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| − | '''''Acorus''''' is a [[genus]] of [[monocot]] [[flowering plant]]s. This genus was once placed within the family [[Araceae]] (aroids), but more recent classifications place it in its own family '''Acoraceae''' and order '''Acorales''', of which it is the sole genus of the oldest surviving line of monocots. Some older studies indicated that it was placed in a lineage (the order Alismatales), that also includes aroids (Araceae), [[Tofieldiaceae]], and several families of aquatic monocots (e.g., Alismataceae, [[Posidoniaceae]]). However, modern phylogenetic studies demonstrate that ''Acorus'' is [[sister group|sister]] to all other monocots. Common names include '''calamus''' and '''[[sweet flag]]'''. | + | The genus '''''Acorus''''' is the only member of the family '''''Acoraceae''''' and the order '''''Acorales'''''. It contains several species of wetland plants whose roots are used as a flavoring. Some other plant parts are said to be edible as well. |
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| − | The genus is native to [[North America]] and northern and eastern [[Asia]], and [[naturalised]] in southern Asia and [[Europe]] from ancient cultivation.<ref name=k/><ref>Govaerts, R. & Frodin, D.G. (2002). World Checklist and Bibliography of Araceae (and Acoraceae): 1-560. The Board of Trustees of the Royal Botanic Gardens, Kew.</ref><ref>[http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=100307 Flora of North America: ''Acorus'']</ref><ref>[http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=100307 Flora of China, Vol. 23 Page 1, <big>菖蒲属</big> chang pu shu, ''Acorus'' Linnaeus, Sp. Pl. 1: 324. 1753. ]</ref><ref name="Boyce, P.C. 2012">Boyce, P.C., Sookchaloem, D., Hetterscheid, W.L.A., Gusman, G., Jacobsen, N., Idei, T. & Nguyen, V.D. (2012). Flora of Thailand 11(2): 101-325. The Forest Herbarium, National Park, Wildlife and Plant Conservation Department, Bangkok.</ref><ref name="Boyce, P.C. 2012"/><ref>Nooteboom, H.P. (ed.) (2011). Flora Malesiana 20: 1-61. Noordhoff-Kolff N.V., Djakarta.</ref> The known wild populations are [[diploid]] except for some [[tetraploid]]s in eastern Asia, while the cultivated plants are sterile [[triploid]]s, probably of [[Hybrid (biology)|hybrid]] origin between the diploid and tetraploid forms.
| + | There is some concern over levels of the carcinogen asarone, so extracts of these plants have been banned in the US. |
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| − | ==Characteristics== | + | ==Species== |
| − | [[File:Acorus calamus1.jpg|thumb|220px|Habit of ''Acorus calamus''.]]
| + | * ''[[Acorus americanus]]'' (or ''A. calamus var. americanus'') |
| − | The inconspicuous [[flower]]s are arranged on a lateral [[spadix (botany)|spadix]] (a thickened, fleshy axis). Unlike aroids, there is no [[spathe]] (large bract, enclosing the spadix). The spadix is 4–10 cm long and is enclosed by the foliage. The bract can be ten times longer than the spadix. The [[leaf|leaves]] are linear with entire margin.
| + | * ''[[Acorus calamus]]'' |
| − | | + | * ''[[Acorus gramineus]]'' |
| − | ==Taxonomy==
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| − | Although the family Acoraceae was originally described in 1820, since then ''Acorus'' has traditionally been included in Araceae in most classification systems, as in the [[Cronquist system]]. The family has recently been resurrected as molecular systematic studies have shown that ''Acorus'' is not closely related to Araceae or any other [[monocot]] family, leading plant systematists to place the genus and family in its own order. This placement currently lacks support from traditional plant morphology studies, and some taxonomists still place it as a subfamily of Araceae, in the order [[Alismatales]]. The [[APG III]] system recognizes order Acorales, distinct from the Alismatales, and as the [[sister group]] to all other monocots. This relationship is confirmed by more recent phylogenetic studies.{{sfn|Soltis et al|2011}}{{sfn|Givnish et al|2010}} Treatment in the [[APG IV system]] is unchanged from APG III.<ref name=APGIV>{{Cite journal|authors=Angiosperm Phylogeny Group|authorlink=Angiosperm Phylogeny Group|year=2016|title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV|journal=[[Botanical Journal of the Linnean Society]]|volume=181|issue=1|pages=1–20|url=http://onlinelibrary.wiley.com/doi/10.1111/boj.12385/epdf|format=PDF|issn=00244074|doi=10.1111/boj.12385}}</ref>
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| − | === Species ===
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| − | In older literature and on many websites, there is still much confusion, with the name ''Acorus calamus'' equally but wrongfully applied to ''Acorus americanus'' (formerly ''Acorus calamus'' var. ''americanus'').
