Botrychium

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Moonwort
Botrychium-4.jpg
Botrychium lunaria
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Ophioglossales
Family: Ophioglossaceae
Subfamily: Botrychioideae
Genus: Botrychium
Sw.
Type species
Botrychium lunaria
(L.) Swartz
Species

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.

Contents


Taxonomy

The Smith et al. classification of 2006, based on molecular phylogeny, placed Botrychium in Ophioglossaceae. [2] Subsequent classifications have maintained this placement. [3] [4] [5] Circumscriptions of the genus vary. The Christenhusz and Chase classification of 2014 recognizes a broadly circumscribed Botrychium. [4] This circumscription corresponds to subfamily Botrychioideae in the PPG I classification, which recognizes Botrypus , Japanobotrychum , and Sceptridium as separate genera. [5]

Phylogeny

Phylogeny of Botrychium [6] [7]

section

B. matricariifolium (Döll) Braun ex Koch 1847 [8] (daisy-leaved/ [9] matricary/chamomile moonwort/grapefern)

B. pedunculosum Wagner 1986 (stalked moonwort) [10]

B. hesperium Maxon & Clausen) Wagner & Lellinger 1981 (western moonwort) [11]

B. pseudopinnatum Wagner 1990 (false northwestern/ [12] daisy-leaved grapefern)

B. acuminatum W.H.Wagner 1990 (pointed moonwort) [13]

B. pinnatum H. St.John (northern moonwort [14] )

B. alaskense Wagner & Grant 2002 [15] (Alaska moonwort) [16]

B. boreale Milde 1857 (northern moonwort) [17]

B. lanceolatum (Gmel) Ångstr. 1854 (triangle moonwort, [18] lance-leaved grapefern)

Lanceolatae
section

B. echo Wagner 1983 [19] (reflected grapfern) [20]

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

B. pumicola Coville ex Underw. 1900 [21] (pumice moonwort/grape-fern) [22]

B. furculatum Popovich & Farrar 2020

B. ×watertonense Wagner (Waterton grapefern) [23]

B. dusenii (Christ 1906) Alston 1960

B. tunux Stensvold & Farrar 2002 [24] [25] (moosewort) [26]

B. lunaria (L.) Sw. 1801 (common moonwort, [27] grapefern moonwort)

B. neolunaria sp. nov. ined. Stensvold & Farrar 2008 (common moonwort) [28]

B. nordicum Stensvold & Farrar 2016

B. crenulatum W.H.Wagner 1981 (dainty [29] /crenulate moonwort)

B. yaaxudakeit Stensvold & Farrar 2002 [30] [31] (giant/Yakutat moonwort) [32]

B. minganense Vict. 1927 (Mingan's moonwort) [33]

B. campestre Wagner & Farrar (prairie moonwort/ [34] dunewort, Iowa moonwort) [35]

B. gallicomontanum Farrar & Johnson-Groh 1991 [36] (Frenchman's Bluff moonwort) [37]

B. pallidum Wagner 1990 (Pale moonwort) [38]

B. ascendens W.H.Wagner 1986 (upswept [39] /triangle-lobed/upward-lobed moonwort)

B. simplex E.Hitchc. 1823 (least moonwort/grapefern, [40] little grapefern)

B. lineare Wagner 1994 (skinny moonwort, [41] narrowleaf grapefern) [42]

B. paradoxum Wagner 1981 (paradox [43] /peculiar moonwort)

B. spathulatum Wagner 1990 (spatulate/ [44] spoon-leaved moonwort)

B. montanum Wagner (western goblin, [45] mountain moonwort)

B. mormo Wagner 1981 [36] (little goblin moonwort) [46]

Botrychium

Unassigned species:

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. [36] 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. [50] Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root. [36]

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. [51]

