Botrychium

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Moonwort
Botrychium-4.jpg
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[edit]

Botrychium lunarioides in beech/oak forest in northern Florida; March 2003.

Botrychium s.s., the moonworts[edit]

Botrychium virginianum

Botrychium species placed in Botrypus[edit]

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[edit]

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]

= Sceptridium australe (R.Br.) Lyon 1905[56]
= Sceptridium biforme (Colenso) Lyon 1905[58]
= Osmunda biternata Savigny 1798
= Sceptridium biternatum (Savigny) Lyon 1905
= Botrychium obliquum Muhl. 1810[64]
= Sceptridium dissectum (Spreng.) Lyon 1905
= 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
= Sceptridium oneidense (Gilbert) Holub 1998[75]
= 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[83]

Conservation[edit]

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[edit]

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  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
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  5. ^ B. alaskense Iowa State Herbarium 27 Dec 2011
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  15. ^ B. echo USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004
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  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. 
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  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 
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  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
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  49. ^ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 247 (2005)
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  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. doi:10.1080/0028825x.1994.10412925. 
  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
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  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. doi:10.2307/1547206. 
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  62. ^ B. dissectum Flora of North America, www.eFloras.org 26 Dec 2011
  63. ^ [1] 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; 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. 
  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 
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  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.