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| − | As of July 2014, the Kew Checklist accepts only 2 species, one of which has three accepted varieties:<ref name=k/>
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| − | * ''[[Acorus calamus]]'' <small>[[Carl Linnaeus|L.]]</small> – '''Common sweet flag'''; sterile triploid (3''n'' = 36); probably of cultivated origin. It is native to [[Europe]], temperate [[India]] and the [[Himalaya]]s and southern [[Asia]], widely cultivated and naturalised elsewhere. | |
| − | **[[Acorus americanus|''Acorus calamus'' var. ''americanus'']] <small>Raf.</small> - Canada, northern United States, Buryatiya region of Russia
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| − | **''Acorus calamus'' var. ''angustatus'' <small>Besser</small> - Siberia, China, Russian Far East, Japan, Korea, Mongolia, Himalayas, Indian Subcontinent, Indochina, Philippines, Indonesia
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| − | **''Acorus calamus'' var. ''calamus'' - Siberia, Russian Far east, Mongolia, Manchuria, Korea, Himalayas; naturalized in Europe, North America, Java and New Guinea
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| − | * ''[[Acorus gramineus]]'' <small>[[Daniel Solander|Sol.]] ex [[William Aiton|Aiton]]</small> – '''Japanese sweet flag''' or '''grassy-leaved sweet flag'''; fertile diploid (2''n'' = 18); - China, Himalayas, Japan, Korea, Indochina, Philippines, Primorye | |
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| − | ''Acorus'' from Europe, [[China]] and Japan have been planted in the [[United States]].
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| − | === Etymology ===
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| − | The name 'acorus' is derived from the Greek word 'acoron', a name used by [[Dioscorides]], which in turn was derived from 'coreon', meaning 'pupil', because it was used in [[herbalism|herbal medicine]] as a treatment for [[inflammation]] of the [[Human eye|eye]].
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| − | == Distribution and habitat ==
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| − | These plants are found in [[wetlands]], particularly marshes, where they spread by means of thick rhizomes. Like many other marsh plants, they depend upon [[aerenchyma]] to transport oxygen to the rooting zone.<ref>Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. Chapter 1.</ref> They frequently occur on shorelines and floodplains where water levels fluctuate seasonally.
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| − | == Ecology ==
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| − | The native North American species appears in many ecological studies. Compared to other species of wetland plants, they have relatively high competitive ability.<ref>{{cite journal | last1 = Gaudet | first1 = C.L. | last2 = Keddy | first2 = P.A. | year = 1988 | title = Predicting competitive ability from plant traits: a comparative approach | url = | journal = Nature | volume = 334 | issue = | pages = 242–243 | doi=10.1038/334242a0}} Figure 1.</ref> Although many marsh plants accumulate large banks of buried seeds,<ref>{{cite journal | last1 = van der Valk | first1 = A. G. | last2 = Davis | first2 = C. B. | year = 1978 | title = The role of seed banks in the vegetation dynamics of prairie glacial marshes | url = | journal = Ecology | volume = 59 | issue = | pages = 322–35 | doi=10.2307/1936377}}</ref> seed banks of ''Acorus'' may not accumulate in some wetlands owing to low seed production.<ref>{{cite journal | last1 = Leck | first1 = Mary Allessio | last2 = Simpson | first2 = Robert L. | year = 1995 | title = Ten-year seed bank and vegetation dynamics of a tidal freshwater marsh | url = | journal = American Journal of Botany | volume = 82 | issue = | pages = 1547–1557 | doi=10.2307/2446183}}</ref> The seeds appear to be adapted to germinate in clearings; after a period of cold storage, the seeds will germinate after seven days of light with fluctuating temperature, and somewhat longer under constant temperature.<ref>{{cite journal | last1 = Shipley | first1 = B. | last2 = Keddy | first2 = P.A. | last3 = Moore | first3 = D.R.J. | last4 = Lemky | first4 = K. | year = 1990 | title = Regeneration and establishment strategies of emergent macrophytes | url = | journal = Journal of Ecology | volume = 77 | issue = | pages = 1093–1110 | doi=10.2307/2260825}} Appendix 3.</ref> A comparative study of its life history traits classified it as a "tussock interstitial", that is, a species that has a dense growth form and tends to occupy gaps in marsh vegetation, not unlike ''[[Iris versicolor]]''.<ref>{{cite journal | last1 = Boutin | first1 = C. | last2 = Keddy | first2 = P. A. | year = 1993 | title = A functional classification of wetland plants | url = | journal = Journal of Vegetation Science | volume = 4 | issue = | pages = 591–600 | doi=10.2307/3236124}} Figure 2</ref>
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| − | ==Toxicity ==
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| − | [[File:Acorus calamus illustration.jpg|thumb|right|Sweet Flag (2006 drawing by USGS Northern Prairie Wildlife Research Center)]]
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| − | Products derived from ''Acorus calamus'' were banned in 1968 as food additives by the United States [[Food and Drug Administration]].<ref>{{cite web|
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| − | url=http://www.accessdata.fda.gov/scripts/cdrh/cfdocs/cfcfr/CFRSearch.cfm?fr=189.110 |title=Code of Federal regulations, title 21}}</ref> The questionable chemical derived from the plant was [[Asarone|β-asarone]]. Confusion exists whether all strains of ''A. calamus'' contain this substance.
