''Allium'' is a genus of

monocotyledon Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

ous

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s that includes hundreds of species, including the cultivated

onion An onion (''Allium cepa'' L., from Latin ''cepa'' meaning "onion"), also known as the bulb onion or common onion, is a vegetable that is the most widely cultivated species of the genus ''Allium''. The shallot is a botanical variety of the onio ...

, garlic,

scallion Scallions (also known as spring onions or green onions) are vegetables derived from various species in the genus '' Allium''. Scallions generally have a milder taste than most onions and their close relatives include garlic, shallot, leek, c ...

shallot The shallot is a botanical variety (a cultivar) of the onion. Until 2010, the (French red) shallot was classified as a separate species, ''Allium ascalonicum''. The taxon was synonymized with '' Allium cepa'' (the common onion) in 2010, as the ...

, leek, and

chives Chives, scientific name ''Allium schoenoprasum'', is a species of flowering plant in the family Amaryllidaceae that produces edible leaves and flowers. Their close relatives include the common onions, garlic, shallot, leek, scallion, and ...

. The generic name ''Allium'' is the Latin word for garlic,Gledhill, David (2008). "The Names of Plants". Cambridge University Press. (hardback), (paperback). pp 43 and the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...

for the genus is '' Allium sativum'' which means "cultivated garlic".''Allium'' In: Index Nominum Genericorum. In:

Regnum Vegetabile The International Association for Plant Taxonomy (IAPT) is an organization established to promote an understanding of plant biodiversity, facilitate international communication of research between botanists, and oversee matters of uniformity and ...

(see ''External links'' below).

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...

first described the genus ''Allium'' in 1753. Some sources refer to Greek ἀλέω (aleo, to avoid) by reason of the smell of garlic. Various ''Allium'' have been cultivated from the earliest times, and about a dozen species are economically important as crops, or garden

vegetables Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems ...

, and an increasing number of species are important as ornamental plants. The decision to include a species in the genus ''Allium'' is

taxonomically In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...

difficult, and

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

boundaries are unclear. Estimates of the number of species are as low as 260, and as high as 979. ''Allium'' species occur in

temperate climate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...

s of the Northern Hemisphere, except for a few species occurring in Chile (such as '' A. juncifolium''), Brazil ('' A. sellovianum''), and tropical Africa ('' A. spathaceum''). They vary in height between 5 cm and 150 cm. The

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...

s form an

umbel In botany, an umbel is an inflorescence that consists of a number of short flower stalks (called pedicels) that spread from a common point, somewhat like umbrella ribs. The word was coined in botanical usage in the 1590s, from Latin ''umbella'' "p ...

at the top of a leafless stalk. The bulbs vary in size between species, from small (around 2–3 mm in diameter) to rather large (8–10 cm). Some species (such as Welsh onion '' A. fistulosum'' and

leeks The leek is a vegetable, a cultivar of ''Allium ampeloprasum'', the broadleaf wild leek ( syn. ''Allium porrum''). The edible part of the plant is a bundle of leaf sheaths that is sometimes erroneously called a stem or stalk. The genus ''Alliu ...

(''A. ampeloprasum'')) develop thickened leaf-bases rather than forming bulbs as such. Plants of the genus ''Allium'' produce chemical compounds, mostly derived from cysteine sulfoxides, that give them a characteristic onion or garlic taste and odor. Many are used as food plants, though not all members of the genus are equally flavorful. In most cases, both bulb and leaves are edible. The characteristic ''Allium'' flavor depends on the

sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ...

content of the soil the plant grows in. In the rare occurrence of sulfur-free growth conditions, all ''Allium'' species completely lose their usual pungency. In the APG III classification system, ''Allium'' is placed in the

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Amaryllidaceae, subfamily

Allioideae Allioideae is a subfamily of monocot flowering plants in the family Amaryllidaceae, order Asparagales. It was formerly treated as a separate family, Alliaceae. The subfamily name is derived from the generic name of the type genus, ''Allium''. ...

(formerly the family Alliaceae). In some of the older

classification systems Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organiza ...

, ''Allium'' was placed in

Liliaceae The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair a ...

. Molecular phylogenetic studies have shown this

circumscription Circumscription may refer to: *Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) *Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...

of Liliaceae is not monophyletic. ''Allium'' is one of about fifty-seven genera of flowering plants with more than 500 species. It is by far the largest genus in the Amaryllidaceae, and also in the Alliaceae in Biological classification, classification List of systems of plant taxonomy, systems in which that family is recognized as separate.Knud Rahn. 1998. "Alliaceae" pages 70-78. In: Klaus Kubitzki (editor). ''The Families and Genera of Vascular Plants'' volume III. Springer-Verlag: Berlin;Heidelberg, Germany.

