''Botrychium'' is a genus of

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except ...

s, seedless

vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...

s in the family Ophioglossaceae. ''Botrychium'' species are known as moonworts. They are small, with fleshy

root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...

s, and reproduce by

spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...

s 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 A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...

. The circumscription of ''Botrychium'' is disputed between different authors; some botanists include the genera ''

Botrypus ''Botrypus virginianus'', synonym ''Botrychium virginianum'', sometimes called rattlesnake fern is a species of perennial fern in the adders-tongue family. It is monotypic within the genus ''Botrypus'', meaning that it is the only species withi ...

'' and ''

Sceptridium ''Sceptridium'' is a genus of seedless vascular plants in the family Ophioglossaceae, closely allied to (and often included as a subgenus of) the genus '' Botrychium'' (the moonworts and grapeferns). It is also closely related to the genus '' B ...

'' within ''Botrychium'', while others treat them as distinct. The latter treatment is provisionally followed here.

Taxonomy

* – pointed moonwort * – Alaska moonwort * – upswept moonwort, triangle-lobed moonwort, upward-lobed moonwort * – northern moonwort * – prairie moonwort, prairie dunewort, Iowa moonwort * – dainty moonwort, crenulate moonwort * – thin-leaved moonwort * ''

Botrychium dusenii ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' (Christ 1906) Alston 1960 * – reflected grapfern * ''

Botrychium furculatum ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' Popovich & Farrar 2020 * – Frenchman's Bluff moonwort * – western moonwort * – triangle moonwort, triangle grapefern, lance-leaved grapefern * '' Botrychium lineare'' – skinny moonwort, narrowleaf grapefern * ''

Botrychium lunaria ''Botrychium lunaria'' is a species of fern in the family Ophioglossaceae known by the common name moonwort or common moonwort. It is the most widely distributed moonwort, growing throughout the Northern Hemisphere across Eurasia and from Alas ...

'' – common moonwort, grapefern moonwort * ''

Botrychium matricariifolium ''Botrychium matricariifolium'' ( orth.var. ''B. matricariaefolium'') is a species of fern in the Ophioglossaceae family. It is referred to by the common names chamomile grape-fern, daisyleaf grape-fern, and matricary grape-fern. It is native to ...

'' – daisy-leaved moonwort, matricary grapefern, matricary moonwort, chamomile grapefern * ''

Botrychium michiganense ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' Wagner ex Gilman, Farrar & Zika 2015 (Michigan moonwort) * ''

Botrychium minganense ''Botrychium minganense'' is a species of fern in the family Ophioglossaceae known by the common name Mingan moonwort. It is native to North America from Alaska and northern Canada to Arizona, where it is uncommon throughout most of its range, ap ...

'' – Mingan's moonwort * '' Botrychium montanum'' – western goblin, mountain moonwort * ''

Botrychium mormo ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' – little goblin moonwort * ''

Botrychium neolunaria ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' – common moonwort * ''

Botrychium pallidum ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 tr ...

'' – Pale moonwort * ''

Botrychium paradoxum ''Botrychium paradoxum'' is a species of fern in the family Ophioglossaceae known by the common name peculiar moonwort. It is native to North America, where there are scattered occurrences in Alberta, British Columbia, Saskatchewan, Montana, Oreg ...

'' – paradox moonwort, peculiar moonwort * ''

Botrychium pedunculosum ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 tr ...

'' – stalked moonwort * '' Botrychium pinnatum'' – northern moonwort * '' Botrychium pseudopinnatum'' – false northwestern moonwort, false daisy-leaved grapefern * '' Botrychium pumicola'' – pumice moonwort, pumice grape-fern * '' Botrychium simplex'' – least moonwort, little grapefern, least grapefern * '' Botrychium socorrense'' – Isla Socorro moonwort * ''

Botrychium spathulatum ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 tr ...

'' – spatulate moonwort, spoon-leaved moonwort * '' Botrychium sutchuanense'' Chien & Chun 1959 * '' Botrychium tolucaense'' Wagner & Mickel 2004 * ''

Botrychium tunux ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' – moosewort * ''

Botrychium × watertonense ''Botrychium'' is a genus of ferns, seedless vascular plants in the family Ophioglossaceae. ''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 ...

'' – Waterton grapefern * '' Botrychium yaaxudakeit'' – giant moonwort, Yakutat moonwort

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. 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. Finally, some species produce gemmae, a form of asexual propagation achieved by budding of the root. 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. This mycorrhizal dependence has also made lab cultivation of moonworts difficult. Thus far, only germination of the gametophyte has been successful.

References

External links

USDA Plants Profile for ''Botrychium'' (grapeferns)

Efloras.org: Flora of North America, treatment of genus ''Botrychium'' ITIS.gov: List of ''Botrychium'' species
— ''with species links''. {{Taxonbar, from=Q1353804 Fern genera Ferns of the Americas Ferns of the United States Taxa named by Olof Swartz