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| − | Four varieties of ''A. calamus'' strains exist in nature: diploid, triploid, tetraploid and hexaploid.<ref>Ginwal, HS, An efficient genomic DNA isolation protocol for RAPD and SSR analysis in {{tx|Acorus calamus|L.}}</ref> Diploids do not produce the carcinogenic β-asarone. Diploids are known to grow naturally in Eastern Asia (Mongolia and C Siberia) and North America. The triploid cytotype probably originated in the Himalayan region, as a hybrid between the diploid and tetraploid cytotypes.<ref>Evstatieva et al., Fitologiya 48: 19–22. 1996; Löve & Löve, ''Proc. Genet. Soc. Canada'' 2: 14–17. 1957</ref> The North American Calamus is known as ''Acorus calamus'' var. ''americanus'' or more recently as simply ''Acorus americanus''. Like the diploid strains of ''A. calamus'' in parts of the Himalayas, Mongolia, and C Siberia, the North American diploid strain does not contain the carcinogenic β-asarone.<ref>{{cite journal | last1 = Marongiu | first1 = L.B | last2 = Piras | first2 = A | last3 = Porcedda | first3 = S | year = 2005 | title = Chemical composition of the essential oil and supercritical CO2 extract of Commiphora myrrha (Nees) Engl. and of ''Acorus calamus'' | url = | journal = J. Agric | volume = | issue = | page = }}</ref><ref>(Rost and Bos, 1979)</ref><ref>Antimicrobial activities of the crude methanol extract of ''Acorus calamus'' Linn., S Phongpaichit, N Pujenjob, J. Songklanakarin</ref> Research has consistently demonstrated that "β-asarone was not detectable in the North American spontaneous diploid Acorus [Calamus var. Americanus]".<ref>{{Cite journal | last1 = Radušienė | first1 = J. | last2 = Judžentienė | first2 = A. | last3 = Pečiulytė | first3 = D. | last4 = Janulis | first4 = V. | title = Essential oil composition and antimicrobial assay of Acorus calamus leaves from different wild populations | doi = 10.1017/S1479262107390928 | journal = Plant Genetic Resources | volume = 5 | pages = 37 | year = 2007 | pmid = | pmc = }}</ref>
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| − | == Uses ==
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| − | The parallel-veined leaves of some species contain ethereal oils that give a sweet scent when dried. Fine-cut leaves used to be strewn across the floor in the Middle Ages, both for the scent, and for presumed efficacy against [[pest (animal)|pest]]s.
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| | ==References== | | ==References== |
| | {{Reflist|30em}} | | {{Reflist|30em}} |
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| − | == Bibliography ==
| + | {{Ack-Wikipedia}} |
| − | {{refbegin|30em}} | |
| − | * Govaerts, R. & Frodin, D.G. (2002). World Checklist and Bibliography of Araceae and Acoraceae. 1-560. The Board of Trustees of the Royal Botanic Gardens, Kew.
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| − | * [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=20008 Flora of North America: Acoraceae]
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| − | * [http://www.mobot.org/MOBOT/Research/APweb/orders/acoralesweb.htm#Acorales Acorales] in Stevens, P. F. (2001 onwards). [http://www.mobot.org/MOBOT/research/APweb Angiosperm Phylogeny Website]. Version 7, May 2006.
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| − | * [http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=91812&lvl=3&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser]
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| − | * [https://web.archive.org/web/20050209163227/http://delta-intkey.com:80/angio/www/acoracea.htm Acoraceae] in L. Watson and M.J. Dallwitz (1992 onwards) ''[http://delta-intkey.com/angio/ The families of flowering plants:] descriptions, illustrations, identification, information retrieval''. Version: 27 April 2006. http://delta-intkey.com.
