Epilobium
Epilobium
| Epilobium | |
|---|---|
| Epilobium hirsutum (great willowherb) | |
| Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Rosids |
| Order: | Myrtales |
| Family: | Onagraceae |
| Subfamily: | Onagroideae |
| Tribe: | Epilobieae |
| Genus: | Epilobium L. |
| Species | |
| 160-200, see text | |
| Synonyms | |
| |
Epilobium is a genus of flowering plants in the family Onagraceae, containing about 197 species.[1] The genus has a worldwide distribution. It is most prevalent in the subarctic, temperate and subantarctic regions, whereas in the subtropics and tropics Epilobium species are restricted to the cool montane biomes, such as the New Guinea Highlands.
The taxonomy of the genus has varied between different botanists, but the modern trend is to include the previously recognised genera Boisduvalia, Pyrogennema and Zauschneria within Epilobium. Chamaenerion, previously (Chamerion), is considered distinct, however,[2] according to Peter H. Raven, who has extensively studied the willowherbs and merges the other segregate genera into Epilobium. Fringed willowherb (E. ciliatum) is likely a cryptic species complex; apparently these plants also commonly hybridize with their congeners.[3]
Most species are known by the common name willowherbs for their willow-like leaves.[4] Those that were once separated in Boisduvalia are called spike-primroses or boisduvalias. Those Epilobium species previously placed in the Chamaenerion group and known as fireweeds are now segregated into the genus Chamaenerion.
| Epilobium | |
|---|---|
| Epilobium hirsutum (great willowherb) | |
| Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Rosids |
| Order: | Myrtales |
| Family: | Onagraceae |
| Subfamily: | Onagroideae |
| Tribe: | Epilobieae |
| Genus: | Epilobium L. |
| Species | |
| 160-200, see text | |
| Synonyms | |
| |
Description
Ripe capsule of Epilobium canum (zauschneria) releasing seed
Epilobiums are mostly herbaceous plants, either annual or perennial; a few are subshrubs. The leaves are opposite or rarely whorled,[5] [] simple and ovate to lanceolate in shape. The flowers are actinomorphic (radially symmetrical) with four petals that may be notched.[5][6] These are usually smallish and pink in most species, but red, orange or yellow in a few. The fruit is a slender cylindrical capsule containing numerous seeds embedded in fine, soft silky fluff which disperses the seeds very effectively in the wind.
A number of Epilobium species with slightly asymmetrical (zygomorphic) magenta flowers and alternate leaves were placed in a Chamaenerion group, and have been separated by some authorities into the genus Chamerion. However, Chamerion has not been universally accepted[7] and there is now emerging consensus that this group of species should be segregated into the genus Chamaenerion.[8][9]
Habitat and ecology
Willowherbs sensu lato are typically very quick to carpet large swathes of ground and may become key or dominant species of local ecosystems. In the United Kingdom, for example, rosebay willowherb (Chamaenerion angustifolium) is widely found on mesotrophic soils dominated by false oat-grass (Arrhenatherum elatius), cock's-foot grass (Dactylis glomerata), and red fescue (Festuca rubra), while great willowherb (Epilobium hirsutum) is found in mesotrophic grassland with stinging nettle (Urtica dioica). These two willowherb species also dominate open habitat early in ecological succession, to the virtual exclusion of other plant life. Broad-leaved willowherb (Epilobium montanum) is found characteristically, though not abundantly, in the mesotrophic grasslands with meadowsweet (Filipendula ulmaria) and sometimes the uncommon Greek valerian (Polemonium caeruleum). Most willowherbs will not tolerate shade trees and thus are limited to more recently disturbed patches, yielding to other plants over time. Consequently, though the genus contains many pioneer plants, rather few of them are invasive weeds of major importance.
Epilobium species are used as food plants by the caterpillars of certain Lepidoptera species, including: Geometridae
Grey pug (Eupithecia subfuscata), recorded on several species
Noctuidae
Mouse moth (Amphipyra tragopoginis), recorded on several species
Small angle shades (Euplexia lucipara), recorded on several species
Australian grapevine moth (Phalaenoides glycinae), recorded on fringed willowherb (E. ciliatum)
Sphingidae
Elephant hawk-moth (Deilephila elpenor), recorded on several species
Small elephant hawk-moth (Deilephila porcellus), recorded on several species
White-lined sphinx (Hyles lineata), recorded on several species
Use by humans
The main use of Epilobium by humans is as a herbal supplement in the treatment of prostate, bladder (incontinence) and hormone disorders. Many of the small willowherb species are nuisance weeds in gardens. Though few are regularly used as ornamental plants, the larger willowherbs may be attractive in ruderal locales. One of the most frequently recognized members of the genus is the circumboreal fireweed (E. angustifolium), known as rosebay willowherb in the United Kingdom. It rapidly colonizes burnt ground; during the bombing of London in World War II many of the derelict bomb sites were soon covered with these plants, bringing a splash of colour to what was otherwise a very grim scene. It is the floral emblem of Yukon in Canada, Hedmark in Norway (where it is called geitrams) and Southern Ostrobothnia in Finland.
