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Epipactis helleborine

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Epipactis helleborine
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Asparagales
Family: Orchidaceae
Subfamily: Epidendroideae
Genus: Epipactis
Species:
E. helleborine
Binomial name
Epipactis helleborine
Synonyms[1][2]
List
  • Serapias helleborine L.
  • Helleborine helleborine (L.) Druce, invalid name
  • Epipactis latifolia subsp. helleborine (L.) Rivas Goday & Borja, illegitimate name
  • Serapias helleborine var. latifolia L.
  • Serapias latifolia (L.) Huds.
  • Epipactis latifolia (L.) All.
  • Cymbidium latifolium (L.) Sw.
  • Helleborine latifolia (L.) Moench
  • Epipactis consimilis D.Don
  • Epipactis uliginosa Vest
  • Epipactis herbacea Lindl. in J.F.Royle
  • Epipactis macrostachya Lindl.
  • Epipactis ovalis Bab. in J.E.Sm.
  • Epipactis pycnostachys K.Koch
  • Epipactis dalhousiae Wight
  • Epipactis intrusa Lindl.
  • Epipactis gutta-sanguinis Arv.-Touv.
  • Calliphyllon latifolium (L.) Bubani
  • Epipactis atroviridis Linton
  • Helleborine ovalis (Bab.) Druce
  • Helleborine atroviridis (Linton) F.Hanb. in H.C.Watson
  • Serapias consimilis (D.Don) A.A.Eaton
  • Helleborine pycnostachys (K.Koch) Druce
  • Amesia consimilis (D.Don) A.Nelson & J.F.Macbr.
  • Amesia latifolia (L.) A.Nelson & J.F.Macbr.
  • Amesia pycnostachys (K.Koch) A.Nelson & J.F.Macbr.
  • Epipactis squamellosa Schltr.
  • Epipactis yunnanensis Schltr.
  • Epipactis discolor Kraenzl.
  • Epipactis tenii Schltr.
  • Epipactis monticola Schltr.
  • Epipactis nephrocordia Schltr.
  • Amesia discolor (Kraenzl.) Hu
  • Amesia monticola (Schltr.) Hu
  • Amesia squamellosa (Schltr.) Hu
  • Amesia tenii (Schltr.) Hu
  • Amesia yunnanensis (Schltr.) Hu
  • Helleborine varians Soó
  • Amesia longibracteata Schweinf.
  • Helleborine macrostachya (Lindl.) Soó
  • Helleborine nephrocardia (Schltr.) Soó
  • Helleborine squamellosa (Schltr.) Soó
  • Helleborine tenii (Schltr.) Soó
  • Helleborine yunnanensis (Schltr.) Soó
  • Epipactis ligulata Hand.-Mazz.
  • Epipactis magnibracteata C.Schweinf.
  • Epipactis youngiana A.J.Richards & A.F.Porter
  • Epipactis voethii Robatsch
  • Epipactis kezlinekii Batoušek
  • Epipactis zirnsackiana Riech.
Flowers

Epipactis helleborine, the broad-leaved helleborine, is a terrestrial species of orchid with a broad distribution. It is a long lived herb which varies morphologically with ability to self-pollinate.[3]

Description

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Epipactis helleborine can grow to a maximum height of 1 m (3 ft 3 in) or more under good conditions, and has broad dull green leaves which are strongly ribbed and flat.[4][5] The flowers are arranged in long drooping racemes with dull green sepals and shorter upper petals. The lower labellum is pale red and is much shorter than the upper petals.[6]

Achlorophyllous, white Epipactis helleborine plants have been found. Achlorophyllous forms tend to be shorter, as small as 17 cm.[4]

Flowering occurs June–September.[7]

Distribution

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This species is widespread across much of Europe and Asia, from Portugal to China, as well as northern Africa.[1][8][9]

In the United Kingdom, the species is widespread and common in England and Wales, but is absent from most of Scotland scattered across Ireland.[10]

