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Botrychium

{{Short description|Genus of ferns in the family Ophioglossaceae}}

{{Italic title}}

{{Automatic taxobox

| name = Moonwort

| image = Botrychium-4.jpg

| image_caption = Botrychium lunaria

| taxon = Botrychium

| authority = Sw.

| type_species = Botrychium lunaria

| type_species_authority = (L.) Swartz

| subdivision_ranks = Species

| subdivision = see text

| synonyms=

}}

Botrychium is a genus of ferns, seedless vascular plants in the family Ophioglossaceae.{{cite journal | last1 = Christenhusz | first1 = Maarten J. M. | last2 = Zhang| first2 = Xian-Chun | last3 = Schneider | first3 = Harald | year = 2011 | title = A linear sequence of extant families and genera of lycophytes and ferns | url = http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf | journal = Phytotaxa | volume = 19 | pages = 7–54 | doi = 10.11646/phytotaxa.19.1.2 }} Botrychium species are known as moonworts. They are small, with fleshy roots, and reproduce by spores shed into the air. One part of the leaf, the trophophore, is sterile and fernlike; the other, the sporophore, is fertile and carries the clusters of sporangia or spore cases. Some species only occasionally emerge above ground and gain most of their nourishment from an association with mycorrhizal fungi.

The circumscription of Botrychium is disputed between different authors; some botanists include the genera Botrypus and Sceptridium within Botrychium, while others treat them as distinct. The latter treatment is provisionally followed here.

Phylogeny

Phylogeny of Botrychium{{cite journal |last1=Nitta |first1=Joel H. |last2=Schuettpelz |first2=Eric |last3=Ramírez-Barahona |first3=Santiago |last4=Iwasaki |first4=Wataru |display-authors=et al. |year=2022 |title=An Open and Continuously Updated Fern Tree of Life |journal=Frontiers in Plant Science |volume=13 |issue= |page= 909768| doi=10.3389/fpls.2022.909768 |pmid= 36092417|pmc= 9449725|bibcode= |doi-access=free}}{{cite web|last1= |first1= |last2= |display-authors=et al. |year=2024 |title=Tree viewer: interactive visualization of FTOL |url=https://fernphy.github.io/viewer.html |version=FTOL v1.7.0 (GenBank release 261) |access-date=1 May 2025}}

{{Clade | style=font-size:90%;line-height:90%;width:600px

|1={{clade

|label1=section

|sublabel1=Lanceolatae

|1={{Clade

|1=B. matricariifolium {{au|(Döll) Braun ex Koch 1847}}{{cite journal|last=Muller|first=S.|title=The impact of a drought in spring on the sporulation of Botrychium matricariifolium (Retz) A. Br. in the Bitcherland (Northern Vosges, France)|journal=Acta Oecologica|year=1992|volume=13|pages=335–343}} (daisy-leaved/[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500282 B. matricariifolium] Flora of North America, www.eFloras.org 26 Dec 2011 matricary/chamomile moonwort/grapefern)

|2={{Clade

|1=B. pedunculosum {{au|Wagner 1986}} (stalked moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500290 B. pedunculosum] Flora of North America, www.eFloras.org 26 Dec 2011

|2={{Clade

|1={{Clade

|1=B. hesperium {{au|Maxon & Clausen) Wagner & Lellinger 1981}} (western moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500278 B. hesperium] Flora of North America, www.eFloras.org 26 Dec 2011

|2=B. pseudopinnatum {{au|Wagner 1990}} (false northwestern/[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500292 B. pseudopinnatum] Flora of North America, www.eFloras.org 26 Dec 2011 daisy-leaved grapefern)

}}

|2={{Clade

|1={{Clade

|1=B. acuminatum {{au|W.H.Wagner 1990}} (pointed moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500271 B. acuminatum] Flora of North America, www.eFloras.org 26 Dec 2011

|2=B. pinnatum {{au|H. St.John}} (northern moonwort[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500291 B. pinnatum] Flora of North America, www.eFloras.org 26 Dec 2011)

}}

|2={{Clade

|1=B. alaskense {{au|Wagner & Grant 2002}}[http://www.tropicos.org/Name/100367914 B. alaskense] Tropicos.org. Missouri Botanical Garden. 27 Dec 2011 (Alaska moonwort)[http://www.public.iastate.edu/~herbarium/botrychium/B-alaskense.pdf B. alaskense] Iowa State Herbarium 27 Dec 2011

|2={{Clade

|1=B. boreale {{au|Milde 1857}} (northern moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500273 B. boreale] Flora of North America, www.eFloras.org 26 Dec 2011

|2=B. lanceolatum {{au|(Gmel) Ångstr. 1854}} (triangle moonwort,[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002875 B. lanceolatum] Flora of North America, www.eFloras.org 26 Dec 2011 lance-leaved grapefern)

