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Lobed spleenwort was first recognized by [[Henry Muhlenberg]] in 1813, who considered it a variety of ''Asplenium rhizophyllum''.{{sfn|Muhlenberg|1813|p=102}} In 1818, Nuttall observed that it was always distinguishable from ''A. rhizophyllum'', and raised it to a species as ''Asplenium pinnatifidum''.{{sfn|Nuttall|1818|p=251}} [[Alphonso Wood]] used the name ''Camptosorus pinnatifidus'' for the species in 1870,{{sfn|Wood|1870|p=425}}, but this was not widely accepted.
Lobed spleenwort was first recognized by [[Henry Muhlenberg]] in 1813, who considered it a variety of ''Asplenium rhizophyllum''.{{sfn|Muhlenberg|1813|p=102}} In 1818, Nuttall observed that it was always distinguishable from ''A. rhizophyllum'', and raised it to a species as ''Asplenium pinnatifidum''.{{sfn|Nuttall|1818|p=251}} [[Alphonso Wood]] used the name ''Camptosorus pinnatifidus'' for the species in 1870,{{sfn|Wood|1870|p=425}}, but this was not widely accepted.


As a member of the "Appalachian ''Asplenium'' complex", ''A.&nbsp;pinnatifidum'' readily participates in the formation of [[Asplenium hybrids|hybrids]]. Graves's spleenwort ([[Asplenium × gravesii|''A.&nbsp;×&nbsp;gravesii'']]) was recognized as a hybrid of ''A.&nbsp;pinnatifidum'' and Bradley's spleenwort (''A.&nbsp;bradleyi'') by [[William Ralph Maxon|W. R. Maxon]] in 1918.{{sfn|Maxon|1918}} [[Edgar T. Wherry]] noted the similarities between ''A.&nbsp;montanum'', ''A.&nbsp;pinnatifidum'', and Trudell's spleenwort ([[Asplenium × trudellii|''A.&nbsp;×&nbsp;trudellii'']]) in 1925,{{sfn|Wherry|1925}} and in 1936 concluded that Trudell's spleenwort was a hybrid between the first two.{{sfn|Wherry|Gray|1936}} In 1951, [[Herb Wagner]], while reviewing [[Irene Manton]]'s ''Problems of Cytology and Evolution in the Pteridophyta'', suggested in passing that ''A.&nbsp;pinnatifidum'' itself might represent a hybrid between ''A.&nbsp;montanum'' and the American walking fern, ''Camptosorus rhizophyllus'' (now [[Asplenium rhizophyllum|''A.&nbsp;rhizophyllum'']]).{{sfn|Wagner|1951}}<!-- his ''Evolution'' review in 1950 predates this? -->
⚫
In 1956, C. V. Morton pointed out that, as the species 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''.{{sfn|Morton|1956}} The ICBN's rules were relaxed in 1972, and in 1974, [[John Thomas Mickel|John Mickel]] published ''Asplenosorus pinnatifidus'' as a new combination for the species to allow the continued recognition of ''Camptosorus''.{{sfn|Mickel|1974}} Since then, phylogenetic studies have shown that ''Camptosorus'' nests within ''Asplenium'',{{sfn|Murakami|Nogami|Watanabe|Iwatsuki|1999}}{{sfn|Schneider et al.|2004}} and current treatments do not recognize it as a separate genus.{{sfn|Wagner|Moran|Werth|1993}}


As part of the "Appalachian ''Asplenium'' complex", ''A.&nbsp;pinnatifidum'' readily forms hybrids. In 1951, [[Herb Wagner]], while reviewing [[Irene Manton]]'s ''Problems of Cytology and Evolution in the Pteridophyta'', suggested in passing that ''A.&nbsp;pinnatifidum'' itself might represent a hybrid between ''A.&nbsp;montanum'' and the American walking fern, ''Camptosorus rhizophyllus'' (now [[Asplenium rhizophyllum|''A.&nbsp;rhizophyllum'']]).{{sfn|Wagner|1951}}<!-- his ''Evolution'' review in 1950 predates this? --> His experiments, published in 1954, strongly suggested that ''A.&nbsp;pinnatifidum'' was an [[allotetraploid]]s, the product of hybridization between ''A.&nbsp;montanum'' and ''A.&nbsp;rhizophyllum'' to form a sterile diploid, followed by chromosome doubling that restored fertility.{{sfn|Wagner|1954}}
In 1953, he made chromosome counts of ''A.&nbsp;×&nbsp;trudellii'', which had been classified by some simply as a variety of ''A.&nbsp;pinnatifidum''. As ''A.&nbsp;pinnatifidum'' proved to be a [[tetraploid]] while ''A.&nbsp;montanum'' was a [[diploid]], a hybrid between them would be a [[triploid]], and Wagner showed that this was in fact the case for ''A.&nbsp;×&nbsp;trudellii''.{{sfn|Wagner|1953}} His further experiments, published the following year, strongly suggested that ''A.&nbsp;pinnatifidum'' is an [[allotetraploid]], the product of hybridization between ''A.&nbsp;montanum'' and ''A.&nbsp;rhizophyllum'' to form a sterile diploid, followed by chromosome doubling that restored fertility.{{sfn|Wagner|1954}} However, the hypothesized sterile diploid has never been found.{{sfn|Wagner|Moran|Werth|1993}} <!-- see also A. gravesii work in 1957, Brittonia -->

