Moonwort
Botrychium lunaria
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Division:	Polypodiophyta
Class:	Polypodiopsida
Order:	Ophioglossales
Family:	Ophioglossaceae
Subfamily:	Botrychioideae
Genus:	Botrychium Sw.
Type species
Botrychium lunaria (L.) Swartz
Species
Several, see text

Botrychium is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ^[1] 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.

Taxonomy

• Botrychium acuminatum W.H.Wagner 1990 – pointed moonwort ^[4]
• Botrychium alaskense W.H.Wagner & J.R.Grant 2002 ^[5] – Alaska moonwort ^[6]
• Botrychium ascendens W.H.Wagner 1986 – upswept moonwort, ^[7] triangle-lobed moonwort, upward-lobed moonwort
• Botrychium boreale J.Milde 1857 – northern moonwort ^[8]
• Botrychium campestre W.H.Wagner & Farrar – prairie moonwort, ^[9] prairie dunewort, Iowa moonwort ^[10]
• Botrychium crenulatum W.H.Wagner 1981 – dainty moonwort, ^[11] crenulate moonwort
• Botrychium daucifolium Wall. ex Hook. & Grev. 1830 ^[12] – thin-leaved moonwort ^[13]
• Botrychium dusenii (Christ 1906) Alston 1960
• Botrychium echo W.H.Wagner 1983 ^[14] – reflected grapfern ^[15]
• Botrychium furculatum Popovich & Farrar 2020
• Botrychium gallicomontanum Farrar & Johnson-Groh 1991 ^[16] – Frenchman's Bluff moonwort ^[17]
• Botrychium hesperium Maxon & R.T.Clausen) W.H.Wagner & Lellinger 1981 – western moonwort ^[18]
• Botrychium lanceolatum (Gmel) Ångstr. 1854 – triangle moonwort, ^[19] triangle grapefern, lance-leaved grapefern
• Botrychium lineare W.H.Wagner 1994 – skinny moonwort, ^[20] narrowleaf grapefern ^[21]
• Botrychium lunaria ( L.) Sw. 1801 – common moonwort, ^[22] grapefern moonwort
• Botrychium matricariifolium (Döll) A. Braun ex W.D.J. Koch 1847 ^[23] – daisy-leaved moonwort, ^[24] matricary grapefern, matricary moonwort, chamomile grapefern
• Botrychium michiganense Wagner ex Gilman, Farrar & Zika 2015 (Michigan moonwort)
• Botrychium minganense Vict. 1927 – Mingan's moonwort ^[25]
• Botrychium montanum W.H.Wagner – western goblin, ^[26] mountain moonwort
• Botrychium mormo W.H.Wagner 1981 ^[16] – little goblin moonwort ^[27]
• Botrychium neolunaria sp. nov. ined. Stensvold & Farrar 2008 – common moonwort ^[28]
• Botrychium onondagense
• Botrychium pallidum W.H.Wagner 1990 – Pale moonwort ^[29]
• Botrychium paradoxum W.H.Wagner 1981 – paradox moonwort, ^[30] peculiar moonwort
• Botrychium pedunculosum W.H.Wagner 1986 – stalked moonwort ^[31]
• Botrychium pinnatum H. St.John – northern moonwort ^[32]
• Botrychium pseudopinnatum W.H.Wagner 1990 – false northwestern moonwort, ^[33] false daisy-leaved grapefern
• Botrychium pumicola Coville ex Underw. 1900 ^[34] – pumice moonwort, pumice grape-fern ^[35]
• Botrychium simplex E.Hitchc. 1823 – least moonwort, ^[36] little grapefern, least grapefern
• Botrychium socorrense W.H.Wagner 1989 ^[37] – Isla Socorro moonwort
• Botrychium spathulatum W.H.Wagner 1990 – spatulate moonwort, ^[38] spoon-leaved moonwort
• Botrychium sutchuanense Chien & Chun 1959
• Botrychium tolucaense Wagner & Mickel 2004
• Botrychium tunux Stensvold & Farrar 2002 ^{[39] [40]} – moosewort ^[41]
• Botrychium × watertonense W.H.Wagner [hesperium × paradoxum] – Waterton grapefern ^[42]
• Botrychium yaaxudakeit Stensvold & Farrar 2002 ^{[43] [44]} – giant moonwort, Yakutat moonwort ^[45]

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. ^[16] 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. ^[46] Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root. ^[16]

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. ^[47]

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

References

External links

Wikimedia Commons has media related to Botrychium.

• USDA Plants Profile for Botrychium (grapefern)
• Efloras.org: Flora of North America, treatment of genus Botrychium
• ITIS.gov: List of Botrychium species — with species links.