Epipactis
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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 In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ...
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 any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
, that grow progressively shorter near the top. The margins are entire, the top is acute. Species with less
chlorophyll Chlorophyll (also chlorophyl) 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 allow plants to ...
have blue-purple leaves. Their bilaterally symmetrical colorful
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
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 The term ''sepalum'' was coine ...
s and the two lateral
petal Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corolla''. Petals are usuall ...
s are ovate and acuminate. Their color can vary from greenish-white to violet and purple. The
lip The lips are the visible body part at the mouth of many animals, including humans. Lips are soft, movable, and serve as the opening for food intake and in the articulation of sound and speech. Human lips are a tactile sensory organ, and can be ...
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 an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
is inferior. It produces a dry capsule with countless minute
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s.


Chemistry

As is characteristic of all orchids, ''Epipactis'' spp. are dependent on a
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, 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 plan ...
l
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
(see also
Orchid mycorrhiza Orchid mycorrhizae are endomycorrhizal fungi which develop symbiotic relationships with the roots and seeds of plants of the family Orchidaceae. Nearly all orchids are myco-heterotrophic at some point in their life cycle. Orchid mycorrhizae are cr ...
). 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 The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, and
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
. 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 ''Epipactis gigantea'' is a species of orchid known as the stream orchid, giant helleborine, and chatterbox. This wildflower is native to western North America from British Columbia to central Mexico. This is one of the most abundant orchids of ...
''). One species from Europe, broad-leaved helleborine ('' Epipactis helleborine''), is invasive in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
. Most species are protected. Most of these hardy orchids grow in a wet environment, but there are exceptions. The marsh helleborine (''
Epipactis palustris ''Epipactis palustris'', the marsh helleborine, is a species of orchid native to Europe and Asia. Description ''Epipactis palustris'' is a perennial herbaceous plant. This species has a stem growing to 60 cm high with as many as ten ere ...
'') is the only European orchid able to survive in a flooded habitat. ''
Epipactis gigantea ''Epipactis gigantea'' is a species of orchid known as the stream orchid, giant helleborine, and chatterbox. This wildflower is native to western North America from British Columbia to central Mexico. This is one of the most abundant orchids of ...
'' is a species found in the American west, and into southern
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by to ...
, 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 ''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 eme ...
'' (Ethiopia to Malawi) *''
Epipactis albensis ''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 eme ...
'' (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 ''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 eme ...
'' (Vietnam) *''
Epipactis atrorubens ''Epipactis atrorubens'', the dark-red helleborine or royal helleborine, is an herbaceous plant in the orchid family, Orchidaceae. As with many other species of orchids, the species is legally protected in some countries. Plantlife designated the ...
'' (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 ''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 eme ...
'' 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 ''Epipactis dunensis'', commonly known as dune helleborine, is a species of plant in the orchid family Orchidaceae and is endemic to Great Britain and Ireland. It typically grows to a height of and the upper half of the flowering stalk is hairy ...
'' (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 ''Epipactis gigantea'' is a species of orchid known as the stream orchid, giant helleborine, and chatterbox. This wildflower is native to western North America from British Columbia to central Mexico. This is one of the most abundant orchids of ...
'' : stream orchid, chatterbox, giant helleborine (W. Canada to N. Mexico). *''
Epipactis greuteri ''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 eme ...
'' 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 ''Epipactis palustris'', the marsh helleborine, is a species of orchid native to Europe and Asia. Description ''Epipactis palustris'' is a perennial herbaceous plant. This species has a stem growing to 60 cm high with as many as ten ere ...
'' : marsh helleborine, marsh orchid (Europe to Caucasus and Mongolia) *'' Epipactis papillosa'' (Russian Far East to Korea,
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the n ...
) *'' 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 called Holy Island, is a tidal island off the northeast coast of England, which constitutes the civil parish of Holy Island in Northumberland. Holy Island has a recorded history from the 6th century AD; it was an important ...
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).


External links

* * {{Taxonbar, from=Q161844 Neottieae genera Terrestrial orchids