equisetum

''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds.

''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which for over 100 million years was much more diverse and dominated the understorey of late Paleozoic forests. Some equisetids were large trees reaching to tall. The genus ''Calamites'' of the family Calamitaceae, for example, is abundant in coal deposits from the Carboniferous period. The pattern of spacing of nodes in horsetails, wherein those toward the apex of the shoot are increasingly close together, is said to have inspired John Napier to invent logarithms. Modern horsetails first appeared during the Jurassic period.

A superficially similar but entirely unrelated flowering plant genus, mare's tail (''Hippuris''), is occasionally referred to as "horsetail", and adding to confusion, the name "mare's tail" is sometimes applied to ''Equisetum''.

Etymology

The name "horsetail", often used for the entire group, arose because the branched species somewhat resemble a horse's tail. Similarly, the scientific name ''Equisetum'' is derived from the Latin ('horse') + ('bristle').

Other names include candock for branching species, puzzlegrass, and snake grass or scouring-rush for unbranched or sparsely branched species. The latter name refers to the rush-like appearance of the plants and to the fact that the stems are coated with abrasive silicates, making them useful for scouring (cleaning) metal items such as cooking pots or drinking mugs, particularly those made of tin. '' Equisetum hyemale'', rough horsetail, is still boiled and then dried in Japan to be used for the final polishing process on woodcraft to produce a smooth finish. In German, the corresponding name is ('tin-herb'). In Spanish-speaking countries, these plants are known as ('horsetail').

Description

''Equisetum'' leaves are greatly reduced and usually non-photosynthetic. They contain a single, non-branching vascular trace, which is the defining feature of microphylls. However, it has recently been recognised that horsetail microphylls are probably not ancestral as in lycophytes (clubmosses and relatives), but rather derived adaptations, evolved by reduction of megaphylls.

The leaves of horsetails are arranged in whorls fused into nodal sheaths. The stems are usually green and photosynthetic, and are distinctive in being hollow, jointed and ridged (with sometimes 3 but usually 6–40 ridges). There may or may not be whorls of branches at the nodes.Streeter D, Hart-Davies C, Hardcastle A, Cole F, Harper L. 2009. ''Collins Flower Guide''. Harper Collins Unusually, the branches often emerge below the leaves in an internode, and grow from buds between their bases.

Spores

The spores are borne under sporangiophores in strobili, cone-like structures at the tips of some of the stems. In many species the cone-bearing shoots are unbranched, and in some (e.g. '' E. arvense'', field horsetail) they are non-photosynthetic, produced early in spring. In some other species (e.g. '' E. palustre'', marsh horsetail) they are very similar to sterile shoots, photosynthetic and with whorls of branches.

Horsetails are mostly homosporous, though in the field horsetail, smaller spores give rise to male prothalli. The spores have four elaters that act as moisture-sensitive springs, assisting spore dispersal through crawling and hopping motions after the sporangia have split open longitudinally. They are photosynthetic and have a lifespan that is usually two weeks at most, but will germinate immediately under humid conditions and develop into a gametophyte.

Cell walls

The crude cell extracts of all ''Equisetum'' species tested contain mixed-linkage glucan : xyloglucan endotransglucosylase (MXE) activity. This is a novel enzyme and is not known to occur in any other plants. In addition, the cell walls of all ''Equisetum'' species tested contain mixed-linkage glucan (MLG), a polysaccharide which, until recently, was thought to be confined to the Poales. The evolutionary distance between ''Equisetum'' and the Poales suggests that each evolved MLG independently. The presence of MXE activity in ''Equisetum'' suggests that they have evolved MLG along with some mechanism of cell wall modification. Non-''Equisetum'' land plants tested lack detectable MXE activity. An observed negative correlation between XET activity and cell age led to the suggestion that XET is catalysing endotransglycosylation in controlled wall-loosening during cell expansion. The lack of MXE in the Poales suggests that there it must play some other, currently unknown, role. Due to the correlation between MXE activity and cell age, MXE has been proposed to promote the cessation of cell expansion.

Taxonomy

Species

Currently, 18 species of ''Equisetum'' are accepted by Plants of the World Online. The living members are divided into three distinct lineages, which are usually treated as subgenera. The name of the type subgenus, ''Equisetum'', means "horse hair" in Latin, while the name of the other large subgenus, ''Hippochaete'', means "horse hair" in Greek. Hybrids are common, but hybridization has only been recorded between members of the same subgenus.

Two ''Equisetum'' plants are sold under the names ''Equisetum japonicum'' (barred horsetail) and ''Equisetum camtschatcense'' (Kamchatka horsetail). These are both types of ''E. hyemale'' var. ''hyemale'', although they may also be listed as separate varieties of ''E. hyemale''.

