Wheat taxonomy
Encyclopedia
During 10,000 years of cultivation, numerous forms of wheat
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

 have evolved under human selection. This diversity has led to much confusion in the naming of wheats. This article explains how genetic and morphological characteristics of wheat influence its classification, and gives the most common botanical names of wheat in current use (see Table of wheat species). Information on the cultivation and uses of wheat is at the main wheat
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

 page.

Aegilops and Triticum

The genus Triticum includes the wild and domesticated species usually thought of as wheat.

In the 1950s growing awareness of the genetic similarity (including some shared genones) of the wild goatgrasses (Aegilops
Aegilops
Aegilops is a genus of plants generally known as goatgrasses and belonging to the grass family, Poaceae. There are about 23 species and numerous sub species in the genus. Various members of the genus are classed as agricultural weeds. Growing through the winter, they resemble winter wheat...

) led some botanists to amalgamate Aegilops and Triticum as one genus, Triticum. This approach is still followed by some (mainly geneticists), but has not been widely adopted by taxonomists. Aegilops is morphologically highly distinct from Triticum, with rounded glumes rather than keeled glumes.

Aegilops is important in wheat evolution because of its role in two important hybridisation events. Wild emmer (T. dicoccoides and T. araraticum) resulted from the hybridisation of a wild wheat, T. urartu, and an as yet unidentified goatgrass, probably similar to Ae. speltoides. Hexaploid wheats (e.g. T. aestivum and T. spelta) are the result of a hybridisation between a domesticated tetraploid wheat, probably T. dicoccum or T. durum, and another goatgrass, Ae. tauschii (also known as Ae. squarrosa).

Early taxonomy

Botanists of the classical period, such as Columella
Columella
Lucius Junius Moderatus Columella is the most important writer on agriculture of the Roman empire. Little is known of his life. He was probably born in Gades , possibly of Roman parents. After a career in the army , he took up farming...

, and in sixteenth and seventeenth century herbals, divided wheats into two groups, Triticum corresponding to free-threshing wheats, and Zea corresponding to hulled ('spelt') wheats.

Carl Linnaeus recognised five species, all domesticated:
  • T. aestivum Bearded spring wheat
  • T. hybernum Beardless winter wheat
  • T. turgidum Rivet wheat
  • T. spelta Spelt wheat
  • T. monococcum Einkorn wheat


Later classifications added to the number of species described, but continued to give species status to relatively minor variants, such as winter vs. spring forms. The wild wheats were not described until the mid-19th century because of the poor state of botanical exploration in the Near East, where they grow.

The development of a modern classification depended on the discovery, in the 1920s, that wheat was divided into 3 ploidy levels.

Ploidy level

As with many grasses, polyploidy
Polyploidy
Polyploid is a term used to describe cells and organisms containing more than two paired sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common...

 is common in wheat. Some wheats are not polyploid. There are two wild diploid wheats, T. boeoticum and T. urartu. T. boeoticum is the wild ancestor of domesticated einkorn, T. monococcum. Cells of the diploid wheats each contain 2 complements of 7 chromosomes, one from the mother and one from the father (2n=2x=14, where 2n is the number of chromosomes in each somatic cell, and x is the basic chromosome number).

The polyploid wheats are tetraploid (4 sets of chromosomes, 2n=4x=28), or hexaploid (6 sets of chromosomes, 2n=6x=42). The tetraploid wild wheats are wild emmer, T. dicoccoides, and T.araraticum. Wild emmer is the ancestor of all the domesticated tetraploid wheats, with one exception: T. araraticum is the wild ancestor of T. timopheevi.

There are no wild hexaploid wheats, although feral forms of common wheat
Common wheat
Common wheat, Triticum aestivum, is a cultivated wheat species.-Nomenclature and taxonomy of the variety and its cultivars:-Evolution:...

 are sometimes found. Hexaploid wheats evolved under domestication. Genetic analysis has shown that the original hexaploid wheats were the result of a cross between a tetraploid domesticated wheat, such as T. dicoccum or T. durum, and a wild goatgrass, Ae. tauschii.

Polyploidy is important to wheat classification for three reasons:
  • Wheats within one ploidy level will be more closely related to each other.
  • Ploidy level influences some plant characteristics. For example, higher levels of ploidy tend to be linked to larger cell size.
  • Polyploidy brings new genomes into a species. For example, Aegilops tauschii brought the D genome into hexaploid wheats, with enhanced cold-hardiness and some distinctive morphological features.