⚫
In 1956, C. V. Morton pointed out that, as ''A.&nbsp;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''.{{sfn|Morton|1956}} The ICBN's rules were relaxed in 1972, and in 1974, [[John Thomas Mickel|John Mickel]] published ''Asplenosorus pinnatifidus'' as a new combination for the species to allow the continued recognition of ''Camptosorus''.{{sfn|Mickel|1974}} Since then, phylogenetic studies have shown that ''Camptosorus'' nests within ''Asplenium'',{{sfn|Murakami|Nogami|Watanabe|Iwatsuki|1999}}{{sfn|Schneider et al.|2004}} and current treatments do not recognize it as a separate genus.{{sfn|Wagner|Moran|Werth|1993}}

In addition to ''A.&nbsp;×&nbsp;gravesii'' and ''A.&nbsp;×&nbsp;trudellii'', ''A.&nbsp;pinnatifidum'' is known to be the parent of several other hybrid species. A sterile triploid hybrid, formed by the crossing of ''A.&nbsp;pinnatifidum'' with a diploid cytotype of maidenhair spleenwort (''A.&nbsp;trichomanes'' ssp. ''trichomanes'') was discovered in 1969{{sfn|Wagner|Wagner|1969}} and named [[Asplenium × herb-wagneri|''A.&nbsp;×&nbsp;herb-wagneri'']] in 1977.{{sfn|Taylor|Mohlenbrock|1977}}{{efn|''Asplenium stotleri'', previously thought to be ''A.&nbsp;pinnatifidum × trichomanes'', was shown to be a variant of ''A.&nbsp;bradleyi''.}} Kentucky spleenwort ([[Asplenium × kentuckiense|''A.&nbsp;×&nbsp;kentuckiense'']]), also a sterile triploid, is formed by crossing ''A.&nbsp;pinnatifidum'' with ebony spleenwort ([[Asplenium platyneuron|''A.&nbsp;platyneuron'']]).{{sfn|Wagner|1954}}<!-- see also Smith, Bryant & Tate, Brittonia 1961 --> Finally, an unnamed hybrid between ''A.&nbsp;pinnatifidum'' and Tutwiler's spleenwort ([[Asplenium tutwilerae|''A.&nbsp;tutwilerae'']]) has been grown in culture.{{sfn|Wagner|Boydston|1961}}