Evolutionary history

The oldest remains of modern horsetails of the genus ''Equisetum'' first appear in the Early Jurassic, represented by '' Equisetum dimorphum'' from the Early Jurassic of Patagonia and '' Equisetum laterale'' from the Early-Middle Jurassic of Australia. Silicified remains of '' Equisetum thermale'' from the Late Jurassic of Argentina exhibit all the morphological characters of modern members of the genus. The estimated split between '' Equisetum bogotense'' and all other living ''Equisetum'' is estimated to have occurred no later than the Early Jurassic.

Subgenus ''Paramochaete''

* – Andean horsetail; upland South America up to Costa Rica; includes ''E. rinihuense'', sometimes treated as a separate species. Previously included in subg. ''Equisetum'', but Christenhusz ''et al.'' (2019) transfer this here, as ''E. bogotense'' appears to be sister to all the remaining species in the genus.

Subgenus ''Equisetum''

* – field horsetail or common horsetail; circumboreal down through temperate zones
* – northern giant horsetail, syn. ''E. telmateia'' subsp. ''braunii'' (Milde) Hauke.; west coast of North America
* – Himalayan horsetail; Himalayan India and China and adjacent nations above about
* – water horsetail; circumboreal down through temperate zones
* – marsh horsetail; circumboreal down through temperate zones
* – shady horsetail, meadow horsetail, shade horsetail; circumboreal except for tundra down through cool temperate zones
* – wood horsetail; circumboreal down through cool temperate zones, more restricted in east Asia
* – great horsetail; Europe to Asia Minor and north Africa. The former North American subspecies ''Equisetum telmateia'' subsp. ''braunii'' (Milde) Hauke is now treated as a separate species

Subgenus ''Hippochaete''

* – southern giant horsetail or giant horsetail; temperate to tropical South America and Central America north to southern Mexico
* – rough horsetail; most of non-tropical Old World. The former North American subspecies ''Equisetum hyemale'' subsp. ''affine'' (Engelm.) A.A.Eat. is now treated as a separate species
* – smooth horsetail, smooth scouringrush; western 3/4 of North America down into northwestern Mexico; also sometimes known as ''Equisetum kansanum''
* – Mexican giant horsetail; from central Mexico south to Peru
* – scouringrush horsetail, syn. ''E. hyemale'' subsp. ''affine'' (Engelm.) A.A.Eat.; temperate North America
* (including '' E. debile'') – branched horsetail; Asia, Europe, Africa, southwest Pacific islands
* – dwarf horsetail, dwarf scouringrush; northern (cool temperate) zones worldwide
* – variegated horsetail, variegated scouringrush; northern (cool temperate) zones worldwide, except for northeasternmost Asia
* – Atacama Desert giant horsetail; southern Peru, northern Chile

Unplaced to subgenus

*'' Equisetum dimorphum'' – Early Jurassic, Argentina
*'' Equisetum laterale –'' Early to Middle Jurassic, Australia
*'' Equisetum thermale'' – Middle to Late Jurassic, Argentina
*'' Equisetum similkamense'' – Ypresian, British Columbia

Named hybrids

Hybrids between species in subgenus ''Equisetum''

* ''Equisetum'' × ''bowmanii'' (''Equisetum sylvaticum'' × ''Equisetum telmateia'')
* ''Equisetum'' × ''dycei'' (''Equisetum fluviatile'' × ''Equisetum palustre'')
* ''Equisetum'' × ''font-queri'' (''Equisetum palustre'' × ''Equisetum telmateia'')
* ''Equisetum'' × ''litorale'' (''Equisetum arvense'' × ''Equisetum fluviatile'')
* ''Equisetum'' × ''mchaffieae'' (''Equisetum fluviatile'' × ''Equisetum pratense'')
* ''Equisetum'' × ''mildeanum'' (''Equisetum pratense'' × ''Equisetum sylvaticum'')
* ''Equisetum'' × ''robertsii'' (''Equisetum arvense'' × ''Equisetum telmateia'')
* ''Equisetum'' × ''rothmaleri'' (''Equisetum arvense'' × ''Equisetum palustre'')
* ''Equisetum'' × ''willmotii'' (''Equisetum fluviatile'' × ''Equisetum telmateia'')

Hybrids between species in subgenus ''Hippochaete''

* ''Equisetum'' × ''ferrissii'' (''Equisetum hyemale'' × ''Equisetum laevigatum'')
* ''Equisetum'' × ''moorei'' (''Equisetum hyemale'' × ''Equisetum ramosissimum'')
* ''Equisetum'' × ''nelsonii'' (''Equisetum laevigatum'' × ''Equisetum variegatum'')
* ''Equisetum'' × ''schaffneri'' (''Equisetum giganteum'' × ''Equisetum myriochaetum'')
* ''Equisetum'' × ''trachyodon'' (''Equisetum hyemale'' × ''Equisetum variegatum'')