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. ↑ Smith, Alan R.; Pryer, Kathleen M.; Schuettpelz, Eric; Korall, Petra; Schneider, Harald; Wolf, Paul G. (2006). "A classification for extant ferns" (PDF). Taxon . 55 (3): 705–731. doi:10.2307/25065646. JSTOR   25065646.
  3. ↑ 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.
  4. 1 2 Christenhusz, Maarten J.M.; Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany . 113 (9): 571–594. doi:10.1093/aob/mct299. PMC   3936591 . PMID   24532607.
  5. 1 2 PPG I (2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution . 54 (6): 563–603. doi: 10.1111/jse.12229 . S2CID   39980610.
  6. ↑ 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.
  7. ↑ "Tree viewer: interactive visualization of FTOL". FTOL v1.7.0 (GenBank release 261). 2024. Retrieved 1 May 2025.
  8. ↑ 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.
  9. ↑ B. matricariifolium Flora of North America, www.eFloras.org 26 Dec 2011
  10. ↑ B. pedunculosum Flora of North America, www.eFloras.org 26 Dec 2011
  11. ↑ B. hesperium Flora of North America, www.eFloras.org 26 Dec 2011
  12. ↑ B. pseudopinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  13. ↑ B. acuminatum Flora of North America, www.eFloras.org 26 Dec 2011
  14. ↑ B. pinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  15. ↑ B. alaskense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  16. ↑ B. alaskense Iowa State Herbarium 27 Dec 2011
  17. ↑ B. boreale Flora of North America, www.eFloras.org 26 Dec 2011
  18. ↑ B. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
  19. ↑ B. echo Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  20. ↑ B. echo USDA Forest Service, Rocky Mountain Region, Species Conservation Project July 22, 2004
  21. ↑ B. pumicola Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  22. ↑ B. pumicola Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012
  23. ↑ NRCS. "Botrychium ×watertonense". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 27 Dec 2011.
  24. ↑ NRCS. "Botrychium tunux". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 27 Dec 2011.
  25. ↑ B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  26. ↑ B. tunux Iowa State Herbarium 03 Jan 2012
  27. ↑ B. lunaria Flora of North America, www.eFloras.org 26 Dec 2011
  28. ↑ B. neolunaria Iowa State Herbarium 27 Dec 2011
  29. ↑ B. crenulatum Flora of North America, www.eFloras.org 26 Dec 2011
  30. ↑ NRCS. "Botrychium yaaxudakeit". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 27 Dec 2011.
  31. ↑ B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  32. ↑ B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
  33. ↑ B. minganense Flora of North America, www.eFloras.org 26 Dec 2011
  34. ↑ B. campestre Flora of North America, www.eFloras.org 26 Dec 2011
  35. ↑ B. campestre Archived 2013-03-17 at the Wayback Machine Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011
  36. 1 2 3 4 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.
  37. ↑ B. gallicomontanum Flora of North America, www.eFloras.org 26 Dec 2011
  38. ↑ B. pallidum Flora of North America, www.eFloras.org 26 Dec 2011
  39. ↑ B. ascendens Flora of North America, www.eFloras.org 26 Dec 2011
  40. ↑ B. simplex Flora of North America, www.eFloras.org 26 Dec 2011
  41. ↑ B. lineare Archived April 26, 2012, at the Wayback Machine Washington State Department of Natural Resources 26-Dec-2011
  42. ↑ NRCS. "Botrychium lineare". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 26 Dec 2011.
  43. ↑ B. paradoxum Flora of North America, www.eFloras.org 26 Dec 2011
  44. ↑ B. spathulatum Flora of North America, www.eFloras.org 26 Dec 2011
  45. ↑ B. montanum Flora of North America, www.eFloras.org 26 Dec 2011
  46. ↑ B. mormo Flora of North America, www.eFloras.org 26 Dec 2011
  47. ↑ B. daucifolium Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  48. ↑ B. daucifolium Taiwan Plant Names, www.eFlora.org 27 Dec 2011
  49. ↑ B. socorrense Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  50. ↑ 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.
  51. ↑ 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.
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