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''Epipactis'', or helleborine, is a genus of terrestrial orchids consisting of approximately 70 species. This genus is abbreviated as Epcts in horticultural trade.


Description

Their creeping, fleshy rhizomes grow offshoots, from which then emerge the 20–70 cm long stems during the next spring. There are four to eight alternate, lanceolate
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
, that grow progressively shorter near the top. The margins are entire, the top is acute. Species with less
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
have blue-purple leaves. Their bilaterally symmetrical colorful
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s grow from a terminal raceme. The three
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s and the two lateral
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s are ovate and acuminate. Their color can vary from greenish-white to violet and purple. The
lip The lips are a horizontal pair of soft appendages attached to the jaws and are the most visible part of the mouth of many animals, including humans. Mammal lips are soft, movable and serve to facilitate the ingestion of food (e.g. sucklin ...
is divided in a bowl-shaped hypochile, with the outer surface greenish-white and threaded with dark veins. The wavy, snow-white epichile is fan-shaped. The
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
is inferior. It produces a dry capsule with countless minute
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s.


Chemistry

As is characteristic of all orchids, ''Epipactis'' spp. are dependent on a
mycorrhiza A mycorrhiza (; , mycorrhiza, or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...
l
symbiosis Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...
(see also Orchid mycorrhiza). This allows some species to have reduced leaves and need little chlorophyll. Violet helleborine (''Epipactis viridiflora'') can even do without chlorophyll. These forms can be recognized by their purple instead of violet flowers.


Habitat

The species occur in temperate and subtropical climates of America,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, and
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. These orchids grow in open spaces in forests, in undergrowth, on calcareous soils and are often found in wet dune-slacks near the sea. The only original American species is the giant helleborine ('' Epipactis gigantea''). One species from Europe, broad-leaved helleborine ('' Epipactis helleborine''), is invasive in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. Most species are protected. Most of these hardy orchids grow in a wet environment, but there are exceptions. The marsh helleborine ('' Epipactis palustris'') is the only European orchid able to survive in a flooded habitat. '' Epipactis gigantea'' is a species found in the American west, and into southern
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
, in wet areas and even streams. It can grow to a height of 1 m. However, '' Epipactis helleborine'' grows in more diverse habitats, from sheltered sandy beaches to open spaces in deciduous or coniferous forests, on roadsides, in meadows, and on moist soils. It is sometimes called the weed orchid.