Phylogeny

Distribution and ecology

The genus ''Equisetum'' as a whole, while concentrated in the non-tropical northern hemisphere, is near-cosmopolitan, being absent naturally only from Antarctica, Australia, New Zealand, and the islands of the Pacific Ocean. They are most common in northern Europe, with ten species (''E. arvense'', ''E. fluviatile'', ''E. hyemale'', ''E. palustre'', ''E. pratense'', ''E. ramosissimum'', ''E. scirpoides'', ''E. sylvaticum'', ''E. telmateia'', and ''E. variegatum''); Great Britain has nine of these species, missing only ''E. scirpoides'' of the European list. Northern North America (Canada and the northernmost United States), also has nine species (''E. arvense'', ''E. fluviatile'', ''E. laevigatum'', ''E. palustre'', ''E. praealtum'', ''E. pratense'', ''E. scirpoides'', ''E. sylvaticum'', and ''E. variegatum''). Only five (''E. bogotense'', ''E. giganteum'', ''E. myriochaetum'', ''E. ramosissimum'', and ''E. xylochaetum'') of the eighteen species are known to be native south of the Equator.

They are perennial plants, herbaceous and dying back in winter in most temperate species, or evergreen as most tropical species and the temperate species '' E. hyemale'' (rough horsetail), '' E. ramosissimum'' (branched horsetail), '' E. scirpoides'' (dwarf horsetail) and '' E. variegatum'' (variegated horsetail). They typically grow 20 cm–1.5 m (8 in–5 ft) tall, though the subtropical "giant horsetails" are recorded to grow as high as ('' E. giganteum'', southern giant horsetail) or ('' E. myriochaetum'', Mexican giant horsetail), and allegedly even more.

One species, '' Equisetum fluviatile'', is an emergent aquatic, rooted in water with shoots growing into the air. The stalks arise from rhizomes that are deep underground and difficult to dig out. Field horsetail ('' E. arvense'') can be a nuisance weed, readily regrowing from the rhizome after being pulled out. It is unaffected by many herbicides designed to kill seed plants. Since the stems have a waxy coat, the plant is resistant to contact weedkillers like glyphosate. However, as ''E. arvense'' prefers an acid soil, lime may be used to assist in eradication efforts to bring the soil pH to 7 or 8. Members of the genus have been declared noxious weeds in Australia and in the US state of Oregon.

All the ''Equisetum'' are classed as "unwanted organisms" in New Zealand and are listed on the National Pest Plant Accord.

Consumption

People have regularly consumed horsetails. The fertile stems bearing strobili of some species can be cooked and eaten like asparagus (a dish called in Japan). Indigenous nations across Cascadia consume and use horsetails in a variety of ways, with the Squamish calling them ''sx̱ém'x̱em'' and the Lushootseed using ''gʷəɫik'', or horsetail roots, for cedar root baskets. The young plants are eaten cooked or raw, but considerable care must be taken.

If eaten over a long enough period of time, some species of horsetail can be poisonous to grazing animals, including horses. The toxicity appears to be due to thiaminase, which can cause thiamin (vitamin B1) deficiency.

''Equisetum'' species may have been a common food for herbivorous dinosaurs. With studies showing that horsetails are nutritionally of high quality, it is assumed that horsetails were an important component of herbivorous dinosaur diets. Analysis of the scratch marks on hadrosaur teeth is consistent with grazing on hard plants like horsetails.

Folk medicine and safety concerns

Extracts and other preparations of ''E. arvense'' have served as herbal remedies, with records dating over centuries. In 2009, the European Food Safety Authority concluded there was no evidence for the supposed health effects of ''E. arvense'', such as for invigoration, weight control, skincare, hair health or bone health. , there is insufficient scientific evidence for its effectiveness as a medicine to treat any human condition.

''E. arvense'' contains thiaminase, which metabolizes the B vitamin, thiamine, potentially causing thiamine deficiency and associated liver damage, if taken chronically. Horsetail might produce a diuretic effect. Further, its safety for oral consumption has not been sufficiently evaluated and it may be toxic, especially to children and pregnant women.

See also

* List of plants poisonous to equines

References

Further reading

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External links

''Equisetum''
at the Tree of Life Web Project

National Collection of ''Equisetum''

International Equisetological Association
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{{Authority control

Fern genera
Medicinal plants
Extant Middle Jurassic first appearances
Callovian first appearances
Callovian genus first appearances
Taxa named by Carl Linnaeus