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''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.


Description

''Asplenium'' × ''trudellii'' is a small fern, with evergreen fronds growing in tufts. It is similar to ''A. pinnatifidum'', with a triangular leaf blade, pinnatifid (lobed) in the upper part, with a long, drawn-out tip. The fronds are monomorphic, showing little or no difference between sterile and fertile fronds. Fronds of a large specimen measured long. The
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...
is presumably covered in clathrate scales (bearing a lattice-like pattern), as in the other '' Asplenium''s, including its parent species. The stipe (the stalk of the leaf, below the blade) is green in color, except at the very base, where it is brown. The leaf blade has a narrowly triangular shape, and is widest at the base or the next pair of pinnae above the base. A large specimen was measured at across. The bottom half, more or less, of the blade is cut into pinnae, while the upper half is merely pinnatifid. The pinnae are rounded, and are broadest at the base or between the base and the middle. They are relatively widely spaced on the stem, and have lobes or teeth at their edges. The rachis (central axis of the leaf) is green, slender below with a wing on the top side. The leaf tissue is coriaceous (leathery) in texture. The sori are pale brown, resembling those of the parent species. The chromosome number of the
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote pr ...
is 2''n'' = 108; it is a triploid. As a hybrid triploid, ''A.'' × ''trudellii'' is unable to undergo meiosis to form spores, and its spores are typically abortive and sterile. Such was the case in examinations of Georgia, Tennessee, and Pennsylvania material by Wherry and Paul Kestner in 1932. Both found only sterile spores. However, the species has been observed, particularly along the lower
Susquehanna River The Susquehanna River (; Lenape: Siskëwahane) is a major river located in the Mid-Atlantic region of the United States, overlapping between the lower Northeast and the Upland South. At long, it is the longest river on the East Coast of the ...
, to form colonies, suggesting that it undergoes independent reproduction. This has been hypothesized to occur by apogamy, yielding triploid spores and gametophytes which then grow directly into a new generation of triploid sporophytes. This apogamous reproduction is thought to account for the formation of the one suspected hybrid descendant of ''A.'' × ''trudellii''. This specimen was collected in 1961 on a sandstone cliff at
Cumberland Falls State Resort Park Cumberland Falls State Resort Park is a park located just southwest of Corbin, Kentucky and is contained entirely within the Daniel Boone National Forest. The park encompasses and is named for its major feature, Cumberland Falls. The falls are ...
, and identified as ''A. pinnatifidum'' × ''trudellii''. This was hypothesized to be a pentaploid formed from spores of ''A. pinnatifidum'' and apogamous triploid spores of ''A.'' × ''trudellii'', but this was not cytologically verified. The presumed hybrid has oblong pinnae, on stalks of intermediate length between its two parents. They are more widely spaced on the frond than in either parent. It is similar in appearance to its parent, ''A. pinnatifidum''. The two can be distinguished by the narrow-stalked pinnae of ''A.'' × ''trudellii'', as the basal pinnae of ''A. pinnatifidum'' are broad-stalked or adnate (fused) with the rachis.


