Botrychium
| Moonwort | |
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| Botrychium lunaria | |
| Scientific classification | |
| Kingdom: | |
| Division: | |
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| Genus: | Botrychium |
| Species | |
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Several, see text | |
Botrychium is a genus of ferns, seedless vascular plants. 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 acuminatum W.H.Wagner 1990 – pointed moonwort[3]
- Botrychium alaskense W.H.Wagner & J.R.Grant 2002[4] – Alaska moonwort[5]
- Botrychium ascendens W.H.Wagner 1986 – upswept moonwort,[6] triangle-lobed moonwort, upward-lobed moonwort
- Botrychium boreale J.Milde 1857 – northern moonwort[7]
- Botrychium chamaeconium Bitter & Hieron. 1900[8]
- Botrychium campestre W.H.Wagner & Farrar – prairie moonwort,[9] prairie dunewort, Iowa moonwort[10]
- Botrychium crenulatum W.H.Wagner 1981 – dainty moonwort,[11] crenulate moonwort
- 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 lunaria (L.) Sw. 1801 – common moonwort,[23] grapefern moonwort
- Botrychium lunarioides (Michx.) Sw. 1806[24] – winter grapefern, prostrate grapefern[25]
- Botrychium matricariifolium (Döll) A. Braun ex W.D.J. Koch 1847[26] – daisy-leaved moonwort,[27] matricary grapefern, matricary moonwort, chamomile grapefern
- Botrychium minganense Vict. 1927 – Mingan's moonwort[28]
- Botrychium montanum W.H.Wagner – western goblin,[29] mountain moonwort
- 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 paradoxum W.H.Wagner 1981 – paradox moonwort,[33] peculiar moonwort
- Botrychium pedunculosum W.H.Wagner 1986 – stalked moonwort[34]
- Botrychium pinnatum H. St.John – northern moonwort[35]
- Botrychium pseudopinnatum W.H.Wagner 1990 – false northwestern moonwort,[36] false daisy-leaved grapefern
- Botrychium pumicola Coville ex Underw. 1900[37] – pumice moonwort, pumice grape-fern[38]
- 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 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 virginianum (L.) Sw. 1801[50] – Rattlesnake fern,[51] common grapefern[52]
= Botrypus virginianus (L.) Michx. 1803
Botrychium species placed in Sceptridium
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
- ^ Botrychium Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
- ^ 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.
- ^ B. acuminatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. alaskense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
- ^ B. alaskense Iowa State Herbarium 27 Dec 2011
- ^ B. ascendens Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. boreale Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. chamaeconium Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
- ^ B. campestre Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. campestre Robert W. Freckmann Herbarium University of Wisconsin - Stevens Point 27 Dec 2011
- ^ B. crenulatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. daucifolium Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
- ^ B. daucifolium Taiwan Plant Names, www.eFlora.org 27 Dec 2011
- ^ B. echo Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
- ^ B. echo USDA Forest Service,Rocky Mountain Region, Species Conservation Project July 22, 2004
- ^ a b c d 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.
- ^ B. gallicomontanum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. hesperium Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. japonicum Tropicos.org. Missouri Botanical Garden. 02 Jan 2012
- ^ B. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. lineare Washington State Department of Natural Resources 26-Dec-2011
- ^ B. lineare, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011
- ^ B. lunaria Flora of North America, www.eFloras.org 26 Dec 2011
- ^ Wagner, W. H. (1993). Flora of North America. New York: Oxford University Press. pp. 85–91.
- ^ B. lunarioides Flora of North America, www.eFloras.org 26 Dec 2011
- ^ 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.
- ^ B. matricariifolium Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. minganense Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. montanum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. mormo Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. neolunaria Iowa State Herbarium 27 Dec 2011
- ^ B. pallidum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. paradoxum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. pedunculosum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. pinnatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. pseudopinnatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. pumicola Tropicos.org. Missouri Botanical Garden. 03 Jan 2012
- ^ B. pumicola Oregon.gov ODA Plant Division, Plant Conservation 03 Dec 2012
- ^ B. simplex Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. socorrense Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. spathulatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. tunux, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
- ^ B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. tunux Iowa State Herbarium 03 Jan 2012
- ^ B. ×watertonense, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
- ^ B. yaaxudakeit, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
- ^ B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
- ^ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 247 (2005)
- ^ B. virginianum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. virginianum, USDA Natural Resources Conservation Service PLANTS Profile, 27 Dec 2011
- ^ B. virginianum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ Cobb, Farsworth & Lowe, Ferns of Northeastern North America 2nd edition, p. 265 (2005)
- ^ 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.
- ^ B. australe Plants for a Future 26 Dec 2011
- ^ International Plant Names Index (IPNI) 17 Jan 2011
- ^ B. biforme International Plant Names Index (IPNI) 02 Jan 2012
- ^ S. biforme International Plant Names Index (IPNI) 02 Jan 2012
- ^ B. biternatum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ 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.
- ^ B. dissectum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. dissectum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ [1] MICHIGAN FLORA ONLINE. A. A. Reznicek, E. G. Voss, & B. S. Walters. February 2011. University of Michigan. Web. January 16, 2012
- ^ B. obliquum Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
- ^ B. jenmanii Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. jenmanii Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. alabamense Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
- ^ 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.
- ^ B. multifidum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. multifidum Flora of North America, www.eFloras.org 26 Dec 2011
- ^ B. multifidum, USDA Natural Resources Conservation Service PLANTS Profile
- ^ B. multifidum University of Wisconsin, Cofrin Center for Biodiversity, Ferns and Fern Allies of Wisconsin. 26 Dec 2011
- ^ Botrychium Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 02 Jan 2012
- ^ B. oneidense Flora of North America, www.eFloras.org 26 Dec 2011
- ^ Botrychium oneidense Tropicos.org. Missouri Botanical Garden. 16 Jan 2012
- ^ B. rugulosum Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ^ B. rugulosum Robert W. Freckmann Herbarium, University of Wisconsin - Stevens Point 27 Dec 2011
- ^ B. schaffneri Tropicos.org. Missouri Botanical Garden. 04 Jan 2012
- ^ B. subbifoliatum International Plant Names Index 27 Dec 2011
- ^ B. subbifoliatum ITIS taxonomy 26 Dec 2011
- ^ S. subbifoliatum, USDA Natural Resources Conservation Service PLANTS Profile, 26 Dec 2011
- ^ B. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
- ^ S. underwoodianum Tropicos.org. Missouri Botanical Garden. 27 Dec 2011
- ^ Whittier D (1973). “The effect of light and other factors on spore germination in Botrychium dissectum”. Can J Bot 51: 1791-1794.
- ^ Winther J, Friedman W (2007). “Arbuscular mycorrhizal symbionts in Botrychium (Ophioglossaceae)”. Am J Bot 94 (7): 1248-1255.
Wikimedia Commons has media related to Botrychium.
- Botrychium in Flora of North America (incl. Botrypus)