Species

*'' Epipactis africana'' (Ethiopia to Malawi) *'' Epipactis albensis'' (C. Europe) **''Epipactis albensis'' var. ''albensis'' (EC. Europe). Rhizome geophyte **''Epipactis albensis'' var. ''fibri'' (France). Hemicryptophye or rhizome geophyte *''Epipactis aspromontana'' (Italy) - now synonym of ''Epipactis leptochila'' subsp. ''aspromontana'' (Bartolo, Pulv. & Robatsch) Kreutz *'' Epipactis atromarginata'' (Vietnam) *'' Epipactis atrorubens'' (Hoffm.) Besser : dark red helleborine, royal helleborine (Europe to Caucasus) **''Epipactis atrorubens'' var. ''atrorubens'' **''Epipactis atrorubens'' var. ''atrata'' A.Waldner & Webernd (Austria) **''Epipactis atrorubens'' subsp. ''danubialis'' (Robatsch & Rydlo) Ciocârlan & R.Rösler **''Epipactis atrorubens'' subsp. ''spiridonovii'' (Devillers-Tersch. & Devillers) Kreutz **''Epipactis atrorubens'' subsp. ''subclausa'' (Robatsch) Kreutz **''Epipactis atrorubens'' var. ''triploidea'' (Gelbr. & G.Hamel) Kreutz *'' Epipactis autumnalis'' D.Doro (Italy) *'' Epipactis baumanniorum'' Ströhle *''Epipactis bithynica'' (Turkey) - now synonym of ''Epipactis helleborine'' subsp. ''bithynica'' (Robatsch) Kreutz *'' Epipactis bugacensis'' (Hungary, near Bugacpusztaháza) *'' Epipactis campeadorii'' (Spain) *'' Epipactis cardina'' (Spain) *'' Epipactis condensata'' (Turkey to Lebanon, Cyprus, W. Transcaucasus) *'' Epipactis cretica'' (Crete) *''Epipactis danubialis'' (Romania) - now synonym of ''Epipactis atrorubens'' subsp. ''danubialis'' (Robatsch & Rydlo) Ciocârlan & R.Rösler *''Epipactis degenii'' (Greece) - now synonym of ''Epipactis halacsyi'' subsp. ''degenii'' (Szentp. & Mónus) Kreutz *'' Epipactis distans'' Arvet-Touvet (Central Europe) *'' Epipactis dunensis'' (Great Britain: N. England to N. Wales) *''Epipactis duriensis'' Bernardos, et al. (Portugal) - now synonym of ''Epipactis tremolsii'' var. ''duriensis'' (Bernardos, D.Tyteca, Revuelta & Amich) P.Delforge *''Epipactis exilis'' P.Delforge - now a synonym of ''Epipactis persica'' subsp. ''exilis'' (P.Delforge) Kreutz *''Epipactis flaminia'' (EC. Europe) - now synonym of ''Epipactis greuteri'' var. ''flaminia'' (P.R.Savelli & Aless.) Kreutz *'' Epipactis flava'' (Laos, Thailand). *''Epipactis futakii'' (EC. Europe) - now synonym of ''Epipactis leptochila'' var. ''futakii'' (Mered'a & Potek) P.Delforge *'' Epipactis gigantea'' : stream orchid, chatterbox, giant helleborine (W. Canada to N. Mexico). *'' Epipactis greuteri'' H.Baumann & Künkele (Greece). **''Epipactis greuteri'' var. ''flaminia'' (P.R.Savelli & Aless.) Kreutz **''Epipactis greuteri'' var. ''preinensis'' (Seiser) P.Delforge **''Epipactis greuteri'' subsp. ''preinensis'' (Austria) *'' Epipactis guegelii'' (Romania) *''Epipactis halacsyi'' Robatsch (Greece) - now basionym of ''Epipactis viridiflora'' subsp. ''halacsyi'' **''Epipactis halacsyi'' subsp. ''degenii'' (Szentp. & Mónus) Kreutz *'' Epipactis helleborine'' (L.) Crantz : broad-leaved helleborine (N. Africa, Europe to C. China, Type species). **''Epipactis helleborine'' subsp. ''bithynica'' (Robatsch) Kreutz **''Epipactis helleborine'' subsp. ''helleborine'' (N. Africa, Europe to C. China). **''Epipactis helleborine'' subsp. ''latina'' (Italy to NW. Balkan Pen). **''Epipactis helleborine'' subsp. ''leutei'' (Robatsch) Kreutz **''Epipactis helleborine'' var. ''minor'' R.Engel (France) **''Epipactis helleborine'' subsp. ''molochina'' (P.Delforge) Kreutz **''Epipactis helleborine'' subsp. ''neerlandica'' (W. Europe). **''Epipactis