Taxonomy

''A.'' × ''trudellii'' was first distinguished from ''A. pinnatifidum'' in 1925 by
Edgar T. Wherry Edgar Theodore Wherry (1885–1982) was an American mineralogist, soil scientist and botanist. He had a deep interest in ferns and ''Sarracenia''. Wherry earned his bachelor's degree in chemistry in 1906 from the University of Pennsylvania. He r ...
. He chose for a type specimen a sample he had collected with Harry W. Trudell in July 1920 from Cully ravine, just below the Holtwood Dam. Wherry named the fern in honor of Trudell, a leather company purchasing agent and amateur botanist whom Wherry would later describe as his "favorite field companion" in hunting for ferns. Other
herbarium A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...
specimens from
York Furnace, Pennsylvania York is a cathedral city with Roman origins, sited at the confluence of the rivers Ouse and Foss in North Yorkshire, England. It is the historic county town of Yorkshire. The city has many historic buildings and other structures, such as a m ...
, Harpers Ferry, West Virginia, and Winston County, Alabama were retrospectively identified with the new taxon, as were specimens collected by Edward W. Graves at
Long Island, Alabama Long Island (also called Carpenter, Carpenter Station, and Carpenters Station)John Robert Kennamer: ''History of Jackson County'', Southern Printing & Publishing Co., Winchester, Tenn., 1935, p. 173. is an unincorporated community in the northeaste ...
in 1917. Wherry initially speculated that it was either a hybrid between ''A. montanum'' and ''A. pinnatifidum'', or a common descendant with them from a hypothesized acid-soil ancestor. The latter theory was soon discarded in favor of the former, which was also endorsed by Graves.
Herb Wagner Warren Herbert Wagner Jr. (August 29, 1920 – January 8, 2000) was an eminent American botanist who was trained at Berkeley with E.B. Copeland and lived most of his professional career in Michigan. History Wagner was instructed in the ways of ...
performed a cytological investigation in 1954 which showed that ''A.'' × ''trudellii'' was a sterile triploid, wherein about 72 chromosomes paired and about 36 remained unpaired. This confirmed its origins as a hybrid between ''A. montanum'' and ''A. pinnatifidum''. Because the latter is a tetraploid hybrid between ''A. montanum'' and walking fern (''A. rhizophyllum''), half of the chromosomes it contributed to ''A.'' × ''trudellii'' pair with those contributed by ''A. montanum'' and half do not. A subsequent
chromatographic In chemical analysis, chromatography is a laboratory technique for the separation of a mixture into its components. The mixture is dissolved in a fluid solvent (gas or liquid) called the ''mobile phase'', which carries it through a system (a ...
analysis showed that ''A.'' × ''trudellii'' produced chromatograms very similar to ''A. pinnatifidum''; these chromatograms contained all the spots seen on ''A. montanum'' and ''A. rhizophyllum'' chromatograms. (In other words, the chromatograms showed that both ''A. pinnatifidum'' and ''A.'' × ''trudellii'' were descended from ''A. montanum'' and ''A. rhizophyllum'', but could not discern the number of chromosomes contributed by each ancestor.) In 1974,
John Mickel John Mickel (born 28 January 1971) is a British professional stock car racing driver and commentator. He has raced in the NASCAR Craftsman Truck Series in the United States and the Pickup Truck Racing Series in the United Kingdom. He was also ...
published ''Asplenosorus trudellii'' as a new combination for the species to allow the continued recognition of the genus ''Camptosorus'' for the walking ferns. Since then, phylogenetic studies have shown that ''Camptosorus'' nests within ''Asplenium'', and current treatments do not recognize it as a separate genus.


Distribution and habitat

It is known from
Blairstown, New Jersey Blairstown is a township in Warren County, New Jersey. As of the 2020 U.S. census, Blairstown's population was 5,704. As of the 2010 United States Census, the township's population was 5,967Appalachian Mountains through Pennsylvania, Maryland, Virginia, West Virginia, Ohio, North Carolina, Kentucky, Tennessee, Georgia and Alabama, and in the Shawnee Hills in Jackson County, Illinois. A population was discovered in
Baxter County, Arkansas Baxter County is a county in the U.S. state of Arkansas. As of the 2020 census, the county's population was 41,627. The county seat is Mountain Home. It is Arkansas's 66th county, formed on March 24, 1873, and named for Elisha Baxter, the tent ...
in 2005. Both parent species prefer acidic soil, typically growing on sandstone. The type specimen was found growing in ( pH 3.5–4.0) soil, on "
quartzose Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
mica-schist ledges". Other occurrences were in "low soil" on gneiss and in cracks of sandstone cliffs.


Cultivation

It has been successfully cultivated in a pot on a mixture of acidic peat and sand.


See also

*
Asplenium hybrids The fern genus ''Asplenium'' is well known for its hybridization capacity, especially in temperate zones. Appalachian ''Asplenium'' hybrid complex The Appalachian hybrid complex in ''Asplenium'' is a polyploid complex composed of 3 diploid spec ...


Notes and references


Notes


References


Works cited

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External links


NatureServe ExplorerType specimen
at JSTOR Plant Science
Pressed fronds
specimen from Georgia
Alan Cressler's comparison of fronds of ''A. × trudellii'' with parental species
{{DEFAULTSORT:Asplenium trudellii trudellii Hybrid plants Plants described in 1925