Botrychium

Moonwort
Botrychium lunaria
Scientific classification
Kingdom: Plantae
Division: Pteridophyta
Class: Psilotopsida
Order: Ophioglossales
Family: Ophioglossaceae
Subfamily: Botrychioideae
Genus: Botrychium
Sw.
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. They are unusual among tracheophytes ("higher plants") in that at least some species produce the sugar trehalose.

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.

Selected species

Botrychium s.s., the moonworts

Botrychium virginianum

Botrychium species placed in Botrypus

The rattlesnake fern has traditionally been placed in the Botrychium subgenus Osmundopteris, the name of which is based on the species' superficial similarities to the Osmunda genus[48] where it was previously placed.

Botrychium species placed in Sceptridium

Botrychium rugulosum

These species (the evergreen grapeferns) have traditionally been placed in the Botrychium subgenus Sceptridium,a name based on the apparent similarity of their sporangia to "little scepters."[52]

= Sceptridium australe (R.Br.) Lyon 1905[55]
= Sceptridium biforme (Colenso) Lyon 1905[57]
= Osmunda biternata Savigny 1798
= Sceptridium biternatum (Savigny) Lyon 1905
= Botrychium obliquum Muhl. 1810[63]
= Sceptridium dissectum (Spreng.) Lyon 1905
= Sceptridium jenmanii (Underw.) Lyon 1905
= Botrychium alabamense Maxon 1906[66]
= Sceptridium alabamense (Maxon) Holub. 1973
= Osmunda multifida S.G. Gmel. 1768
= Botrychium silaifolium C.Presl 1825
= Botrychium matricariae (Schrank) Spreng. 1827[72]
= Sceptridium multifidum (S.G.Gmel.) Nishida ex Tagawa 1958
= Sceptridium oneidense (Gilbert) Holub 1998[74]
= Botrychium ternatum auct. non (Thunb.) Sw. 1801
= Sceptridium rugulosum (W.H.Wagner) Skoda & Holub 1996
= Botrychium ternatum (Thunb.) Sw. 1801
= Sceptridium underwoodianum (Maxon) Lyon 1905[82]

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.[15] 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.[83] Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root.[15]

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.[84]

