Editing 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 = [[Olof Swartz|Sw.]]
| type_species = ''[[Botrychium lunaria]]''
| type_species_authority = (L.) Swartz
| subdivision_ranks = [[Species]]
| subdivision = [[#Phylogeny|see text]]
| synonyms=
}}

'''''Botrychium''''' is a genus of [[fern]]s, seedless [[vascular plant]]s in the family [[Ophioglossaceae]].<ref name="Christenhusz_2011b">{{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 }}</ref> ''Botrychium'' species are known as '''moonworts'''. They are small, with fleshy [[root]]s, and reproduce by [[spore]]s 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 [[fungus|fungi]].

<!-- [[Image:BotrychiumLunaria1.jpg|left|frame|Moonwort (''Botrychium lunaria'')]] [messes up layout real bad. Not in Commons, so it is left here for species article] -->
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''<ref>{{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}}</ref><ref>{{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}}</ref>
{{Clade | style=font-size:90%;line-height:90%;width:600px
|1={{clade
  |label1=section
  |sublabel1=''Lanceolatae''
  |1={{Clade
    |1=''[[Botrychium matricariifolium|B. matricariifolium]]'' {{au|(Döll) Braun ex Koch 1847}}<ref>{{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}}</ref> (daisy-leaved/<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500282 ''B. matricariifolium''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> matricary/chamomile moonwort/grapefern)
    |2={{Clade
      |1=''[[Botrychium pedunculosum|B. pedunculosum]]'' {{au|Wagner 1986}} (stalked moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500290 ''B. pedunculosum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
      |2={{Clade
        |1={{Clade
          |1=''[[Botrychium hesperium|B. hesperium]]'' {{au|Maxon & Clausen) Wagner & Lellinger 1981}} (western moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500278 ''B. hesperium''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
          |2=''[[Botrychium pseudopinnatum|B. pseudopinnatum]]'' {{au|Wagner 1990}} (false northwestern/<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500292 ''B. pseudopinnatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> daisy-leaved grapefern)
           }}
        |2={{Clade
          |1={{Clade
            |1=''[[Botrychium acuminatum|B. acuminatum]]'' {{au|[[W.H.Wagner]] 1990}} (pointed moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500271 ''B. acuminatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
            |2=''[[Botrychium pinnatum|B. pinnatum]]'' {{au|[[Harold St. John|H. St.John]]}} (northern moonwort<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500291 ''B. pinnatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>)
           }}
          |2={{Clade
            |1=''[[Botrychium alaskense|B. alaskense]]'' {{au|Wagner & Grant 2002}}<ref>[http://www.tropicos.org/Name/100367914 ''B. alaskense''] Tropicos.org. Missouri Botanical Garden. 27 Dec 2011</ref> (Alaska moonwort)<ref>[http://www.public.iastate.edu/~herbarium/botrychium/B-alaskense.pdf ''B. alaskense''] Iowa State Herbarium 27 Dec 2011</ref>
            |2={{Clade
              |1=''[[Botrychium boreale|B. boreale]]''  {{au|Milde 1857}} (northern moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500273 ''B. boreale''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
              |2=''[[Botrychium lanceolatum|B. lanceolatum]]'' {{au|(Gmel) Ångstr. 1854}} (triangle moonwort,<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002875 ''B. lanceolatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> lance-leaved grapefern)
             }}
           }}
         }}
       }}
     }}
   }}
  |label2=section
  |sublabel2=''Botrychium''
  |2={{Clade
    |1={{clade
      |1=''[[Botrychium echo|B. echo]]'' {{Au|Wagner 1983}}<ref>[http://www.tropicos.org/Name/26604469 ''B. echo''] Tropicos.org. Missouri Botanical Garden. 03 Jan 2012</ref> (reflected grapfern)<ref>[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</ref>
      |2=''[[Botrychium michiganense|B. michiganense]]'' <small>Wagner ex Gilman, Farrar & Zika 2015</small> (Michigan moonwort)
       }}
    |2={{clade
      |1={{clade
        |1={{clade
          |1=''[[Botrychium pumicola|B. pumicola]]'' {{au|Coville ex Underw. 1900}}<ref>[http://www.tropicos.org/Name/26603636 ''B. pumicola''] Tropicos.org. Missouri Botanical Garden. 03 Jan 2012</ref> (pumice moonwort/grape-fern)<ref>[http://www.oregon.gov/ODA/PLANT/CONSERVATION/profile_bopu.shtml ''B. pumicola''] Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012</ref>
          |2={{Clade
            |1=''[[Botrychium furculatum|B. furculatum]]'' <small>Popovich & Farrar 2020</small>
            |2=''[[Botrychium watertonense|B. ×watertonense]]'' {{au|Wagner}} (Waterton grapefern)<ref>{{PLANTS|id=BOWA2|taxon=Botrychium ×watertonense|access-date=27 Dec 2011}}</ref>
             }}
           }}
        |2={{clade
          |1=''[[Botrychium dusenii|B. dusenii]]'' <small>(Christ 1906) Alston 1960</small>
          |2={{Clade