Description

The genus ''Allium'' (alliums) is characterised by herbaceous geophyte perennials with true bulbs, some of which are borne on rhizomes, and an

or garlic odor and flavor. The bulbs are solitary or clustered and :wikt:tunicate, tunicate and the plants are perennialized by the bulbs reforming annually from the base of the old bulbs, or are produced on the ends of rhizomes or, in a few species, at the ends of stolons. A small number of species have tuberous roots. The bulbs' outer coats are commonly brown or grey, with a smooth texture, and are fibrous, or with cellular reticulation. The inner coats of the bulbs are membranous. Many alliums have basal leaves that commonly wither away from the tips downward before or while the plants flower, but some species have persistent foliage. Plants produce from one to 12 leaves, most species having linear, channeled or flat leaf blades. The leaf blades are straight or variously coiled, but some species have broad leaves, including ''Allium victorialis, A. victorialis'' and ''Allium tricoccum, A. tricoccum''. The leaves are Sessility (botany), sessile, and very rarely narrowed into a Petiole (botany), petiole. The flowers, which are produced on Scape (botany), scapes are erect or in some species pendent, having six petal-like tepals produced in two whorls. The flowers have one Gynoecium, style and six epipetalous stamens; the anthers and pollen can vary in color depending on the species. The Ovary (plants), ovaries are superior, and three-lobed with three locules. The fruits are capsules that open longitudinally along the capsule wall between the partitions of the locule. The seeds are black, and have a rounded shape. The wikt:terete, terete or flattened flowering scapes are normally persistent. The inflorescences are

s, in which the outside flowers bloom first and flowering progresses to the inside. Some species produce bulbils within the umbels, and in some species, such as ''Allium paradoxum'', the bulbils replace some or all the flowers. The umbels are subtended by noticeable spathe bracts, which are commonly fused and normally have around three veins. Some bulbous alliums increase by forming little bulbs or "offsets" around the old one, as well as by seed. Several species can form many bulbils in the flowerhead; in the so-called "tree onion" or Egyptian onion (''A.'' × ''proliferum'') the bulbils are few, but large enough to be pickling, pickled. Many of the species of ''Allium'' have been used as food items throughout their ranges. There are several poisonous species that are somewhat similar in appearance (e.g. in North America, death camas, ''Toxicoscordion venenosum''), but none of these has the distinctive scent of onions or garlic.

Taxonomy

With over 850 species ''Allium'' is the sole genus in the Allieae, one of four Tribe (biology), tribes of subfamily

( Amaryllidaceae). New species continue to be described and ''Allium'' is one of the largest monocotyledonous genera, but the precise Plant taxonomy, taxonomy of ''Allium'' is poorly understood, with incorrect descriptions being widespread. The difficulties arise from the fact that the genus displays considerable polymorphism and has adapted to a wide variety of habitats. Furthermore, traditional classifications had been based on homoplasious characteristics (the independent evolution of similar features in species of different lineages). However, the genus has been shown to be monophyletic, containing three major clades, although some proposed subgenera are not. Some progress is being made using molecular phylogenetic methods, and the internal transcribed spacer (ITS) region, including the 5.8S ribosomal DNA, rDNA and the two spacer DNA, spacers ITS1 and ITS2, is one of the more commonly used markers in the study of the differentiation of the ''Allium'' species. ''Allium'' includes a number of taxonomic groupings previously considered separate genera (''Caloscordum'' Herb., ''Milula'' Prain and ''Nectaroscordum'' Lindl.) ''Allium spicatum'' had been treated by many authors as ''Milula spicata'', the only species in the Monotypic taxon, monospecific genus ''Milula''. In 2000, it was shown to be Paraphyly, embedded in ''Allium''.

Phylogeny

History

When Linnaeus formerly described the genus ''Allium'' in his ''Species Plantarum'' (1753), there were thirty species with this name. He placed ''Allium'' in a grouping he referred to as ''Hexandria monogynia'' (i.e. six stamens and one pistil) containing 51 genera in all.

Subdivision

Linnaeus originally grouped his 30 species into three alliances, ''e.g.'' ''Foliis caulinis planis''. Since then, many attempts have been made to divide the growing number of recognised species into infrageneric subgroupings, initially as sections, and then as subgenera further divided into sections. For a brief history, see Li ''et al.'' (2010) The modern era of phylogenetic analysis dates to 1996. In 2006 Friesen, Fritsch, and Blattner described a new classification with 15 subgenera, 56 section (botany), sections, and about 780 species based on the cell nucleus, nuclear Ribosome, ribosomal gene internal transcribed spacers. Some of the subgenera correspond to the once separate genera (''Caloscordum'', ''Milula'', ''Nectaroscordum'') included in the Gilliesieae. The terminology has varied with some authors subdividing subgenera into Sections and others Alliances. The term Alliance has also been used for subgroupings within species, ''e.g.'' ''Allium nigrum'', and for subsections. Subsequent molecular phylogenetic studies have shown the 2006 classification is a considerable improvement over previous classifications, but some of its subgenera and sections are probably not monophyletic. Meanwhile, the number of new species continued to increase, reaching 800 by 2009, and the pace of discovery has not decreased. Detailed studies have focused on a number of subgenera, including ''Amerallium''. ''Amerallium'' is strongly Resampling (statistics), supported as monophyletic. Subgenus ''Melanocrommyum'' has also been the subject of considerable study (see below), while work on subgenus ''Allium'' has focussed on section ''Allium'', including ''Allium ampeloprasum'', although Sample (material), sampling was not sufficient to test the monophyly of the section. The major evolutionary lineages or lines correspond to the three major clades. Line one (the oldest) with three subgenera is predominantly bulbous, the second, with five subgenera and the third with seven subgenera contain both bulbous and rhizomatous taxa.

