Botrychium

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Moonwort
Botrychium lunaria
Scientific classification
Kingdom: Plantae
Division: Pteridophyta
Class: Psilotopsida
Order: Ophioglossales
Family: Ophioglossaceae[1][2]
Genus: Botrychium
Sw.
Species

Several, see text

Moonworts are ferns, seedless vascular plants, of the genus Botrychium, sensu stricto. 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 lunarioides in beech/oak forest in northern Florida; March 2003.

Botrychium s.s., the moonworts

  • Botrychium acuminatum W.H.Wagner 1990 – pointed moonwort[3]
  • Botrychium alaskense W.H.Wagner & J.R.Grant 2002[4] – Alaska moonwort[5]
  • Botrychium chamaeconium Bitter & Hieron. 1900[8]
  • Botrychium campestre W.H.Wagner & Farrar – prairie moonwort,[9] prairie dunewort, Iowa moonwort[10]
  • Botrychium daucifolium Wall. ex Hook. & Grev. 1830[12] – thin-leaved moonwort[13]
  • Botrychium echo W.H.Wagner 1983[14] – reflected grapfern[15]
  • 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 japonicum (Prantl) Underw. 1898[19]
  • Botrychium lanceolatum (Gmel) Ångstr. 1854 – triangle moonwort,[20] triangle grapefern, lance-leaved grapefern
  • Botrychium lineare W.H.Wagner 1994 – skinny moonwort,[21] narrowleaf grapefern[22]
  • Botrychium lunarioides (Michx.) Sw. 1806[24] – winter grapefern, prostrate grapefern[25]
  • Botrychium mormo W.H.Wagner 1981[16] – little goblin moonwort[30]
  • Botrychium neolunaria sp. nov. ined. Stensvold & Farrar 2008 – common moonwort[31]
  • Botrychium pallidum W.H.Wagner 1990 – Pale moonwort[32]
  • Botrychium pedunculosum W.H.Wagner 1986 – stalked moonwort[34]
  • Botrychium pseudopinnatum W.H.Wagner 1990 – false northwestern moonwort,[36] false daisy-leaved grapefern
  • Botrychium simplex E.Hitchc. 1823 – least moonwort,[39] little grapefern, least grapefern
  • Botrychium socorrense W.H.Wagner 1989[40] – Isla Socorro moonwort
  • Botrychium spathulatum W.H.Wagner 1990 – spatulate moonwort,[41] spoon-leaved moonwort
  • Botrychium tunux Stensvold & Farrar 2002[42][43] – moosewort[44]
  • Botrychium × watertonense W.H.Wagner [hesperium × paradoxum] – Waterton grapefern[45]
  • Botrychium yaaxudakeit Stensvold & Farrar 2002[46][47] – giant moonwort, Yakutat moonwort[48]
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[49] 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."[53]

  • Botrychium australe R.Br. 1810[54] – Parsley Fern[55]
= Sceptridium australe (R.Br.) Lyon 1905[56]
  • Botrychium biforme Colenso 1886[57] – Fine-leaved parsley fern
= Sceptridium biforme (Colenso) Lyon 1905[58]
  • Botrychium biternatum (Savigny) Underw. 1896 – Sparse-lobed grapefern, Southern grapefern[59]
= Osmunda biternata Savigny 1798
= Sceptridium biternatum (Savigny) Lyon 1905
  • Botrychium dissectum Sprengel 1804[60][61] – Dissected grapefern[62] or Cut-leaved grape-fern[63]
= Botrychium obliquum Muhl. 1810[64]
= Sceptridium dissectum (Spreng.) Lyon 1905
  • Botrychium jenmanii Underw. 1900 – Alabama grapefern[65][66]
= Sceptridium jenmanii (Underw.) Lyon 1905
= Botrychium alabamense Maxon 1906[67]
= Sceptridium alabamense (Maxon) Holub. 1973
= Osmunda multifida S.G. Gmel. 1768
= Botrychium silaifolium C.Presl 1825
= Botrychium matricariae (Schrank) Spreng. 1827[73]
= Sceptridium multifidum (S.G.Gmel.) Nishida ex Tagawa 1958
  • Botrychium oneidense (Gilbert) House 1905 – Blunt-lobed grapefern[74]
= Sceptridium oneidense (Gilbert) Holub 1998[75]
  • Botrychium rugulosum W.H.Wagner 1982[76] – St. Lawrence grape fern, Rugulose grape fern, Ternate grape fern[77]
= Botrychium ternatum auct. non (Thunb.) Sw. 1801
= Sceptridium rugulosum (W.H.Wagner) Skoda & Holub 1996
  • Botrychium schaffneri Underw. 1903[78] –
  • Botrychium subbifoliatum Brack. 1854[79] – Island grapefern[80][81]
    = Sceptridium subbifoliatum (Brack.) Lyon 1905
  • Botrychium underwoodianum Maxon 1905[82]
= Botrychium ternatum (Thunb.) Sw. 1801
= Sceptridium underwoodianum (Maxon) Lyon 1905[83]

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.[84] 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.[85]