            |1=''[[Botrychium tunux|B. tunux]]'' {{au|Stensvold & Farrar 2002}}<ref>{{PLANTS|id=BOTU3|taxon=Botrychium tunux|access-date=27 Dec 2011}}</ref><ref>[http://www.tropicos.org/Name/50270442 ''B. tunux''] Tropicos.org. Missouri Botanical Garden. 26 Dec 2011</ref> (moosewort)<ref>[http://www.public.iastate.edu/~herbarium/botrychium/B-tunux.pdf ''B. tunux''] Iowa State Herbarium 03 Jan 2012</ref>
            |2={{Clade
              |1=''[[Botrychium lunaria|B. lunaria]]'' {{au|([[Carl Linnaeus|L.]]) [[Olof Swartz|Sw.]] 1801}} (common moonwort,<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002876 ''B. lunaria''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> grapefern moonwort)
              |2={{Clade
                |1=''[[Botrychium neolunaria|B. neolunaria]]'' {{au|sp. nov. ined. Stensvold & Farrar 2008}} (common moonwort)<ref>[http://www.public.iastate.edu/~herbarium/botrychium/B-neolunaria.pdf ''B. neolunaria''] Iowa State Herbarium 27 Dec 2011</ref>
                |2={{Clade
                  |1=''[[Botrychium nordicum|B. nordicum]]'' <small>Stensvold & Farrar 2016</small>
                  |2={{Clade
                    |1=''[[Botrychium crenulatum|B. crenulatum]]'' {{au|W.H.Wagner 1981}} (dainty<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500275 ''B. crenulatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>/crenulate moonwort)
                    |2=''[[Botrychium yaaxudakeit|B. yaaxudakeit]]'' {{au|Stensvold & Farrar 2002}}<ref>{{PLANTS|id=BOYA|taxon=Botrychium yaaxudakeit|access-date=27 Dec 2011}}</ref><ref>[http://www.tropicos.org/Name/50270445 ''B. yaaxudakeit''] Tropicos.org. Missouri Botanical Garden. 26 Dec 2011</ref> (giant/Yakutat moonwort)<ref>[http://www.public.iastate.edu/~herbarium/botrychium/B-yaaxudakeit.pdf ''B. yaaxudakeit''] Iowa State Herbarium 27 Dec 2011</ref>
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
      |2={{Clade
        |1=''[[Botrychium minganense|B. minganense]]'' {{au|[[Marie-Victorin|Vict.]] 1927}} (Mingan's moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500283 ''B. minganense''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
        |2={{Clade
          |1={{Clade
            |1=''[[Botrychium campestre|B. campestre]]'' {{au|Wagner & Farrar}} (prairie moonwort/<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500274 ''B. campestre''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> dunewort, Iowa moonwort)<ref>[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</ref>
            |2={{Clade
              |1=''[[Botrychium gallicomontanum|B. gallicomontanum]]'' {{au|Farrar & Johnson-Groh 1991}}<ref name="Johnson-Groh2002" /> (Frenchman's Bluff moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500277 ''B. gallicomontanum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
              |2=''[[Botrychium pallidum|B. pallidum]]'' {{au|Wagner 1990}} (Pale moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500288 ''B. pallidum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
               }}
             }}
          |2={{Clade
            |1={{Clade
              |1=''[[Botrychium ascendens|B. ascendens]]'' {{au|W.H.Wagner 1986}} (upswept<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500270 ''B. ascendens''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>/triangle-lobed/upward-lobed moonwort)
              |2=''[[Botrychium simplex|B. simplex]]'' {{au|E.Hitchc. 1823}} (least moonwort/grapefern,<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=200002879 ''B. simplex''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> little grapefern)
               }}
            |2={{Clade
              |1=''[[Botrychium lineare|B. lineare]]'' {{au|Wagner 1994}} (skinny moonwort,<ref>[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</ref> narrowleaf grapefern)<ref>{{PLANTS|id=BOLI7|taxon=Botrychium lineare|access-date=26 Dec 2011}}</ref>
              |2={{Clade
                |1=''[[Botrychium paradoxum|B. paradoxum]]'' {{au|Wagner 1981}} (paradox<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500289 ''B. paradoxum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>/peculiar moonwort)
                |2={{Clade
                  |1=''[[Botrychium spathulatum|B. spathulatum]]'' {{au|Wagner 1990}} (spatulate/<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500295 ''B. spathulatum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> spoon-leaved moonwort)
                  |2={{Clade
                    |1=''[[Botrychium montanum|B. montanum]]'' {{au|[[Warren H. Wagner|Wagner]]}} (western goblin,<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500284 ''B. montanum''] Flora of North America, www.eFloras.org 26 Dec 2011</ref> mountain moonwort)
                    |2=''[[Botrychium mormo|B. mormo]]'' {{au|Wagner 1981}}<ref name="Johnson-Groh2002" /> (little goblin moonwort)<ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=233500285 ''B. mormo''] Flora of North America, www.eFloras.org 26 Dec 2011</ref>
                     }}
                   }}
                 }}
               }}
             }}
           }}
         }}
       }}
     }}
   }}
}}