Fireweed is used as a sweetener in northwestern North America. It is put in candy, jellies, ice cream, syrup, and sxusem ("Indian ice cream"). In the late summer its flowers yield pollen and copious nectar which give a rich spicy honey. Its young leaves, roots, and shoots are edible (if somewhat bitter), and rich in provitamin A and vitamin C. The Dena’ina found them also useful as food supplement for dogs and applied sap from the stem to wounds, believing it to have antiinflammatory properties.
Several researchers have studied this taxon. Heinrich Carl Haussknecht in the late 19th century and Peter H. Raven about a century later researched the phylogeny, systematics, and taxonomy of willowherbs. Peter Michaelis' studies of this genus paved the way for understanding of extranuclear inheritance in plants.
Species
Epilobium brachycarpum (tall willowherb)
Epilobium canum (zauschneria)
Epilobium alpestre
Top left: Epilobium alsinifolium (chickweed willowherb)Bottom left: Epilobium anagallidifolium (alpine willowherb)Center: Epilobium tetragonum (square-stemmed willowherb)
Epilobium billardierianum
Left: Epilobium roseum (pale willowherb)
Epilobium parviflorum (small-flowered willowherb)
Chamaenerion fleischeri
Boisduvalia
Epilobium densiflorum – denseflower willowherb, dense spike-primrose, dense boisduvalia
Epilobium pallidum – largeflower spike-primrose
Epilobium pygmaeum[10] – pygmy willowherb, smooth boisduvalia
Epilobium subdentatum (Meyen) Lievens & Hoch
Epilobium torreyi – Torrey's willowherb, brook spike-primrose
Cordylophorum
- Subsection Nuttalia
Epilobium suffruticosum Nutt. ex Torr. & A.Gray – shrubby willowherb
- Subsection Petrolobium
Epilobium nevadense Munz – Nevada willowherb
Epilobium nivium Brandegee – Snow Mountain willowherb
Crossostigma
Epilobium foliosum – leafy willowherb, California willowherb
Epilobium minutum – chaparral willowherb, desert willowherb, smallflower willowherb
Epilobiopsis
Epilobium campestre (Jeps.) Hoch & W.L.Wagner
Epilobium cleistogamum – selfing willowherb
Macrocarpa
Epilobium rigidum – stiff willowherb, Siskiyou Mountains willowherb
Xerolobium
Epilobium brachycarpum – tall willowherb, tall annual willowherb, tall fireweed, panicled willowherb
Zauschneria
Epilobium canum – Zauschneria, California-fuchsia, hummingbird flower, hummingbird trumpet
Epilobium septentrionale – northern willowherb, Humboldt County fuchsia
Right: Epilobium obscurum (dwarf willowherb)]]
Epilobium
Epilobium aitchisonii P.H.Raven
Epilobium algidum M.Bieb.
Epilobium alpinum
Epilobium alpestre (Jacq.) Krock.
Epilobium alsinifolium Vill. – chickweed willowherb
Epilobium alsinoides A.Cunn. Epilobium alsinoides ssp. alsinoides Epilobium alsinoides ssp. atriplicifolium (A.Cunn.) P.H.Raven & Engelhorn Epilobium alsinoides ssp. tenuipes (Hook.f.) P.H. Raven & Engelhorn
Epilobium amurense Hausskn. Epilobium amurense ssp. amurense Epilobium amurense ssp. cephalostigma (Hausskn.) C.J.Chen, Hoch & P.H.Raven
Epilobium anagallidifolium – alpine willowherb, pimpernel willowherb
Epilobium anatolicum Hausskn. Epilobium anatolicum ssp. anatolicum Epilobium anatolicum ssp. prionophyllum (Hausskn.) P.H.Raven
Epilobium angustum (Cheeseman) P.H. Raven & Engelhorn
Epilobium arcticum Sam. – Arctic willowherb
Epilobium astonii (Allan) P.H.Raven & Engelhorn
Epilobium atlanticum Litard. & Maire
Epilobium australe Poeppig & Hausskn.