helleborine'' subsp. ''orbicularis'' (C. Europe) synonym of ''Epipactis distans'' Arvet-Touvet **''Epipactis helleborine'' subsp. ''schubertiorum'' (Bartolo, Pulv. & Robatsch) Kreutz **''Epipactis helleborine'' subsp. ''transcaucasica'' (Caucasus). Hemicryptophye or rhizome geophyte **''Epipactis helleborine'' subsp. ''tremolsii'' (W. Medit.) Hemicryptophye or rhizome geophyte **''Epipactis helleborine'' var. ''youngiana'' (A.J.Richards & A.F.Porter) Kreutz *''Epipactis ioessa'' Bongiorni, De Vivo, Fori & Romolini (Italy) *'' Epipactis kleinii'' (S. France to E. Spain) *''Epipactis komoricensis'' (EC. Europe) - now synonym of ''Epipactis leptochila'' subsp. ''komoricensis'' (Mered'a) Kreutz *'' Epipactis lapidocampi'' E.Klein & Laminger (Austria) *''Epipactis latifolia'' All., nom. illeg. (synonym of ''Epipactis helleborine'') **''Epipactis latifolia'' lus. rosea Erdner - now synonym of ''Epipactis viridiflora'' var. ''rosea'' (Erdner) Kreutz *'' Epipactis leptochila'' (Godfery) Godfery : narrow-lipped helleborine (Europe). **''Epipactis leptochila'' subsp. ''aspromontana'' (Bartolo, Pulv. & Robatsch) Kreutz **''Epipactis leptochila'' var. ''cleistogama'' (C.Thomas) Kreutz **''Epipactis leptochila'' var. ''dinarica'' (S.Hertel & Riech.) P.Delforge **''Epipactis leptochila'' var. ''futakii'' (Mered'a & Potek) P.Delforge **''Epipactis leptochila'' subsp. ''komoricensis'' (Mered'a) Kreutz **''Epipactis leptochila'' subsp. ''leptochila'' (Europe) **''Epipactis leptochila'' subsp. ''maestrazgona'' (P.Delforge & Gévaudan) Kreutz **''Epipactis leptochila'' subsp. ''naousaensis'' (Robatsch) Kreutz **''Epipactis leptochila'' subsp. ''neglecta'' (WC. Europe). Hemicryptophyte or rhizome geophyte **''Epipactis leptochila'' var. ''peitzii'' (H.Neumann & Wucherpf.) P.Delforge **''Epipactis leptochila'' subsp. ''sancta'' (P.Delforge) Kreutz *''Epipactis leutei'' (Austria) - now synonym of ''Epipactis helleborine'' subsp. ''leutei'' (Robatsch) Kreutz *'' Epipactis liestalensis'' (C. Europe). *''Epipactis lusitanica'' (S. and N. Portugal)- now synonym of ''Epipactis tremolsii'' subsp. ''lusitanica'' (D.Tyteca) Kreutz *'' Epipactis maestrazgona'' P.Delforge & Gévaudan (Spain) *'' Epipactis magnibracteata'' (NC. China) *'' Epipactis mairei'' (Nepal to C. China) *'' Epipactis maricae'' (Campania, Italy *''Epipactis mecsekensis'' (Hungary) - now a synonym of ''Epipactis nordeniorum'' subsp. ''mecsekensis'' (A.Molnár & Robatsch) Kreutz *'' Epipactis meridionalis'' H.Baumann & R.Lorenz (Sicilia to S. Italy) *'' Epipactis microphylla'' (Europe to Iran) *'' Epipactis molochina'' P.Delforge (Spain) *''Epipactis moravica'' Batousek (Czech Republic, Slovakia, Hungary) - now a synonym of ''Epipactis nordeniorum'' subsp. ''moravica'' (Batousek) Kreutz *'' Epipactis muelleri'' (W. & C. Europe). **''Epipactis muelleri'' subsp. ''cerritae'' (Sicilia). Hemicryptophye or rhizome geophyte **''Epipactis muelleri'' subsp. ''muelleri'' (W. & C. Europe) Rhizome geophyte *''Epipactis nauosaensis'' (Greece) - now a synonym of ''Epipactis leptochila'' subsp. ''naousaensis'' (Robatsch) Kreutz *'' Epipactis nordeniorum'' Robatsch (Austria) **''Epipactis nordeniorum'' subsp. ''mecsekensis'' (A.Molnár & Robatsch) Kreutz **''Epipactis nordeniorum'' subsp. ''moravica'' (Batousek) Kreutz *'' Epipactis ohwii'' (C. Taiwan) *'' Epipactis olympica'' (Greece) *'' Epipactis palustris'' : marsh helleborine, marsh orchid (Europe to Caucasus and Mongolia) *'' Epipactis papillosa'' (Russian Far East to Korea,
Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...
) *'' Epipactis persica'' (Soó) Nannf. (SE. Europe to W. Pakistan) **''Epipactis persica'' subsp. ''exilis'' (P.Delforge) Kreutz *'' Epipactis phyllanthes'' G.E.Sm. : green-flowered helleborine (W. & NW. Europe). **''Epipactis phyllanthes'' subsp. ''fageticola'' (C.E.Hermos.) Kreutz **''Epipactis phyllanthes'' var. ''olarionensis'' (France). Hemicryptophye or rhizome geophyte **''Epipactis phyllanthes'' var. ''phyllanthes'' (W. & NW. Europe) *'' Epipactis placentina'' (Switzerland to Italy). *''Epipactis pollinensis'' (Italy) - now synonym of ''Epipactis viridiflora'' var. ''pollinensis'' (B.Baumann & H.Baumann) Kreutz *'' Epipactis pontica'' (EC. Europe to N. Turkey) *'' Epipactis provincialis'' (France) *'' Epipactis pseudopurpurata'' (Slovakia) - ''Epipactis viridiflora'' subsp. ''pseudopurpurata'' (Mered'a) Kreutz *'' Epipactis purpurata'' *'' Epipactis rechingeri'' (N. Iran) *'' Epipactis rhodanensis'' (France to Austria) *'' Epipactis robatschiana'' Bartolo, D'Emerico, Pulv., Terrasi & Stuto (Italy) *'' Epipactis royleana'' (E. Afghanistan to Himalaya) *'' Epipactis sancta'':
Lindisfarne Lindisfarne, also known as Holy Island, is a tidal island off the northeast coast of England, which constitutes the civil parishes in England, civil parish of Holy Island in Northumberland. Holy Island has a recorded history from the 6th centu ...
helleborine *''Epipactis schubertiorum'' (Italy) - now synonym of ''Epipactis helleborine'' subsp. ''schubertiorum'' (Bartolo, Pulv. & Robatsch) Kreutz *''Epipactis spiridonovii'' (Bulgaria) - now synonym of ''Epipactis atrorubens'' subsp. ''spiridonovii'' (Devillers-Tersch. & Devillers) Kreutz *'' Epipactis stellifera'' (Switzerland) *''Epipactis subclausa'' (Greece) - now synonym of ''Epipactis atrorubens'' subsp. ''subclausa'' (Robatsch) Kreutz *'' Epipactis tallosii'' (Slovakia, Hungary) *'' Epipactis tenii'' (China) *'' Epipactis thessala'' (N. & C. Greece) *'' Epipactis thunbergii'' (S. Russian Far East to Korea, Japan to Nansei-shoto) *'' Epipactis tremolsii'' Pau **''Epipactis tremolsii'' subsp. ''densifolia'' (W.Hahn, Passin & R.Wegener) Kreutz **''Epipactis tremolsii'' var. ''duriensis'' (Bernardos, D.Tyteca, Revuelta & Amich) P.Delforge **''Epipactis tremolsii'' subsp. ''heraclea'' (P.Delforge & Kreutz) Kreutz **''Epipactis tremolsii'' subsp. ''lusitanica'' (D.Tyteca) Kreutz **''Epipactis tremolsii'' subsp. ''turcica'' (Kreutz) Kreutz *'' Epipactis troodi'' (Cyprus, Turkey ) *''Epipactis turcica'' (E. Aegean Is. to Turkey) - now synonym of ''Epipactis tremolsii'' subsp. ''turcica'' (Kreutz) Kreutz *'' Epipactis turcomanica'' (C. Asia) *'' Epipactis ulugurica'' (Tanzania) *'' Epipactis veratrifolia'' : scarce marsh helleborine (Caucasus to Somalia and SC. China) *'' Epipactis viridiflora'' (Hoffm.) Krock: clustered helleborine, violet helleborine (Europe) **''Epipactis viridiflora'' subsp. ''halacsyi'' (Robatsch) B.Baumann & H.Baumann (Greece) **''Epipactis viridiflora'' subsp. ''pollinensis'' (B.Baumann & H.Baumann) B.Baumann & H.Baumann (Italy) **''Epipactis viridiflora'' subsp. ''pseudopurpurata'' (Mered'a) Kreutz **''Epipactis viridiflora'' var. ''rosea'' (Erdner) Kreutz *'' Epipactis xanthophaea'' (China) *''Epipactis youngiana'' : Young's helleborine (Great Britain) - now synonym of ''Epipactis helleborine'' var. ''youngiana'' (A.J.Richards & A.F.Porter) Kreutz