Evolutionary lines and subgenera

The three evolutionary lineages and 15 subgenera represent the classification scheme of Friesen ''et al.'' (2006) and Li (2010). (number of sections/number of species) * First evolutionary line *# ''Nectaroscordum'' (Lindl.) Asch. et Graebn Type: ''Allium siculum'' (1/3) Mediterranean bells, Sicilian honey garlic *# ''Microscordum'' (Maxim.) N. Friesen Type: ''Allium monanthum'' (1/1) *# ''Amerallium'' Traub Type: ''Allium canadense'' (12/135) * Second evolutionary line *# ''Caloscordum'' (Herb.) R. M. Fritsch Type: ''Allium neriniflorum'' (1/3) *# ''Anguinum'' (G. Don ex Koch) N. Friesen Type: ''Allium victorialis'' (1/12) *# ''Porphyroprason'' (Ekberg) R. M. Fritsch Type: ''Allium oreophilum'' (1/1) *# ''Vvedenskya'' (Kamelin) R. M. Fritsch Type: ''Allium kujukense'' (1/1) *# ''Melanocrommyum'' (Philip Barker-Webb, Webb et Berthel.) Georges Rouy, Rouy Type: ''Allium nigrum'' (20/160) * Third evolutionary line *# ''Butomissa'' (Salisb.) N. Friesen Type: ''Allium ramosum'' (2/4) fragrant garlic *# ''Cyathophora'' R. M. Fritsch Type: ''Allium cyathophorum'' (3/5) *# ''Rhizirideum'' (G. Don ex Koch) Wendelbo s.s Type: ''Allium senescens'' (5/37) *# ''Allium'' Carl Linnaeus, L. Type: '' Allium sativum'' (15/300) garlic *# ''Reticulatobulbosa'' (Kamelin) N. Friesen Type: ''Allium lineare'' (5/80) *# ''Polyprason'' Radic Type: ''Allium moschatum'' (4/50) *# ''Cepa'' (Mill.) Radic ́ Type: ''Allium cepa'' (5/30) onion, garden onion, bulb onion, common onion

First evolutionary line

Although this lineage consists of three subgenera, nearly all the species are attributed to subgenus ''Amerallium'', the third largest subgenus of ''Allium''. The lineage is considered to represent the most ancient line within ''Allium'', and to be the only lineage that is purely bulbous, the other two having both bulbous and rhizomatous taxa. Within the lineage ''Amerallium'' is a sister group to the other two subgenera (''Microscordum''+''Nectaroscordum'').

Second evolutionary line

Nearly all the species in this lineage of five subgenera are accounted for by subgenus ''Melanocrommyum'', which is most closely associated with subgenera ''Vvedenskya'' and ''Porphyroprason'', phylogenetically. These three genera are late-branching whereas the remaining two subgenera, ''Caloscordum'' and ''Anguinum'', are early branching.

Third evolutionary line

The third evolutionary line contains the greatest number of sections (seven), and also the largest subgenus of the genus ''Allium'': subgenus ''Allium'', which includes the type species of the genus, ''Allium sativum''. This subgenus also contains the majority of the species in its lineage. Within the lineage, the phylogeny is complex. Two small subgenera, ''Butomissa'' and ''Cyathophora'' form a sister clade to the remaining five subgenera, with ''Butomissa'' as the first branching group. Amongst the remaining five subgenera, ''Rhizirideum'' forms a medium-sized subgenus that is the sister to the other four, larger, subgenera. This line may not be monophyletic.

Distribution and habitat

The majority of ''Allium'' species are native to the Northern Hemisphere, being spread throughout the holarctic region, from dry subtropics to the Boreal ecosystem, boreal zone, predominantly in Asia. Of the latter, 138 species occur in China, about a sixth of all ''Allium'' species, representing five subgenera. A few species are native to Africa and Central and South America. A single known exception, ''Allium dregeanum'' occurs in the Southern Hemisphere (South Africa). There are two centres of diversity, a major one from the Mediterranean Basin to Central Asia and Pakistan, while a minor one is found in western North America. The genus is especially diverse in the eastern Mediterranean.