}}

}}

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}}

}}

}}

|label2=section

|sublabel2=Botrychium

|2={{Clade

|1={{clade

|1=B. echo {{Au|Wagner 1983}}[http://www.tropicos.org/Name/26604469 B. echo] Tropicos.org. Missouri Botanical Garden. 03 Jan 2012 (reflected grapfern)[http://www.fs.fed.us/r2/projects/scp/assessments/botrychiumecho.pdf B. echo] USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004

|2=B. michiganense Wagner ex Gilman, Farrar & Zika 2015 (Michigan moonwort)

}}

|2={{clade

|1={{clade

|1={{clade

|1=B. pumicola {{au|Coville ex Underw. 1900}}[http://www.tropicos.org/Name/26603636 B. pumicola] Tropicos.org. Missouri Botanical Garden. 03 Jan 2012 (pumice moonwort/grape-fern)[http://www.oregon.gov/ODA/PLANT/CONSERVATION/profile_bopu.shtml B. pumicola] Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012

|2={{Clade

|1=B. furculatum Popovich & Farrar 2020

|2=B. ×watertonense {{au|Wagner}} (Waterton grapefern){{PLANTS|id=BOWA2|taxon=Botrychium ×watertonense|access-date=27 Dec 2011}}

}}

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|2={{clade

|1=B. dusenii (Christ 1906) Alston 1960

|2={{Clade

|1=B. tunux {{au|Stensvold & Farrar 2002}}{{PLANTS|id=BOTU3|taxon=Botrychium tunux|access-date=27 Dec 2011}}[http://www.tropicos.org/Name/50270442 B. tunux] Tropicos.org. Missouri Botanical Garden. 26 Dec 2011 (moosewort)[http://www.public.iastate.edu/~herbarium/botrychium/B-tunux.pdf B. tunux] Iowa State Herbarium 03 Jan 2012

|2={{Clade

|1=B. lunaria {{au|(L.) Sw. 1801}} (common moonwort,[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002876 B. lunaria] Flora of North America, www.eFloras.org 26 Dec 2011 grapefern moonwort)

|2={{Clade

|1=B. neolunaria {{au|sp. nov. ined. Stensvold & Farrar 2008}} (common moonwort)[http://www.public.iastate.edu/~herbarium/botrychium/B-neolunaria.pdf B. neolunaria] Iowa State Herbarium 27 Dec 2011

|2={{Clade

|1=B. nordicum Stensvold & Farrar 2016

|2={{Clade

|1=B. crenulatum {{au|W.H.Wagner 1981}} (dainty[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500275 B. crenulatum] Flora of North America, www.eFloras.org 26 Dec 2011/crenulate moonwort)

|2=B. yaaxudakeit {{au|Stensvold & Farrar 2002}}{{PLANTS|id=BOYA|taxon=Botrychium yaaxudakeit|access-date=27 Dec 2011}}[http://www.tropicos.org/Name/50270445 B. yaaxudakeit] Tropicos.org. Missouri Botanical Garden. 26 Dec 2011 (giant/Yakutat moonwort)[http://www.public.iastate.edu/~herbarium/botrychium/B-yaaxudakeit.pdf B. yaaxudakeit] Iowa State Herbarium 27 Dec 2011

}}

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|2={{Clade

|1=B. minganense {{au|Vict. 1927}} (Mingan's moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500283 B. minganense] Flora of North America, www.eFloras.org 26 Dec 2011

|2={{Clade

|1={{Clade

|1=B. campestre {{au|Wagner & Farrar}} (prairie moonwort/[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500274 B. campestre] Flora of North America, www.eFloras.org 26 Dec 2011 dunewort, Iowa moonwort)[http://wisplants.uwsp.edu/scripts/SearchResults.asp?Genus=Botrychium B. campestre] {{webarchive|url=https://web.archive.org/web/20130317084213/http://wisplants.uwsp.edu/scripts/SearchResults.asp?Genus=Botrychium |date=2013-03-17 }} Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011

|2={{Clade

|1=B. gallicomontanum {{au|Farrar & Johnson-Groh 1991}} (Frenchman's Bluff moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500277 B. gallicomontanum] Flora of North America, www.eFloras.org 26 Dec 2011

|2=B. pallidum {{au|Wagner 1990}} (Pale moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500288 B. pallidum] Flora of North America, www.eFloras.org 26 Dec 2011

}}

}}

|2={{Clade

|1={{Clade

|1=B. ascendens {{au|W.H.Wagner 1986}} (upswept[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500270 B. ascendens] Flora of North America, www.eFloras.org 26 Dec 2011/triangle-lobed/upward-lobed moonwort)

|2=B. simplex {{au|E.Hitchc. 1823}} (least moonwort/grapefern,[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002879 B. simplex] Flora of North America, www.eFloras.org 26 Dec 2011 little grapefern)