Genome

Observation of chromosome behaviour during meiosis
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....

, and the results of hybridisation experiments, have shown that grass genomes (complete complements of genetic matter) can be grouped into distinctive types. Each type has been given a name, e.g. B or D. Grasses sharing the same genome will be more-or-less interfertile, and might be treated by botanists as one species. Identification of genome types is obviously a valuable tool in investigating hybridisation. For example, if two diploid plants hybridise to form a new polyploid form (an allopolyploid), the two original genomes will be present in the new form. Many thousands of years after the original hybridisation event, identification of the component genomes will allow identification of the original parent species.

In Triticum, five genomes, all originally found in diploid species, have been identified:
  • Am - present in wild einkorn (T. boeoticum).
  • Au - present in T. urartu (closely related to T. boeoticum but not interfertile).
  • B - present in most tetraploid wheats. Source not identified, but similar to Ae. speltoides.
  • G - present in timopheevi group of wheats. Source not identified, but similar to Ae. speltoides.
  • D - present in Ae. squarrosa, and thus in all hexaploid wheats.


The genetic approach to wheat taxonomy (see below)takes the genome composition as defining each species. As there are five known combinations in Triticum this translates into five super species:
  • Am T. monococcum
  • Au T. urartu
  • BAu T. turgidum
  • GAu T. timopheevi
  • BAuD, T. aestivum

Domestication

There are four wild species, all growing in rocky habitats in the fertile crescent
Fertile Crescent
The Fertile Crescent, nicknamed "The Cradle of Civilization" for the fact the first civilizations started there, is a crescent-shaped region containing the comparatively moist and fertile land of otherwise arid and semi-arid Western Asia. The term was first used by University of Chicago...

 of the Near East
Near East
The Near East is a geographical term that covers different countries for geographers, archeologists, and historians, on the one hand, and for political scientists, economists, and journalists, on the other...

. All the other species are domesticated
Domestication
Domestication or taming is the process whereby a population of animals or plants, through a process of selection, becomes accustomed to human provision and control. In the Convention on Biological Diversity a domesticated species is defined as a 'species in which the evolutionary process has been...

. Although relatively few genes control domestication, and wild and domesticated forms are interfertile, wild and domesticated wheats occupy entirely separate habitats. Traditional classification gives more weight to domesticated status.

Hulled vs. Free-threshing

All wild wheats are hulled: they have tough glumes (husks) that tightly enclose the grains. Each package of glumes, lemma and palaea, and grain(s) is known as a spikelet. At maturity the rachis (central stalk of the cereal ear) disarticulates, allowing the spikelets to disperse.

The first domesticated wheats, einkorn and emmer, were hulled like their wild ancestors, but with rachises that (while not entirely tough) did not disarticulate at maturity. During the PPNB period, at about 8000 BC, free-threshing forms of wheat evolved, with light glumes and fully tough rachis.

Hulled or free-threshing status is important in traditional classification because the different forms are usually grown separately, and have very different post-harvesting processing. Hulled wheats need substantial extra pounding or milling to remove the tough glumes.

For more information, see Wheat: Hulled vs. free-threshing wheat

Morphology

In addition to hulled/free-threshing status, other morphological criteria, e.g. spike laxness or glume wingedness, are important in defining wheat forms. Some of these are covered in the individual species accounts linked from this page, but printed Floras must be consulted for full descriptions and identification keys.

Traditional vs. genetic classifications

Although the range of recognised types of wheat has been reasonably stable since the 1930s, there are now sharply differing views as to whether these should be recognised at species level (traditional approach) or at subspecific level (genetic approach). The first advocate of the genetic approach was Bowden, in a 1959 classification (now historic rather than current)http://www.k-state.edu/wgrc/Taxonomy/taxbow.html. He, and subsequent proponents (usually geneticists), argued that forms that were interfertile should be treated as one species (the biological species concept). Thus emmer and hard wheat should both be treated as subspecies (or at other infraspecific ranks) of a single tetraploid species defined by the genome BAu. Van Slageren's 1994 classificationhttp://www.k-state.edu/wgrc/Taxonomy/taxvsltr.html is probably the most widely used genetic-based classification at present.