In North America, it is an introduced species and widely naturalized mostly in the Northeastern United States, eastern Canada and the Great Lakes Region, but also in scattered locations in other parts of the continent.[11][12][1] In the US it is sometimes referred to as the "weed orchid" or "weedy orchid" and continues to spread throughout the country to new areas including Michigan,[13] Wisconsin,[14] and the San Francisco Bay Area.[15]

Habitat

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Found in woods and hedge-banks[16] and often not far from paths near human activity.[17] It is one of the most likely European orchids to be found within a city, with many sites for example in Glasgow, London and Moscow. Sometimes spotted beside car parks.[18]

Epipactis helleborine is known for its successful colonization of human-made or anthropogenic habitats such as parks, gardens or roadsides.[18] These roadside orchids exhibit special features such as large plant size and greater ability to produce flowers.[18] Pollination plays a huge role as pollinators such as Syrphidae, Culicidae, Apidae etc. possess greater species diversity and visits the flowering sites more in anthropogenic habitats as compared to native ones.[18] The visitation rates along with the reproductive success of these orchids are higher in large populations as they are more attractive to pollinators.[3]

Ecology

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Hoverfly depositing an egg on an Epipactis helleborine leaf as a predatory response to aphids which are farmed by ants.

This species of orchid is pollinated by several species of Hymenoptera, particularly the common wasp, but also other species in the genera Vespula and Dolichovespula. Flowers release a sweet nectar to attract the wasps, which has an intoxicating effect on them.[19][20] Eight populations of Epipactis helleborine in central Europe (Lower Silesia, Poland) had their nectar studied and they were found to contain naturally occurring oxycodone (as well as another narcotic-like opioid) in minute amounts.[21]

Epipactis helleborine requires a mycorrhizal symbiosis to germinate successfully and remains partially dependent upon the fungus when plants mature, however it is not particularly selective among fungal species. Fungi associated with the live roots include Tuber, Helotiales, Peziza, Leptodontidium, Hydnotrya and Wilcoxina.[22][23]

It has been suggested that the presence of this orchid species in a woodland is an indicator that edible truffles can be found there,[24] but this is not always the case.

Subspecies

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A rather long list of names have been proposed for subspecies, varieties and forms of Epipactis helleborine, far too many to list here.[25] This is not unusual for such a widespread species. At present (November 2021) only the following are accorded international acceptance:[25]

  1. Epipactis helleborine subsp. bithynica (Robatsch) Kreutz - Turkey
  2. Epipactis helleborine subsp. helleborine - widespread
  3. Epipactis helleborine subsp. neerlandica (Verm.) Buttler - Great Britain, Norway, Denmark, Belgium, Netherlands, France, Germany
  4. Epipactis helleborine var. tangutica (Schltr.) S.C.Chen & G.H.Zhu - China
  5. Epipactis helleborine subsp. tremolsii (Pau) E.Klein - France, Spain, Portugal, Sardinia, Italy, Algeria, Morocco

Chemistry

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According to a study published in 2005 by Jakubska et al. trace amounts of narcotic compounds have been identified in the plants nectar, namely 3-{2-{3-{3-(benzyloxy)propyl}-3-indol, 7,8-didehydro- 4,5-epoxy-3,6-d-morphinan and oxycodone.[21] This is still debated however, as there is no evidence that such molecules could be readily biosynthesized in a plant.