Ecology

Species grow in various conditions from dry, well-drained mineral-based soils to moist, organic soils; most grow in sunny locations, but a number also grow in forests (e.g., ''Allium ursinum, A. ursinum''), or even in swamps or water. Various ''Allium'' species are used as food plants by the larvae of the leek moth and onion fly as well as some Lepidoptera including cabbage moth, Common swift (moth), common swift moth (recorded on garlic), garden dart moth, large yellow underwing moth, Nutmeg (moth), nutmeg moth, setaceous Hebrew character moth, turnip moth and ''Schinia, Schinia rosea'', a moth that feeds exclusively on ''Allium'' species.

Genetics

The genus Allium has very large variation between species in their genome size that is not accompanied by changes in ploidy level. This remarkable variation was noted in the discussion of the evolution of junk DNA and resulted in the Onion Test, a "reality check for anyone who thinks they have come up with a universal function for junk DNA". Genome sizes vary between 7.5 Gb in ''Allium schoenoprasum, A. schoenoprasum'' and 30.9 Gb in ''Allium ursinum, A. ursinum'', both of which are diploid.

Cultivation

Many ''Allium'' species have been harvested through human history, but only about a dozen are still economically important today as crops or garden vegetables.

Ornamental

Many ''Allium'' species and Hybrid (biology), hybrids are Horticulture, cultivated as Ornamental plant, ornamentals.Anthony Huxley, Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. (set). These include ''Allium cristophii, A. cristophii'' and ''Allium giganteum, A. giganteum'', which are used as herbaceous border, border plants for their ornamental flowers, and their "architectural" qualities.Brickell, Christopher (Editor-in-chief),''The Royal Horticultural Society A–Z Encyclopedia of Garden Plants'', p.95, Dorling Kindersley, London, 1996, Several hybrids have been bred, or selected, with rich purple flowers. ''Allium hollandicum, A. hollandicum'' 'Purple Sensation' is one of the most popular and has been given an Award of Garden Merit (H4).''RHS Plant Finder 2009–2010'', p68, Dorling Kindersley, London, 2009, These ornamental onions produce spherical umbels on single stalks in spring and summer, in a wide variety of sizes and colours, ranging from white (''Allium'' 'Mont Blanc'), blue (''Allium caeruleum, A. caeruleum''), to yellow (''Allium flavum, A. flavum'') and purple (''A. giganteum''). By contrast, other species (such as invasive ''Allium triquetrum, A. triquetrum'' and ''Allium ursinum, A. ursinum'') can become troublesome garden weeds.Lloyd, Christopher & Rice, Graham, (1991) ''Garden Flowers From Seed'', p45, Viking, The following cultivars, of uncertain or mixed parentage, have gained the Royal Horticultural Society's Award of Garden Merit: *’Ambassador’ *’Beau Regard’ *’Gladiator’ *’Globemaster’ *'Michael H' Hoog' (Allium rosenorum, ''A. rosenorum'') *’Round 'n' Purple’ *'Universe'

Toxicity

Dogs and cats are very susceptible to poisoning after the consumption of certain species.Cope RB. Toxicology Brief: Allium species poisoning in dogs and cats. Veterinary Medicine 2005
/ref>

Uses

The genus includes many economically important species. These include

s (''A. cepa''), French shallots (''A. oschaninii''),

(''A. ampeloprasum''), garlic (''A. sativum''), and herbs such as scallions (various ''Allium'' species) and

(''A. schoenoprasum''). Some have been used as traditional medicines. This genus also includes species that are abundantly gathered from the wild such as wild garlic (''Allium ursinum'') and ramps (''Allium tricoccum'').

References

Bibliography

Books

* * * * * * *

Chapters

* , in

Articles

* * * * * * * * * * * * * * * * [In Chinese.] * *
A. Samoylov, N. Friesen, S. Pollner, P. Hanelt. Use of chloroplast DNA polymorphisms for the phylogenetic study of ''Allium'' subgenus ''Amerallium'' and subgenus ''Bromatorrhiza'' (Alliaceae) II. ''Feddes Repertorium'' Volume 110 Issue 1–2, Pages 103–109, 1999
* * * * *

Websites

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DutchGrown: Alliums

Pacific Bulb Society: Allium
*
Pacific Bulb Society: Rhizomatous Alliums

External links

*
Allium
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Index Nominum Genericorum
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Bloomsta.com Florist Community

Reinhard M. Fritsch. Checklist of ornamental ''Allium'' species and cultivars currently offered in the trade. 2015
{{Authority control Allium, Onions, *Allium Amaryllidaceae genera Bulbous plants Taxa named by Carl Linnaeus