==Notes==
{{Notelist}}


==Citations==
==Citations==
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==References==
==References==
*{{cite journal | ref=harv | last=Maxon | first=W.R. | authorlink=William Ralph Maxon | title=A new hybrid ''Asplenium'' | journal=[[American Fern Journal]] | volume=8 | issue=1 | year=1918 | pages=1–3 | url=http://biodiversitylibrary.org/page/30816807}}
*{{cite journal | ref=harv | last=Mickel | first=John T. | authorlink=John Thomas Mickel | title=The status and composition of ''Asplenosorus'' | journal=American Fern Journal | year=1974 | volume=64 | issue=4 | page=119 | url=http://biodiversitylibrary.org/page/32084449}}
*{{cite journal | ref=harv | last=Mickel | first=John T. | authorlink=John Thomas Mickel | title=The status and composition of ''Asplenosorus'' | journal=American Fern Journal | year=1974 | volume=64 | issue=4 | page=119 | url=http://biodiversitylibrary.org/page/32084449}}
*{{cite journal | ref=harv | last=Morton | first=C. V. | authorlink=Conrad Vernon Morton | title=A new name for an ''Asplenium'' hybrid | journal=[[American Fern Journal]] | volume=46 | issue=4 | year=1956 | pages=152–155 | url=http://biodiversitylibrary.org/page/31990471}}
*{{cite journal | ref=harv | last=Morton | first=C. V. | authorlink=Conrad Vernon Morton | title=A new name for an ''Asplenium'' hybrid | journal=[[American Fern Journal]] | volume=46 | issue=4 | year=1956 | pages=152–155 | url=http://biodiversitylibrary.org/page/31990471}}
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*{{cite book | ref=harv | last=Nuttall | first=Thomas | authorlink=Thomas Nuttall | title=The genera of North American plants | volume= | year=1818 | publisher=D. Heartt | location=Philadelphia | url=http://botanicus.org/item/31753002739776}}
*{{cite book | ref=harv | last=Nuttall | first=Thomas | authorlink=Thomas Nuttall | title=The genera of North American plants | volume= | year=1818 | publisher=D. Heartt | location=Philadelphia | url=http://botanicus.org/item/31753002739776}}
*{{cite journal | last1=Schneider | first1=Harald | last2=Russell | first2=Steve J. | last3=Cox | first3=Cymon J. | last4=Henderson | first4=Sally | last5=Rumsey | first5=Fred | last6=Barrett | first6=John | last7=Gibby | first7=Mary | last8=Vogel | first8=Johannes C. | journal=[[Systematic Botany]] | volume=29 | issue=2 | year=2004 | pages=260–274 | doi=10.1600/036364404774195476 | ref = {{harvid|Schneider et al.|2004}}}}
*{{cite journal | last1=Schneider | first1=Harald | last2=Russell | first2=Steve J. | last3=Cox | first3=Cymon J. | last4=Henderson | first4=Sally | last5=Rumsey | first5=Fred | last6=Barrett | first6=John | last7=Gibby | first7=Mary | last8=Vogel | first8=Johannes C. | journal=[[Systematic Botany]] | volume=29 | issue=2 | year=2004 | pages=260–274 | doi=10.1600/036364404774195476 | ref = {{harvid|Schneider et al.|2004}}}}
*{{cite journal | ref=harv | last1=Taylor | first1=W. Carl | last2=Mohlenbrock | first2=Robert H. | title=''Asplenium × herb-wagneri''—a collective epithet for ''A. pinnatifidum × trichomanes'' | journal=[[American Fern Journal]] | volume=67 | issue=3 | year=1977 | pages=65–67 | url=http://biodiversitylibrary.org/page/31876549}}
*{{cite journal | ref=harv | last=Wagner | first=Warren H., Jr. | authorlink=Warren H. Wagner | title=Recent Fern Literature | journal=[[American Fern Journal]] | volume=41 | issue=3 | year=1951 | pages=90–93 | url=http://biodiversitylibrary.org/page/32415676}}
*{{cite journal | ref=harv | last=Wagner | first=Warren H., Jr. | authorlink=Warren H. Wagner | title=A cytological study of the Appalachian spleenworts | journal=[[American Fern Journal]] | volume=43 | issue=3 | year=1953 | pages=109–114 | url=http://biodiversitylibrary.org/page/32266446}}
*{{cite journal | ref=harv | last=Wagner | first=Warren H., Jr. | authorlink=Warren H. Wagner | title=Reticulate evolution in the Appalachian ''Asplenium''s | journal=[[Evolution (journal)|Evolution]] | volume=8 | year=1954 | pages=103–118}}
*{{citation | last1=Wagner | first1=Warren H., Jr. | authorlink1=Warren H. Wagner | last2=Boydston | first2=Kathryn E. | title=A new hybrid showing homology between ''Asplenium ebenoides'' and ''A. pinnatifidum'' | journal=[[Brittonia]] | volume=13 | issue=3 | year=1961 | pages=286–289 | doi=10.2307/2805344}}
*{{cite book | ref=harv |year=1993 |title=Flora of North America | chapter=''Asplenium pinnatifidum'' |editors=Flora of North America Editorial Committee | last1=Wagner | first1=Warren H., Jr. | authorlink1=Warren H. Wagner | last2=Moran | first2=Robbin C. | last3=Werth | first3=Charles R. |encyclopedia=Flora of North America North of Mexico |volume=2 |publisher=Oxford University Press |location=New York and Oxford |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200004154 |accessdate=2012-08-19}}
*{{cite book | ref=harv |year=1993 |title=Flora of North America | chapter=''Asplenium pinnatifidum'' |editors=Flora of North America Editorial Committee | last1=Wagner | first1=Warren H., Jr. | authorlink1=Warren H. Wagner | last2=Moran | first2=Robbin C. | last3=Werth | first3=Charles R. |encyclopedia=Flora of North America North of Mexico |volume=2 |publisher=Oxford University Press |location=New York and Oxford |url=http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200004154 |accessdate=2012-08-19}}
*{{cite journal | ref=harv | last1=Wagner | first1=Warren H., Jr. | authorlink1=Warren H. Wagner | last2=Wagner | first2=Florence S. | title=A new natural hybrid in the Appalachian ''Asplenium'' complex and its taxonomic significance | journal=[[Brittonia]] | volume=21 | issue=2 | year=1969 | pages=178–186 | doi=10.2307/2805524}}
*{{cite journal | ref=harv | last=Wherry | first=Edgar T. | authorlink=Edgar T. Wherry | title=The Appalachian ''Asplenium''s | journal=[[American Fern Journal]] | volume=15 | issue=2 | year=1925 | pages=48–54 | url=http://biodiversitylibrary.org/page/31033989 | jstor=1544001}}
*{{cite journal | ref=harv | last1=Wherry | first1=Edgar T. | authorlink1=Edgar T. Wherry | last2=Gray | first2=William D. | title=Variety of some Appalachian ''Asplenium''s | journal=[[American Fern Journal]] | volume=26 | issue=3 | year=1936 | pages=77–86 | url=http://biodiversitylibrary.org/page/32264333}}
*{{cite book | ref=harv | last=Wood | first=Alphonso | authorlink=Alphonso Wood | title=The American botanist and florist | publisher=A.S. Barnes & Co. | year=1870 | url=http://www.archive.org/stream/americanbotanist00wood#page/n5/mode/2up}}
*{{cite book | ref=harv | last=Wood | first=Alphonso | authorlink=Alphonso Wood | title=The American botanist and florist | publisher=A.S. Barnes & Co. | year=1870 | url=http://www.archive.org/stream/americanbotanist00wood#page/n5/mode/2up}}



