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    calamus has been dispersed by this means. The seeds arealso dispersed by water along stream margins.

    Acorus is found mainlyin the northern hemisphere, fromthe temperate zone to the subtropics and higher altitudes inthe tropics in Asia, in Europe and North America. A.calamus was widely dispersed in Europe and parts ofeastern North America by human agency due to its

    medicinal importance and later became naturalized inmany regions. Records from Celebes (Sulawesi), Philip-pines and New Guinea (including the Bismarck Archipe-lago) are not able to confirm if these are natural orintroduced and later naturalized; but A. calamus iscommon and widespread in the New Guinea highlands.

    Acorus is a herbaceous helophyte which forms densepopulations by strongly branching rhizomatous stems. Itgrows 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 inChina. The two species of Acorus are apparently not

    threatened and both are easy to grow; A. calamus especiallyis prone to naturalize easily and become quite weedy underfavourable conditions.

    Fossil History

    The family Acoraceae is documented by fossils from theEocene of North America; it was first described as Acorusheeri 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 wasdescribed from the Eocene (not Miocene as originallyreported) of Spitzbergen and for over a century wasthought to belong to Acorus, but new material and further

    studies showed that this fossil is identical with short shooof Nordienskioeldia borealis Heer (TrochodendralceaeAcoropsis eximia (Go pp.&Menge) Bogner (syn. Acoropsminor Conw.) is an infructescence from the Baltic amber oEocene age, but does not belong to the Acoraceae; it is member of the AraceaeMonsteroideae, tribe Monsterea

    Phylogeny

    Acorus was for a long time considered to be a member othe Araceae and it has only recently been removed fromthis family, although the family Acoraceae was describeas early as 1820. There are significant characters thadistinguish Acorus from the Araceae: presence of speciaethereal oil cells, absence of raphides, presence operisperm, cellular endosperm development, trichomeon the micropyle of the ovules, unifacial leaves, twseparate vascular systems in the peduncle and a few othe

    anatomical characters. DNA studies have shown thaAcorus is a sister taxon to all other monocots and somauthors believe that it is the most primitive living monocotoday.

    Further Reading

    Bogner J and Mayo SJ (1998) Acoraceae. In: Kabitzki K (ed.) Th

    Families and Genera of Vascular Plants 4, pp. 711. Berlin: Springe

    Engler A (1905) Araceae Pothoideae. In: Engler A (ed.) D

    Pflanzenreich 4, 23B, pp. 308313. Leipzig: W Englemann.

    Grayum MH (1987) A summary of evidence and arguments supportin

    the removal ofAcorus from the Araceae. Taxon 36: 723729.

    Mayo SJ, Bogner J and Boyce PC (1997) The Genera of Araceae. Roy

    Botanical Gardens, Kew, London.Petersen G (1989) Cytology and systematics of Araceae. Nordic Journ

    of Botany 9: 119166.

    Acorales (Sweet Flag)

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