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| − | * [https://web.archive.org/web/20050910122209/http://www.woodlotalt.com:80/publications/BotNotesv1n2.PDF Taxonomy and distribution of Acorus in Maine]
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| − | * [http://www.karenplatt.co.uk/garden-books/gold-fever.html Platt, Karen. Gold Fever 2004] {{ISBN|978-0954576417}}
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| − | * [http://www.pnas.org/content/90/10/4641.short Phylogenetic analysis of rbcL sequences identifies Acorus calamus as the primal extant monocotyledon. Duvall 1993]
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| − | * {{cite journal|last1=Duvall|first1=Melvin R.|last2=Clegg|first2=Michael T.|last3=Chase|first3=Mark W.|authorlink3=Mark Wayne Chase|last4=Clark|first4=W. Dennis|last5=Kress|first5=W. John|last6=Hills|first6=Harold G.|last7=Eguiarte|first7=Luis E.|last8=Smith|first8=James F.|last9=Gaut|first9=Brandon S.|last10=Zimmer|first10=Elizabeth A.|last11=Learn|first11=Gerald H.|title=Phylogenetic Hypotheses for the Monocotyledons Constructed from ''rbc''L Sequence Data|journal=[[Annals of the Missouri Botanical Garden]]|date=1 January 1993|volume=80|issue=3|pages=607–619|doi=10.2307/2399849|jstor=2399849}}
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| − | * [http://mbe.oxfordjournals.org/content/22/9/1813.short Analysis of Acorus calamus Chloroplast Genome and Its Phylogenetic Implications Vadim V. Goremykin 2005]
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| − | * {{cite journal|last1=Givnish|first1=Thomas J.|authorlink1=Thomas J. Givnish|last2=Ames|first2=Mercedes|last3=McNeal|first3=Joel R.|last4=McKain|first4=Michael R.|last5=Steele|first5=P. Roxanne|last6=dePamphilis|first6=Claude W.|last7=Graham|first7=Sean W.|last8=Pires|first8=J. Chris|last9=Stevenson|first9=Dennis W.|last10=Zomlefer|first10=Wendy B.|last11=Briggs|first11=Barbara G.|last12=Duvall|first12=Melvin R.|last13=Moore|first13=Michael J.|last14=Heaney|first14=J. Michael|last15=Soltis|first15=Douglas E.|authorlink15=Douglas E. Soltis|last16=Soltis|first16=Pamela S.|authorlink16 = Pamela S. Soltis|last17=Thiele|first17=Kevin|authorlink17=Kevin Thiele|last18=Leebens-Mack|first18=James H.|title=Assembling the Tree of the Monocotyledons: Plastome Sequence Phylogeny and Evolution of Poales|url=http://www.bioone.org/doi/abs/10.3417/2010023|journal=[[Annals of the Missouri Botanical Garden]]|date=27 December 2010|volume=97|issue=4|pages=584–616|doi=10.3417/2010023|ref={{harvid|Givnish et al|2010}}}}
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| − | * {{cite journal|last1=Soltis|first1=D. E.|authorlink1=Douglas E. Soltis|last2=Smith|first2=S. A.|last3=Cellinese|first3=N.|last4=Wurdack|first4=K. J.|last5=Tank|first5=D. C.|last6=Brockington|first6=S. F.|last7=Refulio-Rodriguez|first7=N. F.|last8=Walker|first8=J. B.|last9=Moore|first9=M. J.|last10=Carlsward|first10=B. S.|last11=Bell|first11=C. D.|last12=Latvis|first12=M.|last13=Crawley|first13=S.|last14=Black|first14=C.|last15=Diouf|first15=D.|last16=Xi|first16=Z.|last17=Rushworth|first17=C. A.|last18=Gitzendanner|first18=M. A.|last19=Sytsma|first19=K. J.|last20=Qiu|first20=Y.-L.|last21=Hilu|first21=K. W.|last22=Davis|first22=C. C.|last23=Sanderson|first23=M. J.|last24=Beaman|first24=R. S.|last25=Olmstead|first25=R. G.|last26=Judd|first26=W. S.|authorlink26=Walter Stephen Judd|last27=Donoghue|first27=M. J.|last28=Soltis|first28=P. S.|authorlink28 = Pamela S. Soltis|title=Angiosperm phylogeny: 17 genes, 640 taxa|url=http://www.amjbot.org/content/98/4/704.full|journal=[[American Journal of Botany]]|date=8 April 2011|volume=98|issue=4|pages=704–730|doi=10.3732/ajb.1000404|ref={{harvid|Soltis et al|2011}}|pmid=21613169}}
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| − | {{refend}}
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| − | {{NIE Poster|year=1905}}
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| − | {{monocotyledons}}
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| − | {{taxonbar}}
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| | [[Category:Monocots]] | | [[Category:Monocots]] |
| − | [[Category:Monocot genera]]
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| − | [[Category:Absinthe]]
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| − | [[Category:Edible plants]]
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| − | [[Category:Freshwater plants]]
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