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.
  2. ↑ B. acuminatum Flora of North America, www.eFloras.org 26 Dec 2011
  3. ↑ B. alaskense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  4. ↑ B. alaskense Iowa State Herbarium 27 Dec 2011
  5. ↑ B. ascendens Flora of North America, www.eFloras.org 26 Dec 2011
  6. ↑ B. boreale Flora of North America, www.eFloras.org 26 Dec 2011
  7. ↑ B. chamaeconium Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
  8. ↑ B. campestre Flora of North America, www.eFloras.org 26 Dec 2011
  9. ↑ B. campestre Archived 2013-03-17 at the Wayback Machine. Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011
  10. ↑ B. crenulatum Flora of North America, www.eFloras.org 26 Dec 2011
  11. ↑ B. daucifolium Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  12. ↑ B. daucifolium Taiwan Plant Names, www.eFlora.org 27 Dec 2011
  13. ↑ B. echo Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  14. ↑ B. echo USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004
  15. 1 2 3 4 Johnson-Groh, C. L. and J. Lee (2002). “Phenology and demography of two species of Botrychium (Ophioglassaceae)”. American Journal of Botany 89 (10): 1624-1633.
  16. ↑ B. gallicomontanum Flora of North America, www.eFloras.org 26 Dec 2011
  17. ↑ B. hesperium Flora of North America, www.eFloras.org 26 Dec 2011
  18. ↑ B. japonicum Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
  19. ↑ B. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
  20. ↑ B. lineare Archived April 26, 2012, at the Wayback Machine. Washington State Department of Natural Resources 26-Dec-2011
  21. ↑ B. lineare, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011
  22. ↑ B. lunaria Flora of North America, www.eFloras.org 26 Dec 2011
  23. ↑ Wagner, W. H. (1993). Flora of North America. New York: Oxford University Press. pp. 85–91.
  24. ↑ B. lunarioides Flora of North America, www.eFloras.org 26 Dec 2011
  25. ↑ 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.
  26. ↑ B. matricariifolium Flora of North America, www.eFloras.org 26 Dec 2011
  27. ↑ B. minganense Flora of North America, www.eFloras.org 26 Dec 2011
  28. ↑ B. montanum Flora of North America, www.eFloras.org 26 Dec 2011
  29. ↑ B. mormo Flora of North America, www.eFloras.org 26 Dec 2011
  30. ↑ B. neolunaria Iowa State Herbarium 27 Dec 2011
  31. ↑ B. pallidum Flora of North America, www.eFloras.org 26 Dec 2011
  32. ↑ B. paradoxum Flora of North America, www.eFloras.org 26 Dec 2011
  33. ↑ B. pedunculosum Flora of North America, www.eFloras.org 26 Dec 2011
  34. ↑ B. pinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  35. ↑ B. pseudopinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  36. ↑ B. pumicola Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  37. ↑ B. pumicola Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012
  38. ↑ B. simplex Flora of North America, www.eFloras.org 26 Dec 2011
  39. ↑ B. socorrense Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  40. ↑ B. spathulatum Flora of North America, www.eFloras.org 26 Dec 2011
  41. ↑ B. tunux, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
  42. ↑ B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  43. ↑ B. tunux Iowa State Herbarium 03 Jan 2012
  44. ↑ B. ×watertonense, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
  45. ↑ B. yaaxudakeit, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
  46. ↑ B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  47. ↑ B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
  48. ↑ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 247 (2005)
  49. ↑ B. virginianum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  50. ↑ B. virginianum, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
  51. ↑ B. virginianum Flora of North America, www.eFloras.org 26 Dec 2011
  52. ↑ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 265 (2005)
  53. ↑ Kelly, Dave (1994). "Demography and conservation of Botrychium australe, a peculiar, sparse mycorrhizal fern". New Zealand Journal of Botany. 32: 393–400. doi:10.1080/0028825x.1994.10412925.
  54. ↑ B. australe Plants for a Future 26 Dec 2011
  55. ↑ International Plant Names Index (IPNI) 17 Jan 2011
  56. ↑ B. biforme International Plant Names Index (IPNI) 02 Jan 2012
  57. ↑ S. biforme International Plant Names Index (IPNI) 02 Jan 2012
  58. ↑ B. biternatum Flora of North America, www.eFloras.org 26 Dec 2011
  59. ↑ Montgomery, James D. (1990). "Survivorship and Predation Changes in Five Populations of Botrychium dissectum in Eastern Pennsylvania". American Fern Journal. 4. 80: 173–182. doi:10.2307/1547206.
  60. ↑ B. dissectum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  61. ↑ B. dissectum Flora of North America, www.eFloras.org 26 Dec 2011
  62. ↑ MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. January 16, 2012
  63. ↑ B. obliquum Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
  64. ↑ B. jenmanii Flora of North America, www.eFloras.org 26 Dec 2011
  65. ↑ B. jenmanii Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  66. ↑ B. alabamense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  67. ↑ Mesipuu, Meeli; R. P. Shefferson; T. Kull (2009). "Weather and herbivores influence fertility in the endangered fern Botrychium multifidum (S.G. Gmel.) Rupr". Plant Ecology. 203: 23–31. doi:10.1007/s11258-008-9501-3.
  68. ↑ B. multifidum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  69. ↑ B. multifidum Flora of North America, www.eFloras.org 26 Dec 2011
  70. ↑ B. multifidum, USDA Natural Resources Conservation Service PLANTS Profile
  71. ↑ B. multifidum University of Wisconsin, Cofrin Center for Biodiversity, Ferns and Fern Allies of Wisconsin. 26 Dec 2011
  72. ↑ Botrychium Archived 2013-03-17 at the Wayback Machine. Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 02 Jan 2012
  73. ↑ B. oneidense Flora of North America, www.eFloras.org 26 Dec 2011
  74. ↑ Botrychium oneidense Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
  75. ↑ B. rugulosum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  76. ↑ B. rugulosum Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 27 Dec 2011
  77. ↑ B. schaffneri Tropicos.org. Missouri Botanical Garden. 04 Jan 2012
  78. ↑ B. subbifoliatum International Plant Names Index 27 Dec 2011
  79. ↑ B. subbifoliatum ITIS taxonomy 26 Dec 2011
  80. ↑ S. subbifoliatum, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011
  81. ↑ B. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  82. ↑ S. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  83. ↑ Whittier D (1973). “The effect of light and other factors on spore germination in Botrychium dissectum”. Can J Bot 51: 1791-1794.
  84. ↑ Winther J, Friedman W (2007). “Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae)”. Am J Bot 94 (7): 1248-1255.
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