Epilobium barbeyanum H.Lév.
Epilobium billardierianum Ser. – glabrous willowherb Epilobium billardierianum ssp. billardierianum Epilobium billardierianum ssp. cinereum (A.Rich.) P.H.Raven & Engelhorn Epilobium billardierianum ssp. hydrophilum P.H. Raven & Engelhorn Epilobium billardierianum ssp. intermedium P.H. Raven & Engelhorn
Epilobium blinii H.Lév.
Epilobium brevifolium D.Don Epilobium brevifolium ssp. brevifolium Epilobium brevifolium ssp. trichoneurum (Hausskn.) P.H.Raven
Epilobium brevipes Hook.f.
Epilobium brevisquamatum P.H.Raven
Epilobium brunnescens (Cockayne) P.H. Raven & Engelhorn – New Zealand willowherb Epilobium brunnescens ssp. beaugleholei K.R.West & P.H.Raven Epilobium brunnescens ssp. brunnescens Epilobium brunnescens ssp. minutiflorum (Cockayne) P.H. Raven & Engelhorn
Epilobium capense Buchinger ex Hochst.
Epilobium chionanthum Hausskn.
Epilobium chitralense P.H.Raven
Epilobium chlorifolium Hausskn.
Epilobium ciliatum – fringed willowherb, American willowherb
Epilobium clarkeanum Hausskn.
Epilobium clavatum – talus willowherb, clavatefruit willowherb
Epilobium collinum C.C.Gmelin
Epilobium coloratum Biehler – purpleleaf willowherb
Epilobium confertifolium Hook.f.
Epilobium confusum Hausskn.
Epilobium conjungens Skottsb.
Epilobium crassum Hook.f.
Epilobium curtisiae P.H.Raven
Epilobium cylindricum D.Don
Epilobium davuricum Fisch. ex Hornem. – Daurian willowherb
Epilobium densifolium Hausskn.
Epilobium denticulatum Ruiz & Pav.
Epilobium detzneranum Schltr. ex Diels
Epilobium duriaei Gay ex Godr.
Epilobium fangii C.J.Chen, Hoch & P.H.Raven
Epilobium fastigiatoramosum Nakai
Epilobium fauriei H.Lév.
Epilobium forbesii Allan
Epilobium fragile Sam.
Epilobium frigidum Hausskn.
Epilobium fugitivum P.H. Raven & Engelhorn
Epilobium gemmascens C.A.Mey.
Epilobium glabellum G.Forst. – smooth spike-primrose
Epilobium glaberrimum – glaucous willowherb
Epilobium glaciale P.H.Raven
Epilobium glaucum Phil.
Epilobium gouldii P.H.Raven
Epilobium gracilipes Kirk
Epilobium griffithianum Hausskn.
Epilobium gunnianum Hausskn.
Epilobium halleanum – glandular willowherb
Epilobium hectorii Hausskn.
Epilobium hirsutum – great willowherb, great hairy willowherb, hairy willowherb, codlins-and-cream, apple-pie, cherry-pie
Epilobium hirtigerum A.Cunn.
Epilobium hohuanense S.S.Ying
Epilobium hooglandii P.H.Raven
Epilobium hornemannii Rchb. – Hornemann's willowherb Epilobium hornemannii ssp. behringianum (Hausskn.) Hoch & P.H.Raven Epilobium hornemannii ssp. hornemannii
Epilobium howellii – Yuba Pass willowherb, subalpine fireweed
Epilobium indicum Hausskn.
Epilobium insulare Hausskn.
Epilobium kermodei P.H.Raven
Epilobium keysseri Diels
Epilobium kingdonii P.H.Raven
Epilobium komarovianum H.Lév. – bronzy willowherb
Epilobium lactiflorum – milkflower willowherb, whiteflower willowherb
Epilobium lanceolatum Sebast. & Mauri – spear-leaved willowherb
Epilobium laxum Royle
Epilobium leiophyllum Hausskn.
Epilobium leptocarpum Hausskn. – slenderfruit willowherb
Epilobium leptophyllum – bog willowherb, linear-leaved willowherb
Epilobium luteum Pursh – yellow willowherb
Epilobium macropus Hook.
Epilobium margaretiae Brockie
Epilobium matthewsii Petrie
Epilobium maysillesii Munz
Epilobium melanocaulon Hook.
Epilobium microphyllum A.Rich.
