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Acorales (Sweet Flag)

Josef Bogner, Munich Botanical Gardens, Munich, Germany


The Acorales contains only one family, the Acoraceae, with only one genus, Acorus, with two species and a few varieties. The species Acorus calamus and A. gramineus are both medicinally and horticulturally of importance.

Introductory article
Article Contents
. Medicinal and Horticultural Importance . Morphology . Biogeography . Fossil History . Phylogeny

Medicinal and Horticultural Importance


The ancient Egyptians, Greeks and Romans used the dried rhizomes (rhizoma calami) of Acorus calamus for medicaments and they are still in use today. A tea from chopped rhizomes is made to treat stomach ailments and to produce a liqueur, and also formerly for perfume. The bitter rhizomes of A. gramineus have been used medicinally too. Substances extracted from the rhizome have been proved to have sedative, analgesic, insecticidal and antibacterial properties. Important chemical compounds from the rhizomes are acorin and polyphenols and varying concentrations of ethereal oils depending on the geographical source and cytotype of the plants. The composition of ethereal oils in the leaves of A. calamus diers from that of the rhizome, but there is a correlation with the cytotype. Bruised leaves give o a strong sweet odour and that is why it is called sweet ag in English. Both species are also cultivated as ornamentals and are suitable for wet places, such as bog gardens, margins of ponds, aquatic gardens etc.; both are hardy outside in temperate regions. Cultivars with variegated leaves are quite popular in horticulture. Acorus gramineus var. pusillus is sometimes grown as an aquarium plant. oblongobovoid with thin, leathery pericarp, whitish with brownish stigma remnant when fresh, soon drying to straw-brown, enclosed by tepals, 15 (9)-seeded; seeds oblong to ellipsoid; with perisperm and abundant endosperm; testa light brown, foveolate or smooth; embryo axile, cylindric. Acorus calamus L.: leaves 60150 cm long and 0.7 2.0 cm wide, with a prominent midrib; spadix 410 cm long and 0.61.5 cm in diameter; owers greenish-yellow. Acorus gramineus Solander: leaves 3050 cm long and (0.3) 0.51 cm wide, without prominent midrib; spadix 5 10 cm long and 0.30.4 cm in diameter; owers greenishyellow; var. pusillus (Sieb.) Engl. with leaves only about 10 cm long. Vessels are present in the roots and the rhizome. The leaves have air cavities beneath a chlorenchyma. The leaf venation is strictly parallel with minor transverse veins joining the main, longitudinally oriented system; it represents a typical monocot venation type. Raphides are lacking and laticifers are also absent. The ethereal oils occur in idioblasts. The formation of the endosperm in the seeds is cellular; the perisperm consists of a single cell layer only.

Morphology
Acorus L.: rhizomatous herbs; rhizome much branched, lacunose, with specialized oil cells; leaves distichous, unifacial, ensiform, not dierentiated into petiole and blade, venation parallel; inorescence solitary, borne laterally on leaf-like axis, the so-called spathe longer than the spadix, erect, and appearing as an extension of the leaflike peduncle; spadix more or less conical or slender and tail-like, densely owered; owers bractless, bisexual, trimerous, perigoniate; tepals in two whorls of three, thin, fornicate; stamens in two whorls of three, laments linear elongate, anthers introrse, thecae roundedelliptical, opening by longitudinal slit; pollen grains ellipsoid, sulcate, exine more or less foveolate, otherwise psilate; gynoecium obconic-oblong, two-to three-locular, placenta apical, ovules several in each locule, pendent, atropous (orthotropous), both integuments bearing trichomes, inner integument longer than outer; stigma small, sessile; berry

Biogeography
Acorus gramineus, found in Asia, occurs only in diploid populations (2n 5 24). A. calamus is di-, tri- and tetraploid; diploids with chromosomes 2n 5 24 are known to grow naturally in North America, tetraploids (2n 5 48) are only known from Asia and the triploids (2n 5 36) are typical of the European A. calamus, from the Near East, India (Himalaya region) and in eastern North America, where they are very probably introductions from Europe by early settlers. This is why the European A. calamus never sets fruits. The pollination agency of Acorus is not known, but entomophily is very likely since the pollen is sticky. Hand pollinated diploid plants set 100% seeds, but otherwise free growing plants set fruits only rarely and visiting insects are hardly ever seen in Europe. Pieces of rhizome are easily dispersed by water along rivers and creeks. In particular, the sterile triploid Acorus
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ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

Acorales (Sweet Flag)

calamus has been dispersed by this means. The seeds are also dispersed by water along stream margins. Acorus is found mainly in the northern hemisphere, from the temperate zone to the subtropics and higher altitudes in the tropics in Asia, in Europe and North America. A. calamus was widely dispersed in Europe and parts of eastern North America by human agency due to its medicinal importance and later became naturalized in many regions. Records from Celebes (Sulawesi), Philippines and New Guinea (including the Bismarck Archipelago) are not able to conrm if these are natural or introduced and later naturalized; but A. calamus is common and widespread in the New Guinea highlands. Acorus is a herbaceous helophyte which forms dense populations by strongly branching rhizomatous stems. It grows along margins of rivers, streams, ponds, lakes, ditches, standing or slow running water or swampy areas. Acorus is found from sea level to 1100 m in central Europe (nearly the same altitude in Turkey) and up to 2600 m in China. The two species of Acorus are apparently not threatened and both are easy to grow; A. calamus especially is prone to naturalize easily and become quite weedy under favourable conditions.

studies showed that this fossil is identical with short shoots of Nordienskioeldia borealis Heer (Trochodendralceae). Acoropsis eximia (Go pp.&Menge) Bogner (syn. Acoropsis minor Conw.) is an infructescence from the Baltic amber of Eocene age, but does not belong to the Acoraceae; it is a member of the AraceaeMonsteroideae, tribe Monstereae.

Phylogeny
Acorus was for a long time considered to be a member of the Araceae and it has only recently been removed from this family, although the family Acoraceae was described as early as 1820. There are signicant characters that distinguish Acorus from the Araceae: presence of special ethereal oil cells, absence of raphides, presence of perisperm, cellular endosperm development, trichomes on the micropyle of the ovules, unifacial leaves, two separate vascular systems in the peduncle and a few other anatomical characters. DNA studies have shown that Acorus is a sister taxon to all other monocots and some authors believe that it is the most primitive living monocot today.

Fossil History
The family Acoraceae is documented by fossils from the Eocene of North America; it was rst described as Acorus heeri by Berry and later closely studied by Crepet (1977) and transferred to its own genus as Acorites heeri (Berry) Crepet. The fossil Acorus brachystachys Heer was described from the Eocene (not Miocene as originally reported) of Spitzbergen and for over a century was thought to belong to Acorus, but new material and further

Further Reading
Bogner J and Mayo SJ (1998) Acoraceae. In: Kabitzki K (ed.) The Families and Genera of Vascular Plants 4, pp. 711. Berlin: Springer. Engler A (1905) Araceae Pothoideae. In: Engler A (ed.) Das Panzenreich 4, 23B, pp. 308313. Leipzig: W Englemann. Grayum MH (1987) A summary of evidence and arguments supporting the removal of Acorus from the Araceae. Taxon 36: 723729. Mayo SJ, Bogner J and Boyce PC (1997) The Genera of Araceae. Royal Botanical Gardens, Kew, London. Petersen G (1989) Cytology and systematics of Araceae. Nordic Journal of Botany 9: 119166.

ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net

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