Botrychium

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

Phylogeny of Botrychium^[2][3]

section B. matricariifolium (Döll) Braun ex Koch 1847[4] (daisy-leaved/[5] matricary/chamomile moonwort/grapefern) B. hesperium Maxon & Clausen) Wagner & Lellinger 1981 (western moonwort)[6] B. pseudopinnatum Wagner 1990 (false northwestern/[7] daisy-leaved grapefern) B. pedunculosum Wagner 1986 (stalked moonwort)[8] B. acuminatum W.H.Wagner 1990 (pointed moonwort)[9] B. pinnatum H. St.John (northern moonwort[10]) B. alaskense Wagner & Grant 2002[11] (Alaska moonwort)[12] B. boreale Milde 1857 (northern moonwort)[13] B. lanceolatum (Gmel) Ångstr. 1854 (triangle moonwort,[14] lance-leaved grapefern) Lanceolatae section B. echo Wagner 1983[15] (reflected grapfern)[16] B. michiganense Wagner ex Gilman, Farrar & Zika 2015 (Michigan moonwort) B. pumicola Coville ex Underw. 1900[17] (pumice moonwort/grape-fern)[18] B. furculatum Popovich & Farrar 2020 B. ×watertonense Wagner (Waterton grapefern)[19] B. dusenii (Christ 1906) Alston 1960 B. tunux Stensvold & Farrar 2002[20][21] (moosewort)[22] B. lunaria (L.) Sw. 1801 (common moonwort,[23] grapefern moonwort) B. neolunaria sp. nov. ined. Stensvold & Farrar 2008 (common moonwort)[24] B. nordicum Stensvold & Farrar 2016 B. crenulatum W.H.Wagner 1981 (dainty[25]/crenulate moonwort) B. yaaxudakeit Stensvold & Farrar 2002[26][27] (giant/Yakutat moonwort)[28] B. minganense Vict. 1927 (Mingan's moonwort)[29] B. campestre Wagner & Farrar (prairie moonwort/[30] dunewort, Iowa moonwort)[31] B. mormo Wagner 1981[32] (little goblin moonwort)[33] B. lineare Wagner 1994 (skinny moonwort,[34] narrowleaf grapefern)[35] B. ascendens W.H.Wagner 1986 (upswept[36]/triangle-lobed/upward-lobed moonwort) B. simplex E.Hitchc. 1823 (least moonwort/grapefern,[37] little grapefern) B. paradoxum Wagner 1981 (paradox[38]/peculiar moonwort) B. spathulatum Wagner 1990 (spatulate/[39] spoon-leaved moonwort) B. montanum Wagner (western goblin,[40] mountain moonwort) B. gallicomontanum Farrar & Johnson-Groh 1991[32] (Frenchman's Bluff moonwort)[41] B. pallidum Wagner 1990 (Pale moonwort)[42] Botrychium

Unassigned species:

• Botrychium daucifolium Wall. ex Hook. & Grev. 1830^[43] (thin-leaved moonwort)^[44]
• Botrychium onondagense
• Botrychium socorrense W.H.Wagner 1989^[45] (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.^[32] 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.^[32]

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

1. Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19: 7–54. doi:10.11646/phytotaxa.19.1.2.
2. Nitta, Joel H.; Schuettpelz, Eric; Ramírez-Barahona, Santiago; Iwasaki, Wataru; et al. (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science. 13: 909768. doi:10.3389/fpls.2022.909768. PMC 9449725. PMID 36092417.
3. "Tree viewer: interactive visualization of FTOL". FTOL v1.6.0 [GenBank release 259]. 2023. Retrieved 27 July 2024.
4. Muller, S. (1992). "The impact of a drought in spring on the sporulation of Botrychium matricariifolium (Retz) A. Br. in the Bitcherland (Northern Vosges, France)". Acta Oecologica. 13: 335–343.
5. B. matricariifolium Flora of North America, www.eFloras.org 26 Dec 2011
6. B. hesperium Flora of North America, www.eFloras.org 26 Dec 2011
7. B. pseudopinnatum Flora of North America, www.eFloras.org 26 Dec 2011
8. B. pedunculosum Flora of North America, www.eFloras.org 26 Dec 2011
9. B. acuminatum Flora of North America, www.eFloras.org 26 Dec 2011
10. B. pinnatum Flora of North America, www.eFloras.org 26 Dec 2011
11. B. alaskense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
12. B. alaskense Iowa State Herbarium 27 Dec 2011
13. B. boreale Flora of North America, www.eFloras.org 26 Dec 2011
14. B. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
15. B. echo Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
16. B. echo USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004
17. B. pumicola Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
18. B. pumicola Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012
19. USDA, NRCS (n.d.). "​Botrychium ×watertonense​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 27 Dec 2011.
20. USDA, NRCS (n.d.). "​Botrychium tunux​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 27 Dec 2011.
21. B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
22. B. tunux Iowa State Herbarium 03 Jan 2012
23. B. lunaria Flora of North America, www.eFloras.org 26 Dec 2011
24. B. neolunaria Iowa State Herbarium 27 Dec 2011
25. B. crenulatum Flora of North America, www.eFloras.org 26 Dec 2011
26. USDA, NRCS (n.d.). "​Botrychium yaaxudakeit​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 27 Dec 2011.
27. B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
28. B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
29. B. minganense Flora of North America, www.eFloras.org 26 Dec 2011
30. B. campestre Flora of North America, www.eFloras.org 26 Dec 2011
31. B. campestre Archived 2013-03-17 at the Wayback Machine Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011
32. Johnson-Groh, C. L.; Lee, J. (2002). "Phenology and demography of two species of Botrychium (Ophioglassaceae)". American Journal of Botany. 89 (10): 1624–1633. doi:10.3732/ajb.89.10.1624. PMID 21665590.
33. B. mormo Flora of North America, www.eFloras.org 26 Dec 2011
34. B. lineare Archived April 26, 2012, at the Wayback Machine Washington State Department of Natural Resources 26-Dec-2011
35. USDA, NRCS (n.d.). "​Botrychium lineare​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 26 Dec 2011.
36. B. ascendens Flora of North America, www.eFloras.org 26 Dec 2011
37. B. simplex Flora of North America, www.eFloras.org 26 Dec 2011
38. B. paradoxum Flora of North America, www.eFloras.org 26 Dec 2011
39. B. spathulatum Flora of North America, www.eFloras.org 26 Dec 2011
40. B. montanum Flora of North America, www.eFloras.org 26 Dec 2011
41. B. gallicomontanum Flora of North America, www.eFloras.org 26 Dec 2011
42. B. pallidum Flora of North America, www.eFloras.org 26 Dec 2011
43. B. daucifolium Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
44. B. daucifolium Taiwan Plant Names, www.eFlora.org 27 Dec 2011
45. B. socorrense Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
46. Whittier, D (1973). "The effect of light and other factors on spore germination in Botrychium dissectum". Can J Bot. 51 (10): 1791–1794. Bibcode:1973CaJB...51.1791W. doi:10.1139/b73-230.
47. Winther, J; Friedman, W (2007). "Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae)". Am J Bot. 94 (7): 1248–1255. doi:10.3732/ajb.94.7.1248. PMID 21636490.

External links

• 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.