Epipactis helleborine is a terrestrial species of orchid that is morphologically diverse[1] but identifiable by its star-shaped flowers.[2] It originated in the
| Louis.Hight/Epipactis helleborine | |
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Scientific classification 
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| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Monocots |
| Order: | Asparagales |
| Family: | Orchidaceae |
| Subfamily: | Epidendroideae |
| Genus: | Epipactis |
| Species: | E. helleborine
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| Binomial name | |
| Epipactis helleborine | |
| Synonyms[3][4] | |
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List
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Description edit
Epipactis helleborine varies in flower morphology and nutritional mode[1] and is rhizomatous.[5] Roots are thick and grow in dense masses.[2] Some plants are photoautotrophic and partially myco-heterotrophic to varying degrees while others are completely mycohetertrophic.[6][7] E. helleborine also has a wide range of height variation, growing to a height of between 15 centimeters to one meter.[2] Chlorophyl-free, mycohetertrophic orchids may be smaller than photosynthetic orchids.[6]
Leaves are broad, ribbed, and flat.[6][8] Leaves and stems of fully myco-heterotrophic E. helleborine are white, and the color of partially myco-heterotrophic plants ranges from yellow-ish green to dark green depending on the level of chlorophyll deficiency.[6] E. helleborine may also have variegated leaves.[6]
Star-shaped, greenish white flowers with labella that may be partly rose, brownish, or red[9][2] grow on spear-shaped bracts that are longer towards the base of the plant than at the top.[2] Flower number may be between 30 and 50[2] and is variable between populations.[1] Lateral sepals are mostly oval in shape, and the dorsal sepal is elliptical and slightly larger.[2] The size of the sepals varies between regions where E. helleborine is found.[1]
In the United States, E. helleborine blooms in the summer, and the blooming seasons are longer in areas with a greater number of plants.[2] The orchid may reproduce through pollination and germination but more commonly spreads through rhizomatous, clonal growth.[5]
Distribution edit
Epipactis helleborine is widespread across Europe and Asia, where its range stretches as far north as Scandinavia and Siberia down to northern Africa and the Himalayas and from Portugal to Japan[7][3][10][11] [12][13] It was found growing in the United States for the first time in 1879 after being introduced by uncertain means.[2] Since its discovery in the United States, it has spread throughout the country and into Canada.[2] In its non-native range in North America it may be considered invasive.[12][14][15]
Habitat edit
Epipactis helleborine is often found in woodland areas and in hedge-banks.[2][16] It grows in forested areas of various types of trees depending on region. For example, it is commonly found near pine forests in Japan[16] and near cottonwoods, willows, and other trees of the cottonwood forests along the Rio Grande.[2] It is also known for its colonization of anthropogenic habitats including parks, roadsides, quarries, and gravel pits, with many sites in Glasgow, London, and Moscow for example.[12][17][18][19] E. helleborine can live in low to bright light conditions.[2] This orchid species has adapted to multiple niches[16][2] and has spread rapidly in areas where it has been naturalized, particularly in soils rich in calcium.[12]
Ecology edit
Pollination edit
Epipactis helleborine relies on insects for pollination and is commonly pollinated by insects from the families Apidae, Culicidae, Formicidae, and Syrphidae.[19]
Some populations of E. helleborine are opportunistic regarding pollinators while others are adapted to be pollinated by insects with specific mouth types.[19] Insects are attracted to nectar secreted by the orchid, and eight populations in central Europe were found to produce nectar containing opioid compounds including oxycodone which may have an intoxicating effect on insects.[20] E. helleborine in dry habitats may also achieve pollination through selfing.[21]
Mycorrhizal Associations edit
E. helleborine forms symbiotic relationships with mycorrhizal fungi for its entire life cycle.[16] Germination requires mycorrhizal fungal symbionts that provide the dust seeds[22] of the orchid with nutrients, allowing them to develop.[16][23] Adult orchids may be partially or completely myco-heterotrophic.[7]
Over its global range, E. helleborine has been documented associating with several fungi of genera Hydnotrya, Leptodontidium, Peziza, Tuber, and Wilcoxina.[16][23] Locally, there is evidence that fungal selection by E. helleborine may be highly specific.[16][7] Because Wilcoxina also forms associations with pine trees, it has been speculated that when E. helleborine forms a symbiotic association with Wilcoxina fungi, the orchid may obtain carbon from pine trees through the fungi.[7] E. helleborine can survive off of the nutrients provided by its mycorrhizal fungi for over a year and, therefore, does not have to grow above ground every year.[2]
Subspecies (unchanged from current page) edit
A rather long list of names have been proposed for subspecies, varieties and forms of Epipactis helleborine, far too many to list here.[24] This is not unusual for such a widespread species. At present (November 2021) only the following are accorded international acceptance:[24]
- Epipactis helleborine subsp. bithynica (Robatsch) Kreutz - Turkey
- Epipactis helleborine subsp. helleborine - widespread
- Epipactis helleborine subsp. neerlandica (Verm.) Buttler - Great Britain, Norway, Denmark, Belgium, Netherlands, France, Germany
- Epipactis helleborine var. tangutica (Schltr.) S.C.Chen & G.H.Zhu - China
- Epipactis helleborine subsp. tremolsii (Pau) E.Klein - France, Spain, Portugal, Sardinia, Italy, Algeria, Morocco
References edit
- ^ a b c d 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.