}}

|2={{Clade

|1=B. lineare {{au|Wagner 1994}} (skinny moonwort,[http://www1.dnr.wa.gov/nhp/refdesk/fguide/pdf/botlin.pdf B. lineare] {{webarchive |url=https://web.archive.org/web/20120426072518/http://www1.dnr.wa.gov/nhp/refdesk/fguide/pdf/botlin.pdf |date=April 26, 2012 }} Washington State Department of Natural Resources 26-Dec-2011 narrowleaf grapefern){{PLANTS|id=BOLI7|taxon=Botrychium lineare|access-date=26 Dec 2011}}

|2={{Clade

|1=B. paradoxum {{au|Wagner 1981}} (paradox[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500289 B. paradoxum] Flora of North America, www.eFloras.org 26 Dec 2011/peculiar moonwort)

|2={{Clade

|1=B. spathulatum {{au|Wagner 1990}} (spatulate/[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500295 B. spathulatum] Flora of North America, www.eFloras.org 26 Dec 2011 spoon-leaved moonwort)

|2={{Clade

|1=B. montanum {{au|Wagner}} (western goblin,[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500284 B. montanum] Flora of North America, www.eFloras.org 26 Dec 2011 mountain moonwort)

|2=B. mormo {{au|Wagner 1981}} (little goblin moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500285 B. mormo] Flora of North America, www.eFloras.org 26 Dec 2011

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Unassigned species:

  • {{txil|Botrychium daucifolium|Wall. ex Hook. & Grev. 1830}}[http://www.tropicos.org/Name/26609745 B. daucifolium] Tropicos.org. Missouri Botanical Garden. 27 Dec 2011 (thin-leaved moonwort)[http://www.efloras.org/florataxon.aspx?flora_id=101&taxon_id=242308863 B. daucifolium] Taiwan Plant Names, www.eFlora.org 27 Dec 2011
  • Botrychium farrarii Legler & Popovich 2024
  • Botrychium onondagense Underw. 1903
  • Botrychium rubellum Stensvold & Farrar 2024
  • Botrychium socorrense {{au|W.H.Wagner 1989}}[http://www.tropicos.org/Name/26616257 B. socorrense] Tropicos.org. Missouri Botanical Garden. 26 Dec 2011 (Isla Socorro moonwort)
  • Botrychium sutchuanense Chien & Chun 1959
  • †Botrychium ternatopsis Kuzitchkina 1960
  • Botrychium tolucaense Wagner & Mickel 2004

Conservation

Moonworts can be found in many environments, including prairies, forests, and mountains. While some Botrychium species are quite rare, conservation efforts can be difficult. Determining the rarity of a species is complicated by the plants’ small leaves, which stand only 2-10 centimeters above the soil.{{cite journal | last1 = Johnson-Groh | first1 = C. L. | last2 = Lee | first2 = J. | year = 2002 | title = Phenology and demography of two species of Botrychium (Ophioglassaceae) | journal = American Journal of Botany | volume = 89 | issue = 10| pages = 1624–1633 | doi = 10.3732/ajb.89.10.1624 | pmid = 21665590 }} Even more of a challenge in obtaining an accurate population count is the genus's largely subterranean life cycle. The vast majority of any one population of moonworts actually exists below ground in banks consisting of several types of propagules. One type of propagule is the ungerminated spores, which must percolate through the soil beyond the reach of light in order to germinate. This presumably increases the probability that the spore will be in range of a mycorrhizal symbiont before it produces the tiny, roughly heart-shaped gametophyte, which also exists entirely below ground.{{cite journal | last1 = Whittier | first1 = D | year = 1973 | title = The effect of light and other factors on spore germination in Botrychium dissectum | journal = Can J Bot | volume = 51 | issue = 10| pages = 1791–1794 | doi = 10.1139/b73-230 | bibcode = 1973CaJB...51.1791W }} Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root.

Juvenile and dormant sporophytes can also be hidden in the soil for long periods of time. Mature sporophytes do not necessarily produce a leaf annually; they can remain viable underground for up to 10 years without putting up a photosynthetic component. This feat is made possible by their dependence on symbiotic partnership with AM fungi of the genus Glomus, which supply most fixed carbon for growth and reproduction.{{cite journal | last1 = Winther | first1 = J | last2 = Friedman | first2 = W | year = 2007 | title = Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae) | journal = Am J Bot | volume = 94 | issue = 7| pages = 1248–1255 | doi = 10.3732/ajb.94.7.1248 | pmid = 21636490 }}

This mycorrhizal dependence has also made lab cultivation of moonworts difficult. Thus far, only germination of the gametophyte has been successful.

References

{{Reflist|24em}}

External links

{{Commons category|Botrychium}}

  • [https://plants.usda.gov/core/profile?symbol=BOTRY USDA Plants Profile for Botrychium (grapefern)]
  • [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=104332 Efloras.org: Flora of North America, treatment of genus Botrychium]
  • [https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=17170#null ITIS.gov: List of Botrychium species] — with species links.

{{Plant classification}}

{{Fern classification}}

{{Taxonbar|from=Q1353804}}

{{Authority control}}

Category:Fern genera

Category:Ferns of the Americas

Category:Ferns of the United States

Category:Taxa named by Olof Swartz