Biology:Botrychium
| Moonwort | |
|---|---|
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| Botrychium lunaria | |
Scientific classification 
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| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Division: | Polypodiophyta |
| Class: | Polypodiopsida |
| Order: | Ophioglossales |
| Family: | Ophioglossaceae |
| Subfamily: | Botrychioideae |
| Genus: | Botrychium Sw. |
| Type species | |
| Botrychium lunaria (L.) Swartz
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| Species | |
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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.
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.
Taxonomy
| Phylogeny of Botrychium[2][3] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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- Template:Txil – pointed moonwort[4]
- Template:Txil[5] – Alaska moonwort[6]
- Template:Txil – upswept moonwort,[7] triangle-lobed moonwort, upward-lobed moonwort
- Template:Txil – northern moonwort[8]
- Template:Txil – prairie moonwort,[9] prairie dunewort, Iowa moonwort[10]
- Template:Txil – dainty moonwort,[11] crenulate moonwort
- Template:Txil[12] – thin-leaved moonwort[13]
- Botrychium dusenii (Christ 1906) Alston 1960
- Template:Txil[14] – reflected grapfern[15]
- Botrychium furculatum Popovich & Farrar 2020
- Template:Txil[16] – Frenchman's Bluff moonwort[17]
- Template:Txil – western moonwort[18]
- Template:Txil – triangle moonwort,[19] triangle grapefern, lance-leaved grapefern
- Botrychium lineare W.H.Wagner 1994 – skinny moonwort,[20] narrowleaf grapefern[21]
- Botrychium lunaria (L.) Sw. 1801 – common moonwort,[22] grapefern moonwort
- Botrychium matricariifolium (Döll) A. Braun ex W.D.J. Koch 1847[23] – daisy-leaved moonwort,[24] matricary grapefern, matricary moonwort, chamomile grapefern
- Botrychium michiganense Wagner ex Gilman, Farrar & Zika 2015 (Michigan moonwort)
- Botrychium minganense Vict. 1927 – Mingan's moonwort[25]
- Botrychium montanum W.H.Wagner – western goblin,[26] mountain moonwort
- Botrychium mormo W.H.Wagner 1981[16] – little goblin moonwort[27]
- Botrychium neolunaria sp. nov. ined. Stensvold & Farrar 2008 – common moonwort[28]
- Botrychium onondagense
- Botrychium pallidum W.H.Wagner 1990 – Pale moonwort[29]
- Botrychium paradoxum W.H.Wagner 1981 – paradox moonwort,[30] peculiar moonwort
- Botrychium pedunculosum W.H.Wagner 1986 – stalked moonwort[31]
- Botrychium pinnatum H. St.John – northern moonwort[32]
- Botrychium pseudopinnatum W.H.Wagner 1990 – false northwestern moonwort,[33] false daisy-leaved grapefern
- Botrychium pumicola Coville ex Underw. 1900[34] – pumice moonwort, pumice grape-fern[35]
- Botrychium simplex E.Hitchc. 1823 – least moonwort,[36] little grapefern, least grapefern
- Botrychium socorrense W.H.Wagner 1989[37] – Isla Socorro moonwort
- Botrychium spathulatum W.H.Wagner 1990 – spatulate moonwort,[38] spoon-leaved moonwort
- Botrychium sutchuanense Chien & Chun 1959
- Botrychium tolucaense Wagner & Mickel 2004
- Botrychium tunux Stensvold & Farrar 2002[39][40] – moosewort[41]
- Botrychium × watertonense W.H.Wagner [hesperium × paradoxum] – Waterton grapefern[42]
- Botrychium yaaxudakeit Stensvold & Farrar 2002[43][44] – giant moonwort, Yakutat moonwort[45]
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.[46] 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.[47]
This mycorrhizal dependence has also made lab cultivation of moonworts difficult. Thus far, only germination of the gametophyte has been successful.
References
- ↑ Christenhusz, Maarten J. M.; Zhang, Xian-Chun; Schneider, Harald (2011). "A linear sequence of extant families and genera of lycophytes and ferns". Phytotaxa 19: 7–54. doi:10.11646/phytotaxa.19.1.2. http://www.mapress.com/phytotaxa/content/2011/f/pt00019p054.pdf.
- ↑ Nitta, Joel H.Expression error: Unrecognized word "et". (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science 13: 909768. doi:10.3389/fpls.2022.909768. PMID 36092417.
- ↑ "Tree viewer: interactive visualization of FTOL". 2023. https://fernphy.github.io/viewer.html.
- ↑ 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. 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
- ↑ 16.0 16.1 16.2 16.3 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.
- ↑ 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. lanceolatum Flora of North America, www.eFloras.org 26 Dec 2011
- ↑ B. lineare Washington State Department of Natural Resources 26-Dec-2011
- ↑ "Botrychium lineare". Natural Resources Conservation Service PLANTS Database. USDA. https://plants.usda.gov/core/profile?symbol=BOLI7. Retrieved 26 Dec 2011.
- ↑ B. lunaria 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 Oecologica 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
- ↑ "Botrychium tunux". Natural Resources Conservation Service PLANTS Database. USDA. https://plants.usda.gov/core/profile?symbol=BOTU3. Retrieved 27 Dec 2011.
- ↑ B. tunux Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ↑ B. tunux Iowa State Herbarium 03 Jan 2012
- ↑ "Botrychium ×watertonense". Natural Resources Conservation Service PLANTS Database. USDA. https://plants.usda.gov/core/profile?symbol=BOWA2. Retrieved 27 Dec 2011.
- ↑ "Botrychium yaaxudakeit". Natural Resources Conservation Service PLANTS Database. USDA. https://plants.usda.gov/core/profile?symbol=BOYA. Retrieved 27 Dec 2011.
- ↑ B. yaaxudakeit Tropicos.org. Missouri Botanical Garden. 26 Dec 2011
- ↑ B. yaaxudakeit Iowa State Herbarium 27 Dec 2011
- ↑ Whittier, D (1973). "The effect of light and other factors on spore germination in Botrychium dissectum". Can J Bot 51 (10): 1791–1794. doi:10.1139/b73-230.
- ↑ 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.
External links
- USDA Plants Profile for Botrychium (grapefern)
- Efloras.org: Flora of North America, treatment of genus Botrychium
- ITIS.gov: List of Botrychium species — with species links.
Wikidata ☰ Q1353804 entry
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