- ^ a b c d e f g h i j k l m n o p Coleman, Ronald A. (2002). The Wild Orchids of Arizona and New Mexico. Cornell University Press. pp. 75–78. ISBN 978-0-8014-3950-6.
- ^ a b Kew World Checklist of Selected Plant Species, Epipactis helleborine
- ^ Kew World Checklist of Selected Plant Families, Epipactis helleborine subsp. helleborine, synonyms
- ^ a b Jakubska-Busse, Anna; Dudkiewicz, Małgorzata; Jankowski, Paweł; Sikora, Radosław (2009-12-01). "Mathematical inference of the underground clonal growth of Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae)". Botanica Helvetica. 119 (2): 69–76. doi:10.1007/s00035-009-0057-1. ISSN 1420-9063.
- ^ a b c d e General Morphology and Anatomy of Chlorophyll-free and Green Forms of Epipactis helleborine
- ^ a b c d e Suetsugu, Kenji; Yamato, Masahide; Miura, Chihiro; Yamaguchi, Katsushi; Takahashi, Kazuya; Ida, Yoshiko; Shigenobu, Shuji; Kaminaka, Hironori. "Comparison of green and albino individuals of the partially mycoheterotrophic orchid Epipactis helleborine on molecular identities of mycorrhizal fungi, nutritional modes and gene expression in mycorrhizal roots". Molecular Ecology. 26 (6): 1652–1669. doi:10.1111/mec.14021.
- ^ 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.)
- ^ Webb, D.A., Parnell, J. and Doogue,D . 1996. An Irish Flora. Dundalgan Press (W. Tempest) Ltd. Dundalk.ISBN 0-85221-131-7
- ^ Altervista Flora Italiana, Elleborine comune, Epipactis helleborine (L.) Crantz
- ^ Flora of China v 25 p 180, 火烧兰 huo shao lan, Epipactis helleborine (Linnaeus) Crantz
- ^ a b c d Kolanowska, Marta (2013-10-15). "Niche Conservatism and the Future Potential Range of Epipactis helleborine (Orchidaceae)". PLOS ONE. 8 (10): e77352. doi:10.1371/journal.pone.0077352. ISSN 1932-6203. PMC 3797094. PMID 24143222.
{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link) - ^ Łobas, Zbigniew; Khapugin, Anatoliy; Żołubak, Elżbieta; Jakubska-Busse, Anna (2021-09-04). "The Epipactis helleborine Group (Orchidaceae): An Overview of Recent Taxonomic Changes, with an Updated List of Currently Accepted Taxa". Plants. 10 (9): 1839. doi:10.3390/plants10091839. ISSN 2223-7747. PMC 8469704. PMID 34579372.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Tenney, Angela; Hill, Erin (2022-06-24). "Broad-leaved helleborine: A weedy orchid invading lawns and flowerbeds". Michigan State University Extension.
- ^ "Plant Invaders of Mid-Atlantic Natural Areas: Plants to Watch". Archived from the original on 2010-12-10.
- ^ a b c d e f g Ogura-Tsujita, Yuki; Yukawa, Tomohisa. "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. ISSN 0940-6360. PMID 18661158.
- ^ 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
- ^ 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
- ^ a b c 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.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ 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.
- ^ Suetsugu, Kenji. "Delayed autonomous self-pollination in two Japanese varieties of E pipactis helleborine (Orchidaceae): Autogamy in Epipactis Helleborine". Botanical Journal of the Linnean Society. 173 (4): 733–743. doi:10.1111/boj.12111.
- ^ Eriksson, Ove; Kainulainen, Kent. "The evolutionary ecology of dust seeds". Perspectives in Plant Ecology, Evolution and Systematics. 13 (2): 73–87. doi:10.1016/j.ppees.2011.02.002.
- ^ a b Jacquemyn, Hans; Waud, Michael; Lievens, Bart; Brys, Rein. "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. ISSN 0305-7364. PMC 4934391. PMID 26946528.
- ^ a b Kew World Checklist of Selected Plant Families - List of names for Epipactis helleborine
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