Epilobium minutiflorum Hausskn.
Epilobium mirabile Trel. – Olympic Mountain willowherb
Epilobium montanum – broad-leaved willowherb
Epilobium nankotaizanense Yamam. – Nankotaizan willowherb
Epilobium nerteroides A.Cunn.
Epilobium nivale Meyen
Epilobium nummulariifolium R.Cunn. ex A.Cunn.
Epilobium nutans F.W.Schmidt
Epilobium obcordatum – rockfringe willowherb
Epilobium obscurum Schreb. – dwarf willowherb, short-fruited willowherb
Epilobium oreganum – Grants Pass willowherb, Oregon fireweed
Epilobium oregonense – Oregon willowherb
Epilobium pallidiflorum Sol. ex A.Cunn.
Epilobium palustre – marsh willowherb
Epilobium pannosum Hausskn.
Epilobium parviflorum Schreb. – small-flowered willowherb, hoary willowherb
Epilobium pedicellare C.Presl
Epilobium pedunculare A.Cunn. – rockery willowherb
Epilobium pengii C.J.Chen, Hoch & P.H.Raven
Epilobium pernitens Cockayne & Allan
Epilobium perpusillum Hausskn.
Epilobium petraeum Heenan
Epilobium pictum Petrie
Epilobium platystigmatosum C.B.Rob.
Epilobium ponticum Hausskn.
Epilobium porphyrium G.Simpson
Epilobium prostratum Warb.
Epilobium psilotum Maire & Sam.
Epilobium pubens A.Rich.
Epilobium puberulum Hook. & Arn.
Epilobium purpuratum Hook.f.
Epilobium pycnostachyum Hausskn.
Epilobium pyrricholophum Franch. & Sav.
Epilobium rechingeri P.H.Raven
Epilobium rhynchospermum Boiss.
Epilobium roseum Schreb. – pale willowherb Epilobium roseum ssp. consimile (Hausskn.) P.H.Raven Epilobium roseum ssp. roseum Epilobium roseum ssp. subsessile (Boiss.) P.H.Raven
Epilobium rostratum Cheeseman
Epilobium rotundifolium G.Forst.
Epilobium royleanum Hausskn.
Epilobium rupicolum Pavlov
Epilobium salignum Hausskn.
Epilobium sarmentaceum Hausskn.
Epilobium saximontanum Hausskn. – Rocky Mountain willowherb
Epilobium sikkimense Hausskn.
Epilobium sinense H.Lév.
Epilobium siskiyouense – Siskiyou willowherb, Siskiyou fireweed
Epilobium smithii H.Lév.
Epilobium staintonii P.H.Raven
Epilobium stereophyllum Fresen.
Epilobium stracheyanum Hausskn.
Epilobium strictum Muhl. – downy willowherb, stiff spike-primrose
Epilobium subalgidum Hausskn.
Epilobium subcoriaceum Hausskn.
Epilobium subnivale Popov ex Pavlov
Epilobium taiwanianum C.J.Chen, Hoch & P.H.Raven
Epilobium tasmanicum Hausskn.
Epilobium tetragonum L. – square-stemmed willowherb Epilobium tetragonum ssp. lamyi (F.W.Schultz) Nyman Epilobium tetragonum ssp. tetragonum Epilobium tetragonum ssp. tournefortii (Michalet) H.Lév.
Epilobium tianschanicum Pavlov
Epilobium tibetanum Hausskn.
Epilobium trichophyllum Hausskn.
Epilobium vernonicum Snogerup
Epilobium wallichianum Hausskn.
Epilobium warakense Nábelek
Epilobium wattianum Hausskn.
Epilobium williamsii P.H.Raven
Epilobium willisii P.H. Raven & Engelhorn
Epilobium wilsonii Cheeseman
Epilobium × wisconsinense Ugent – Wisconsin willowherb (=E. ciliatum ssp. ciliatum × E. coloratum)
former Chamaenerion group
The following species are now segregated into the genus Chamaenerion Ség. The generic name Chamaenerion is preferred to Chamerion.[11])
Chamaenerion
Epilobium angustifolium
Epilobium conspersum
Epilobium latifolium
Epilobium speciosum
Rosmarinifolium
Epilobium colchicum
Epilobium dodonaei
Epilobium fleischeri
Epilobium stevenii
Formerly placed here
Stylidium tenellum (as E. tonkinense)
It is possible to distinguish between leaves of different Epilobium species using high-accuracy FT-IR method based on attenuated total reflection (ATR) without time-consuming preparation.[12]