Hybrids

*''Epipactis × amigoi'' (''E. helleborine'' × ''E. kleinii'') (Europe). *''Epipactis × barlae'' (''E. helleborine'' × ''E. microphylla'') (C. Europe). *''Epipactis × barreana'' (''E. latina'' × ''E. muelleri'') (Italy). *''Epipactis × breinerorum'' (''E. helleborine'' subsp. ''helleborine'' × ''E. greuteri'') (EC. Europe). *''Epipactis × bruxellensis'' (''E. helleborine'' × ''E. phyllanthes'') (W. Europe). *''Epipactis × capellonensis'' (''E. atrorubens'' × ''E. latina'') (Italy). *''Epipactis × cardonneae'' (''E. atrorubens'' × ''E. kleinii'') (W. Europe). *''Epipactis × conquensis'' (''E. cardina'' × ''E. kleinii'') (Spain). *''Epipactis × gerbaudiorum'' (''E. provincialis'' × ''E. tremolsii'') (France) *''Epipactis × gevaudanii'' (''E. helleborine'' × ''E. rhodanensis'') (France). *''Epipactis × graberi'' (''E. atrorubens'' × ''E. microphylla'') (Europe). *''Epipactis × heterogama'' (''E. atrorubens'' × ''E. muelleri'') (Europe). *''Epipactis x nicolosii'' M.P.Grasso & Grillo (. (E. helleborine (L.) Crantz x E. meridionalis H.Baumann & R.Lorenz) (Sicilia) *''Epipactis × populetorum'' ( ''E. helleborine'' × ''E. hispanica'') (Spain). *''Epipactis × pupplingensis'' ( ''E. atrorubens'' × ''E. palustris'') (Europe). *''Epipactis × reinekei'' (''E. helleborine'' × ''E. muelleri'') (Europe). *''Epipactis × robatschii'' Gévaudan & P.Delforge (''E. bugacensis'' Robatsch x ''E. atrorubens'' (Hoffm.) Besser subvar. ''borbasii'' ) *''Epipactis × schmalhausenii'' (''E. atrorubens'' × ''E. helleborine'') (Europe). **''Epipactis × schmalhausenii'' nothosubsp. ''fleischmannii'' (''E. atrorubens'' × ''E. helleborine'' subsp. ''orbicularis'') (Europe). Hemicryptophye or rhizome geophyte **''Epipactis × schmalhausenii'' nothosubsp. ''schmalhausenii'' (Europe). Rhizome geophyte *''Epipactis × schulzei'' (Europe). *''Epipactis × soguksuensis''(''E. helleborine'' × ''E. turcica'') (Turkey). *''Epipactis × stephensonii'' (''E. helleborine'' × ''E. leptochila'') (Europe). *''Epipactis × trikalana'' (''E. helleborine'' × ''E. thessala'') (Greece). *''Epipactis × vermionensis'' (''E. gracilis'' × ''E. helleborine'') (Greece).


References


External links

* * {{Authority control Neottieae genera Terrestrial orchids