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''Acorus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are flowering plants whose seeds contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but ...
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s. This genus was once placed within the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
(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 A sister is a woman or a girl who shares parents or a parent with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to ref ...
to all other monocots. Common names include calamus and sweet flag. The genus is native to
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 ...
and northern and eastern
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 naturalised in southern Asia 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 ...
from ancient cultivation. The known wild populations are
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
except for some
tetraploid Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
s in eastern Asia, while the cultivated plants are sterile triploids, probably of hybrid origin between the diploid and tetraploid forms.


Characteristics

The inconspicuous
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 are arranged on a lateral spadix (a thickened, fleshy axis). Unlike aroids, there is no
spathe In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
(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
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, ...
are linear with entire margin.


Taxonomy

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 Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...
. The family has recently been resurrected as molecular systematic studies have shown that ''Acorus'' is not closely related to Araceae or any other
monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are flowering plants whose seeds contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but ...
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 In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to all other monocots. This relationship is confirmed by more recent phylogenetic studies. Treatment in the
APG IV system The APG IV system of flowering plant classification is the fourth version of a modern, mostly molecular-based, system of plant taxonomy for flowering plants (angiosperms) being developed by the Angiosperm Phylogeny Group (APG). It was publish ...
is unchanged from APG III.


Species

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''). As of July 2014, the Kew Checklist accepts only 2 species, one of which has three accepted varieties: * '' Acorus calamus'' L. – common sweet flag; sterile triploid (3''n'' = 36); probably of cultivated origin. It is native to
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 ...
, temperate
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
and the
Himalaya The Himalayas, or Himalaya ( ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the Earth's highest peaks, including the highest, Mount Everest. More than 100 pea ...
s and southern
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 ...
, widely cultivated and naturalised elsewhere. **''Acorus calamus'' var. ''angustatus'' Besser - Siberia, China, Russian Far East, Japan, Korea, Mongolia, Himalayas, Indian Subcontinent, Indochina, Philippines, Indonesia **''Acorus calamus'' var. ''calamus'' - Siberia, Russian Far east, Mongolia, Manchuria, Korea, Himalayas; naturalized in Europe, North America, Java and New Guinea *'' Acorus americanus'' Raf. - Canada, northern United States, Buryatiya region of Russia * '' Acorus gramineus'' Sol. ex Aiton – Japanese sweet flag or grassy-leaved sweet flag; fertile diploid (2''n'' = 18); - China, Himalayas, Japan, Korea, Indochina, Philippines, Primorye ''Acorus'' from Europe,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
and Japan have been planted in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
.


Etymology

The name 'acorus' is derived from the Greek word 'acoron', a name used by
Dioscorides Pedanius Dioscorides (, ; 40–90 AD), "the father of pharmacognosy", was a Greek physician, pharmacologist, botanist, and author of (in the original , , both meaning "On Materia medica, Medical Material") , a 5-volume Greek encyclopedic phar ...
, which in turn was derived from 'coreon', meaning 'pupil', because it was used in
herbal medicine Herbal medicine (also called herbalism, phytomedicine or phytotherapy) is the study of pharmacognosy and the use of medicinal plants, which are a basis of traditional medicine. Scientific evidence for the effectiveness of many herbal treatments ...
as a treatment for
inflammation Inflammation (from ) is part of the biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or irritants. The five cardinal signs are heat, pain, redness, swelling, and loss of function (Latin ''calor'', '' ...
of the
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
.


Distribution and habitat

These plants are found in
wetlands A wetland is a distinct semi-aquatic ecosystem whose groundcovers are flooded or saturated in water, either permanently, for years or decades, or only seasonally. Flooding results in oxygen-poor ( anoxic) processes taking place, especially ...
, particularly marshes, where they spread by means of thick rhizomes. Like many other marsh plants, they depend upon
aerenchyma Aerenchyma or aeriferous parenchyma or lacunae, is a modification of the parenchyma to form a spongy tissue that creates spaces or air channels in the leaves, stems and roots of some plants, which allows exchange of gases between the shoot and ...
to transport oxygen to the rooting zone. They frequently occur on shorelines and floodplains where water levels fluctuate seasonally.


Ecology

The native North American species appears in many ecological studies. Compared to other species of wetland plants, they have relatively high competitive ability. Although many marsh plants accumulate large banks of buried seeds, seed banks of ''Acorus'' may not accumulate in some wetlands owing to low seed production. 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. 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''.


Toxicity

Products derived from ''Acorus calamus'' were banned in 1968 as food additives by the United States
Food and Drug Administration The United States Food and Drug Administration (FDA or US FDA) is a List of United States federal agencies, federal agency of the United States Department of Health and Human Services, Department of Health and Human Services. The FDA is respo ...
. The primary chemical responsible for the plant's toxic and carcinogenic effects is β-asarone. Confusion exists whether all strains of ''A. calamus'' contain this substance. Four varieties of ''A. calamus'' strains exist in nature: diploid, triploid, tetraploid and hexaploid. 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. 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. Research has consistently demonstrated that "β-asarone was not detectable in the North American spontaneous diploid ''Acorus'' 'calamus'' var. ''americanus''.


Uses

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 pests.


References


Bibliography

* 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.
Flora of North America: Acoraceae


in Stevens, P. F. (2001 onwards)
Angiosperm Phylogeny Website
Version 7, May 2006.
NCBI Taxonomy Browser


https://web.archive.org/web/20050209163227/http://delta-intkey.com/angio/www/acoracea.htm Acoraceae] in L. Watson and M.J. Dallwitz (1992 onwards)
The families of flowering plants:
descriptions, illustrations, identification, information retrieval''. Version: 27 April 2006. http://delta-intkey.com.
Taxonomy and distribution of Acorus in Maine



Phylogenetic analysis of rbcL sequences identifies Acorus calamus as the primal extant monocotyledon. Duvall 1993
*
Analysis of Acorus calamus Chloroplast Genome and Its Phylogenetic Implications Vadim V. Goremykin 2005
* * {{Authority control Acorales Monocot genera Absinthe Edible plants Freshwater plants