridization capacity, especially in temperate zones.
The Appalachian hybrid complex in Asplenium is a polyploid complex composed of three diploid species, three allotetraploid species formed by hybridisation and chromosome doubling from the former, and assorted diploid, triploid and tetraploid hybrids between the six species.
The three diploid parent species are mountain spleenwort ( Asplenium montanum ), ebony spleenwort ( Asplenium platyneuron ), and American walking fern ( Asplenium rhizophyllum ). Their chromosome complements are abbreviated "MM", "PP" and "RR", respectively. Three possible sterile diploid hybrids can form from their hybridization:
In addition, A. platyneuron and A. rhizophyllum, which have a more extensive range, hybridize with other spleenwort species outside the Appalachian complex.
At some point in the past, each of the sterile diploid hybrids experienced a chromosome doubling event that made them fertile allotetraploids; allozyme analysis shows that this happened on several independent occasions for A. montanum × platyneuron and A. montanum × rhizophyllum. These doubling events gave rise to the following tetraploids:
| Common name | Scientific name | Parents | Chromosomes | Notes/Distribution |
|---|---|---|---|---|
| Bradley's spleenwort | Asplenium bradleyi | A. montanum × platyneuron | MMPP | The Appalachians, Ozarks, and Ouachita Mountains. |
| Lobed spleenwort | Asplenium pinnatifidum | A. montanum × rhizophyllum | MMRR | The Appalachians and the Shawnee Hills. |
| Tutwiler's spleenwort | Asplenium tutwilerae | A. platyneuron × rhizophyllum | PPRR | Havana Glen, Alabama |
Since these allotetraploids are fertile, they are capable of crossing with each other and with the three diploid species to form the following sterile triploid and tetraploid hybrids, most of which are quite rare and occur sporadically where their parents' ranges overlap:
| Common name | Scientific name | Parents | Chromosomes | Ploidy | Notes/Distribution |
|---|---|---|---|---|---|
| Wherry's spleenwort | Asplenium × wherryi | A. bradleyi × montanum | MMP | triploid | |
| A. bradleyi × platyneuron | MPP | triploid | Believed to have been collected once at an early date in Pennsylvania; the site is now destroyed. Also reported from Sequatchie County, Tennessee. | ||
| Graves' spleenwort | Asplenium × gravesii | A. bradleyi × pinnatifidum | MMPR | tetraploid | |
| Trudell's spleenwort | Asplenium × trudellii | A. pinnatifidum × montanum | MMR | triploid | The Appalachians and the Shawnee Hills. |
| Kentucky spleenwort | Asplenium × kentuckiense | A. pinnatifidum × platyneuron | MPR | triploid | |
| A. tutwilerae × pinnatifidum | MPRR | tetraploid | Created in culture. | ||
| Boydston's spleenwort | Asplenium × boydstoniae | A. tutwilerae × platyneuron | PPR | triploid | Havana Glen, Alabama |
| A. tutwilerae × rhizophyllum | PRR | triploid | Created accidentally in culture. |
19 hybrids within the genus have been described from New Zealand, but have not been given binomials.
Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one from each of two parents; each set contains the same number of chromosomes, and the chromosomes are joined in pairs of homologous chromosomes. However, some organisms are polyploid. Polyploidy is especially common in plants. Most eukaryotes have diploid somatic cells, but produce haploid gametes by meiosis. A monoploid has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Males of bees and other Hymenoptera, for example, are monoploid. Unlike animals, plants and multicellular algae have life cycles with two alternating multicellular generations. The gametophyte generation is haploid, and produces gametes by mitosis; the sporophyte generation is diploid and produces spores by meiosis.
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 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.
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.
Hybridization and polyploidy are common phenomena in ferns, and the genus Dryopteris is known to be one of the most freely-hybridizing fern genera. North American botanists recognized early that there were close relationships between many of the species of Dryopteris on the continent, and that these relationships reflected hybrid ancestry. The complex includes six sexual diploid parents, six sexual allopolyploids, and numerous sterile hybrids at various ploidal levels.
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.
Polypodium appalachianum is a fern species native to eastern North America. Sometimes called the Appalachian polypody or Appalachian rockcap fern, it is very similar in appearance to Polypodium virginianum. For years, P. virginianum—long considered a variety of the British Polypodium vulgare—was recognized as having cryptic races, with diploid, triploid, and tetraploid representatives. Since the triploid specimens bore abortive spores, it was apparently the hybrid between the diploid and tetraploid groups. In 1991, it was resolved that the type of P. virginianum was the tetraploid series, and that it is an allotetraploid species of hybrid origin, with the diploid species as one parent. The diploid species was then named P. appalachianum. The other parent of P. virginianum was found to be Polypodium sibiricum. The tetraploid of hybrid derivation tolerates warmer climates than either parent.
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.
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 onopteris, known as the Irish spleenwort or western black spleenwort, is a species of fern mostly found throughout the Mediterranean Basin but also around the Eastern Atlantic.
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.