References

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  1. ^ a b c Kew World Checklist of Selected Plant Species, Epipactis helleborine
  2. ^ Kew World Checklist of Selected Plant Families, Epipactis helleborine subsp. helleborine, synonyms
  3. ^ a b Ehlers, B. K.; Olesen, J. M.; Ågren, J. (2002). "Floral morphology and reproductive success in the orchid Epipactis helleborine: regional and local across-habitat variation". Plant Systematics and Evolution. 236 (1/2): 19–32. doi:10.1007/s00606-002-0197-x. ISSN 0378-2697. JSTOR 23644960. S2CID 9878820.
  4. ^ a b General Morphology and Anatomy of Chlorophyll-free and Green Forms of Epipactis helleborine
  5. ^ Solbraa, Knut (2013). 50 norske og svenske orkideer (in Norwegian). Oplandske Bokforlag, Vallset. Norway. p. 26. ISBN 978-82-7518-211-9. Under gode vekstforhold kan den nå høyder på over meteren. (Under good conditions it can reach heights of more than a metre.)
  6. ^ Webb, D.A., Parnell, J. and Doogue,D . 1996. An Irish Flora. Dundalgan Press (W. Tempest) Ltd. Dundalk.ISBN 0-85221-131-7
  7. ^ First Nature - Epipactis Helleborine
  8. ^ Altervista Flora Italiana, Elleborine comune, Epipactis helleborine (L.) Crantz
  9. ^ Flora of China v 25 p 180, 火烧兰 huo shao lan, Epipactis helleborine (Linnaeus) Crantz
  10. ^ Cole, Sean (2020). Britain's Orchids. WildGuides. p. 126.
  11. ^ Flora of North America v 26 p 586, Epipactis helleborine (Linnaeus) Crantz
  12. ^ Biota of North America Program, county range map
  13. ^ Tenney, Angela; Hill, Erin (2022-06-24). "Broad-leaved helleborine: A weedy orchid invading lawns and flowerbeds". Michigan State University Extension.
  14. ^ "Plant Invaders of Mid-Atlantic Natural Areas: Plants to Watch". Archived from the original on 2010-12-10.
  15. ^ "Epipactis helleborine".
  16. ^ Clapham, A.R., Tutin, T.G. and Warburg, E.F. 1968. Excursion Flora of the British Isles. Cambridge University Press ISBN 0 521 04656 4
  17. ^ Beesley, S. and Wild, J. 1997. Urban Flora of Belfast. Institute of Irish Studies and The Queen's University of Belfast. ISBN 0-85389-695 X
  18. ^ a b c d Rewicz, Agnieszka; Jaskuła, Radomir; Rewicz, Tomasz; Tończyk, Grzegorz (2017-04-18). "Pollinator diversity and reproductive success of Epipactis helleborine (L.) Crantz (Orchidaceae) in anthropogenic and natural habitats". PeerJ. 5: e3159. doi:10.7717/peerj.3159. ISSN 2167-8359. PMC 5398293. PMID 28439457.
  19. ^ "London Wildlife Trust - Orchid for July". Archived from the original on 2018-12-19. Retrieved 2018-12-19.
  20. ^ Bioinfo - Epipactis helleborine
  21. ^ a b Jakubska, A.; Przado, D.; Steininger, M.; Aniol-Kwaitkowska, J.; Kadej, M. (2005). "Why do pollinators become "sluggish"? Nectar chemical constituents from Epipactus helleborine (L.) Crantz (Orchidaceae)". Applied Ecology and Environmental Research. 3 (2): 29–38. doi:10.15666/aeer/0302_029038.
  22. ^ Jacquemyn, H.; Waud, M.; Lievens, B.; Brys, R. (2016). "Differences in mycorrhizal communities between Epipactis palustris, e. Helleborine and its presumed sister species e. Neerlandica". Annals of Botany. 118 (1): 105–114. doi:10.1093/aob/mcw015. PMC 4934391. PMID 26946528.
  23. ^ Ogura-Tsujita, Yuki; Yukawa, Tomohisa (2008). "Epipactis helleborine shows strong mycorrhizal preference towards ectomycorrhizal fungi with contrasting geographic distributions in Japan". Mycorrhiza. 18 (6–7): 331–338. doi:10.1007/s00572-008-0187-0. PMID 18661158. S2CID 19796924.
  24. ^ Ouanphanivanh, Noémi; Merényi, Zsolt; Orczán, Ákos Kund; Bratek, Zoltán; Szigeti, Zoltán; Illyés, Zoltán (January 2008). "Could orchids indicate truffle habitats? Mycorrhizal association between orchids and truffles". Acta Biologica Szegediensis. 52 (1): 229–232.
  25. ^ a b Kew World Checklist of Selected Plant Families - List of names for Epipactis helleborine
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