[[Category:Asplenium|pinnatifidum]]
[[Category:Asplenium|pinnatifidum]]

Revision as of 19:03, 19 August 2012

Lobed spleenwort

Apparently Secure  (NatureServe)
Scientific classification
Kingdom:
Plantae
Division:
Pteridophyta
Class:
Pteridopsida
Order:
Blechnales
Family:
Aspleniaceae
Genus:
Asplenium
Species:
A. pinnatifidum
Binomial name
Asplenium pinnatifidum
Synonyms

Asplenium rhizophyllum var. pinnatifidum Muhl.
×Asplenosorus pinnatifidus (Nutt.) Mickel
Camptosorus pinnatifidus (Nutt.) Alph.Wood

Asplenium pinnatifidum is known as the lobed spleenwort or pinnatifid spleenwort. It is an Appalachian rock fern, growing in rock crevices in moderately acid to subacid strata. It is often confused with Asplenium bradleyi.

This fern species is an allotetraploid of hybrid origin. Its parent species are mountain spleenwort (Asplenium montanum) and walking fern (Asplenium rhizophyllum). The genus Asplenium is known for its hybrids.

Taxonomy

Lobed spleenwort was first recognized by Henry Muhlenberg in 1813, who considered it a variety of Asplenium rhizophyllum.[1] In 1818, Nuttall observed that it was always distinguishable from A. rhizophyllum, and raised it to a species as Asplenium pinnatifidum.[2] Alphonso Wood used the name Camptosorus pinnatifidus for the species in 1870,[3], but this was not widely accepted.

As a member of the "Appalachian Asplenium complex", A. pinnatifidum readily participates in the formation of hybrids. Graves's spleenwort (A. × gravesii) was recognized as a hybrid of A. pinnatifidum and Bradley's spleenwort (A. bradleyi) by W. R. Maxon in 1918.[4] Edgar T. Wherry noted the similarities between A. montanum, A. pinnatifidum, and Trudell's spleenwort (A. × trudellii) in 1925,[5] and in 1936 concluded that Trudell's spleenwort was a hybrid between the first two.[6] 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).[7]

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.[8] 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.[9] However, the hypothesized sterile diploid has never been found.[10]

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.[11] 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.[12] Since then, phylogenetic studies have shown that Camptosorus nests within Asplenium,[13][14] and current treatments do not recognize it as a separate genus.[10]

In addition to A. × gravesii and A. × trudellii, 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[15] and named A. × herb-wagneri in 1977.[16][a] Kentucky spleenwort (A. × kentuckiense), also a sterile triploid, is formed by crossing A. pinnatifidum with ebony spleenwort (A. platyneuron).[9] Finally, an unnamed hybrid between A. pinnatifidum and Tutwiler's spleenwort (A. tutwilerae) has been grown in culture.[17]

Notes

  1. ^ Asplenium stotleri, previously thought to be A. pinnatifidum × trichomanes, was shown to be a variant of A. bradleyi.

Citations

  1. ^ Muhlenberg 1813, p. 102.
  2. ^ Nuttall 1818, p. 251.
  3. ^ Wood 1870, p. 425.
  4. ^ Maxon 1918.
  5. ^ Wherry 1925.
  6. ^ Wherry & Gray 1936.
  7. ^ Wagner 1951.
  8. ^ Wagner 1953.
  9. ^ a b Wagner 1954.
  10. ^ a b Wagner, Moran & Werth 1993.
  11. ^ Morton 1956.
  12. ^ Mickel 1974.
  13. ^ Murakami et al. 1999.
  14. ^ Schneider et al. 2004.
  15. ^ Wagner & Wagner 1969.
  16. ^ Taylor & Mohlenbrock 1977.
  17. ^ Wagner & Boydston 1961.

References


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