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


References

  1. ↑ Botrychium Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
  2. ↑ 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. 
  3. ↑ B. acuminatum Flora of North America, www.eFloras.org 26 Dec 2011
  4. ↑ B. alaskense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  5. ↑ B. alaskense Iowa State Herbarium 27 Dec 2011
  6. ↑ B. ascendens Flora of North America, www.eFloras.org 26 Dec 2011
  7. ↑ B. boreale Flora of North America, www.eFloras.org 26 Dec 2011
  8. ↑ B. chamaeconium Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
  9. ↑ B. campestre Flora of North America, www.eFloras.org 26 Dec 2011
  10. ↑ B. campestre Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011
  11. ↑ B. crenulatum Flora of North America, www.eFloras.org 26 Dec 2011
  12. ↑ B. daucifolium Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  13. ↑ B. daucifolium Taiwan Plant Names, www.eFlora.org 27 Dec 2011
  14. ↑ B. echo Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  15. ↑ B. echo USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004
  16. ↑ 16.0 16.1 16.2 16.3 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.
  17. ↑ B. gallicomontanum Flora of North America, www.eFloras.org 26 Dec 2011
  18. ↑ B. hesperium Flora of North America, www.eFloras.org 26 Dec 2011
  19. ↑ B. japonicum Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
  20. ↑ B. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
  21. ↑ B. lineare Washington State Department of Natural Resources 26-Dec-2011
  22. ↑ B. lineare, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011 
  23. ↑ B. lunaria Flora of North America, www.eFloras.org 26 Dec 2011
  24. ↑ Wagner, W. H. (1993). Flora of North America. New York: Oxford University Press. pp. 85–91. 
  25. ↑ B. lunarioides Flora of North America, www.eFloras.org 26 Dec 2011
  26. ↑ 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 Oecologia 13: 335–343. 
  27. ↑ B. matricariifolium Flora of North America, www.eFloras.org 26 Dec 2011
  28. ↑ B. minganense Flora of North America, www.eFloras.org 26 Dec 2011
  29. ↑ B. montanum Flora of North America, www.eFloras.org 26 Dec 2011
  30. ↑ B. mormo Flora of North America, www.eFloras.org 26 Dec 2011
  31. ↑ B. neolunaria Iowa State Herbarium 27 Dec 2011
  32. ↑ B. pallidum Flora of North America, www.eFloras.org 26 Dec 2011
  33. ↑ B. paradoxum Flora of North America, www.eFloras.org 26 Dec 2011
  34. ↑ B. pedunculosum Flora of North America, www.eFloras.org 26 Dec 2011
  35. ↑ B. pinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  36. ↑ B. pseudopinnatum Flora of North America, www.eFloras.org 26 Dec 2011
  37. ↑ B. pumicola Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
  38. ↑ B. pumicola Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012
  39. ↑ B. simplex Flora of North America, www.eFloras.org 26 Dec 2011
  40. ↑ B. socorrense Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  41. ↑ B. spathulatum Flora of North America, www.eFloras.org 26 Dec 2011
  42. ↑ B. tunux, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011 
  43. ↑ B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  44. ↑ B. tunux Iowa State Herbarium 03 Jan 2012
  45. ↑ B. ×watertonense, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011 
  46. ↑ B. yaaxudakeit, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011 
  47. ↑ B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  48. ↑ B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
  49. ↑ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 247 (2005)
  50. ↑ B. virginianum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  51. ↑ B. virginianum, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011 
  52. ↑ B. virginianum Flora of North America, www.eFloras.org 26 Dec 2011
  53. ↑ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 265 (2005)
  54. ↑ Kelly, Dave (1994). "Demography and conservation of Botrychium australe, a peculiar, sparse mycorrhizal fern". New Zealand Journal of Botany 32: 393–400. 
  55. ↑ B. australe Plants for a Future 26 Dec 2011
  56. ↑ International Plant Names Index (IPNI) 17 Jan 2011
  57. ↑ B. biforme International Plant Names Index (IPNI) 02 Jan 2012
  58. ↑ S. biforme International Plant Names Index (IPNI) 02 Jan 2012
  59. ↑ B. biternatum Flora of North America, www.eFloras.org 26 Dec 2011
  60. ↑ Montgomery, James D. (1990). "Survivorship and Predation Changes in Five Populations of Botrychium dissectum in Eastern Pennsylvania". American Fern Journal. 4 80: 173–182. 
  61. ↑ B. dissectum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  62. ↑ B. dissectum Flora of North America, www.eFloras.org 26 Dec 2011
  63. ↑ MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. January 16, 2012
  64. ↑ B. obliquum Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
  65. ↑ B. jenmanii Flora of North America, www.eFloras.org 26 Dec 2011
  66. ↑ B. jenmanii Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  67. ↑ B. alabamense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  68. ↑ Mesipuu, Meeli; R. P. Shefferson and T. Kull (2009). "Weather and herbivores influence fertility in the endangered fern Botrychium multifidum (S.G. Gmel.) Rupr". Plant Ecology 203: 23–31. 
  69. ↑ B. multifidum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  70. ↑ B. multifidum Flora of North America, www.eFloras.org 26 Dec 2011
  71. ↑ B. multifidum, USDA Natural Resources Conservation Service PLANTS Profile 
  72. ↑ B. multifidum University of Wisconsin, Cofrin Center for Biodiversity, Ferns and Fern Allies of Wisconsin. 26 Dec 2011
  73. ↑ Botrychium Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 02 Jan 2012
  74. ↑ B. oneidense Flora of North America, www.eFloras.org 26 Dec 2011
  75. ↑ Botrychium oneidense Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
  76. ↑ B. rugulosum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
  77. ↑ B. rugulosum Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 27 Dec 2011
  78. ↑ B. schaffneri Tropicos.org. Missouri Botanical Garden. 04 Jan 2012
  79. ↑ B. subbifoliatum International Plant Names Index 27 Dec 2011
  80. ↑ B. subbifoliatum ITIS taxonomy 26 Dec 2011
  81. ↑ S. subbifoliatum, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011 
  82. ↑ B. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  83. ↑ S. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
  84. ↑ Whittier D (1973). “The effect of light and other factors on spore germination in Botrychium dissectum”. Can J Bot 51: 1791-1794.
  85. ↑ Winther J, Friedman W (2007). “Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae)”. Am J Bot 94 (7): 1248-1255.
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