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

==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.<ref name="Johnson-Groh2002">{{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 }}</ref>  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.<ref>{{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 }}</ref>  Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root.<ref name="Johnson-Groh2002" />

Juvenile and dormant [[sporophyte]]s 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 (fungus)|''Glomus'']], which supply most fixed carbon for growth and reproduction.<ref>{{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 }}</ref>

This mycorrhizal dependence has also made lab cultivation of moonworts difficult.  Thus far, only germination of the gametophyte has been successful.
<!-- 
The only British species is the ''Botrychium lunaria'' ('moonwort'), a little plant pretty frequent in dry mountain pastures.  About a dozen species occur in the United States, ''Botrychium lunaria'' ranges from [[Colorado]] to [[New England]], [[Lake Superior]], and sparingly northward.  ''Botrychium simplex'', a rare species, with small fronds, is found from [[California]] and [[Yellowstone National Park|Yellowstone Park]] to Lake Superior and eastward.  ''Botrychium lanceolatum'' ranges from Colorado to Lake Superior, [[Ohio]], [[New Jersey]], and New England.  ''Botrychium lunarioides'' is found from [[Massachusetts]] to [[Florida]] in dry, rich woods and shady pastures.  ''Botrychium ternatum'' occurs throughout [[North America]].
[this is pretty useless information, as it deals with Botrychium sensu lato, and only from a USUK perspective.]
-->

==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:Botrychium| ]]
[[Category:Fern genera]]
[[Category:Ferns of the Americas]]
[[Category:Ferns of the United States]]
[[Category:Taxa named by Olof Swartz]]