Equisetum

Equisetum (/ˌɛkwɪˈsiːtəm/ ; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds.^[2]

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 30 m (98 ft) tall.^[3] 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.^[4] 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.^[5]

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 equus ('horse') + seta ('bristle').^[6]

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.^[7] 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.^[8] In German, the corresponding name is Zinnkraut ('tin-herb') or Schachtelhalm ('nesting stalk'). In Spanish-speaking countries, these plants are known as cola de caballo ('horsetail').

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.^[9]

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.^[10] Unusually, the branches often emerge below the leaves in an internode, and grow from buds between their bases.

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.^{[11] : 12–15}

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.^[12] 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.^[13]

The crude cell extracts of all Equisetum species tested contain mixed-linkage glucan : xyloglucan endotransglucosylase (MXE) activity.^[14] 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.^{[15] [16]} 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.^[17] 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.^{[citation needed ]}

Currently, 18 species of Equisetum are accepted by Plants of the World Online.^[1] 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.^[18]

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.^{[19] [citation needed ]}

The Smith et al. classification of 2006, based on molecular phylogeny, placed Equisetum in Equisetaceae.^[20] Subsequent classifications have maintained this placement.^{[21] [22] [23]}

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^[24] and Equisetum laterale from the Early-Middle Jurassic of Australia.^{[25] [26]} Silicified remains of Equisetum thermale from the Late Jurassic of Argentina exhibit all the morphological characters of modern members of the genus.^[27] The estimated split between Equisetum bogotense and all other living Equisetum is estimated to have occurred no later than the Early Jurassic.^[26]

• Equisetum bogotense Kunth – 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)^[28] transfer this here, as E. bogotense appears to be sister to all the remaining species in the genus.

• Equisetum arvense L. – field horsetail or common horsetail; circumboreal down through temperate zones
• Equisetum braunii Milde – northern giant horsetail, syn. E. telmateia subsp. braunii (Milde) Hauke.; west coast of North America
• Equisetum diffusum D.Don – Himalayan horsetail; Himalayan India and China and adjacent nations above about 450 metres (1,480 ft)
• Equisetum fluviatile L. – water horsetail; circumboreal down through temperate zones
• Equisetum palustre L. – marsh horsetail; circumboreal down through temperate zones
• Equisetum pratense Ehrh. – shady horsetail, meadow horsetail, shade horsetail; circumboreal except for tundra down through cool temperate zones
• Equisetum sylvaticum L. – wood horsetail; circumboreal down through cool temperate zones, more restricted in east Asia
• Equisetum telmateia Ehrh. – 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 Equisetum braunii Milde^{[28] [1]}

• Equisetum giganteum L. – southern giant horsetail or giant horsetail; temperate to tropical South America and Central America north to southern Mexico
• Equisetum hyemale L. – 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 Equisetum praealtum Raf.^{[28] [1]}
• Equisetum laevigatum A.Braun – smooth horsetail, smooth scouringrush; western 3/4 of North America down into northwestern Mexico; also sometimes known as Equisetum kansanum
• Equisetum myriochaetum Schltdl. & Cham. – Mexican giant horsetail; from central Mexico south to Peru
• Equisetum praealtum Raf. – scouringrush horsetail, syn. E. hyemale subsp. affine (Engelm.) A.A.Eat.; temperate North America
• Equisetum ramosissimum Desf. (including E. debile ) – branched horsetail; Asia, Europe, Africa, southwest Pacific islands
• Equisetum scirpoides Michx. – dwarf horsetail, dwarf scouringrush; northern (cool temperate) zones worldwide
• Equisetum variegatum Schleich. ex Weber & Mohr – variegated horsetail, variegated scouringrush; northern (cool temperate) zones worldwide, except for northeasternmost Asia
• Equisetum xylochaetum Mett. – Atacama Desert giant horsetail; southern Peru, northern Chile

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

• ×ばつ bowmanii (page does not exist)">Equisetum ×ばつ bowmanii C.N.Page (Equisetum sylvaticum ×ばつ Equisetum telmateia)
• ×ばつ dycei (page does not exist)">Equisetum ×ばつ dycei C.N.Page (Equisetum fluviatile ×ばつ Equisetum palustre)
• ×ばつ font-queri (page does not exist)">Equisetum ×ばつ font-queri Rothm. (Equisetum palustre ×ばつ Equisetum telmateia)
• ×ばつ litorale (page does not exist)">Equisetum ×ばつ litorale Kühlew ex Rupr. (Equisetum arvense ×ばつ Equisetum fluviatile)
• ×ばつ mchaffieae (page does not exist)">Equisetum ×ばつ mchaffieae C.N.Page (Equisetum fluviatile ×ばつ Equisetum pratense)
• ×ばつ mildeanum (page does not exist)">Equisetum ×ばつ mildeanum Rothm. (Equisetum pratense ×ばつ Equisetum sylvaticum)
• ×ばつ robertsii (page does not exist)">Equisetum ×ばつ robertsii Dines (Equisetum arvense ×ばつ Equisetum telmateia)
• ×ばつ rothmaleri (page does not exist)">Equisetum ×ばつ rothmaleri C.N.Page (Equisetum arvense ×ばつ Equisetum palustre)
• ×ばつ willmotii (page does not exist)">Equisetum ×ばつ willmotii C.N.Page (Equisetum fluviatile ×ばつ Equisetum telmateia)

• ×ばつ ferrissii (page does not exist)">Equisetum ×ばつ ferrissii Clute (Equisetum hyemale ×ばつ Equisetum laevigatum)
• ×ばつ moorei">Equisetum ×ばつ moorei Newman (Equisetum hyemale ×ばつ Equisetum ramosissimum)
• ×ばつ nelsonii (page does not exist)">Equisetum ×ばつ nelsonii (A.A.Eaton) Schaffn. (Equisetum laevigatum ×ばつ Equisetum variegatum)
• ×ばつ schaffneri">Equisetum ×ばつ schaffneri Milde (Equisetum giganteum ×ばつ Equisetum myriochaetum)
• ×ばつ trachyodon">Equisetum ×ばつ trachyodon (A.Braun) W.D.J.Koch (Equisetum hyemale ×ばつ Equisetum variegatum)