Users of traditional classifications give more weight to the separate habitats of the traditional species, which means that species that could hybridise do not, and to morphological characters. There are also pragmatic arguments for this type of classification: it means that most species can be described in Latin binomials, e.g. Triticum aestivum, rather than the trinomials necessary in the genetic system, e.g. Triticum aestivum subsp. aestivum. Both approaches are widely used.

Infraspecific classification

In the nineteenth century, elaborate schemes of classification were developed in which wheat ears were classified to botanical variety
Variety (biology)
In botanical nomenclature, variety is a taxonomic rank below that of species: as such, it gets a three-part infraspecific name....

 on the basis of morphological criteria such as glume hairiness and colour or grain colour. These variety names are now largely abandoned, but are still sometimes used for distinctive types of wheat such as miracle wheat, a form of T. turgidum with branched ears, known as T. turgidum L. var. mirabile Körn.

The term cultivar (abbreviated as cv.)is often confused with species or domesticate. In fact, it has a precise meaning in botany: it is the term for a distinct population of a crop, usually commercial and resulting from deliberate plant-breeding. Cultivar names are always capitalised, often placed between apostrophes, and not italicised. An example of a cultivar name is T. aestivum cv. 'Pioneer 2163'. A cultivar is often referred to by farmers as a variety, but this is best avoided in print, because of the risk of confusion with botanical varieties.

Advice for users

Anyone wishing to use a botanical name for wheat is best advised to follow an existing classification, such as those listed as current at the Wheat Classification Tables Sitehttps://www.ksu.edu/wgrc/Taxonomy/taxintro.html. The classifications given in the following table are among those suitable for use. If a genetic classification is favoured, the GRIN classification is comprehensive, based on van Slageren's work but with some extra taxa recognised. If the traditional classification is favoured, Dorofeev's work is a comprehensive scheme that meshes well with other less complete treatments.Wikipedia's wheat pages generally follow a version of the Dorofeev scheme - see the taxobox
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

 on the Wheat page.

The most critical point is that different taxonomic schemes should not be mixed in one context. In a given article, book or web page, only one scheme should be used at a time. Otherwise, it will be unclear to others how the botanical name is being used.

Table of wheat species

Wheat taxonomy - two schemes
Common name Genome(s) Genetic (GRIN Taxonomy for Plants http://www.ars-grin.gov/cgi-bin/npgs/html/index.pl) Traditional (Dorofeev et al. 1979 https://www.ksu.edu/wgrc/Taxonomy/taxdor.html)
Diploid (2x), Wild, Hulled
Wild einkorn Am Triticum monococcum L. subsp. aegilopoides (Link
Johann Heinrich Friedrich Link
Johann Heinrich Friedrich Link was a German naturalist and botanist.Link was born at Hildesheim as a son of the minister August Heinrich Link , who taught him the love for nature through collection of 'natural objects'...

) Thell.
Albert Thellung
-Works:*Illustrierte Flora von Mittel-Europa 1906*Die Gattung Lepidium R. Br. : eine monographische Studie 1906 *Die Entstehung der Kulturpflanzen 1930 *Flora der Schweiz - Exkursionsflora, 1923...

Triticum boeoticum Boiss.
Au Triticum urartu Tumanian ex Gandilyan Triticum urartu Tumanian ex Gandilyan
Diploid (2x), Domesticated, Hulled
Einkorn Am Triticum monococcum L. subsp. monococcum Triticum monococcum L.
Tetraploid (4x), Wild, Hulled
Wild emmer BAu Triticum turgidum L. subsp. dicoccoides (Korn. ex Asch. & Graebn.) Thell.
Albert Thellung
-Works:*Illustrierte Flora von Mittel-Europa 1906*Die Gattung Lepidium R. Br. : eine monographische Studie 1906 *Die Entstehung der Kulturpflanzen 1930 *Flora der Schweiz - Exkursionsflora, 1923...

Triticum dicoccoides
Emmer
Emmer wheat , also known as farro especially in Italy, is a low yielding, awned wheat. It was one of the first crops domesticated in the Near East...

(Körn. ex Asch. & Graebner) Schweinf.
Tetraploid (4x), Domesticated, Hulled
Emmer BAu Triticum turgidum L. subsp. dicoccum (Schrank
Franz Paula von Schrank
Franz von Paula Schrank was a German botanist and entomologist.Schrank was the first director of the botanical gardens in Munich from 1809 to 1832.Shrank was the first author to use the genus name Triops...

 ex Schübl.
Gustav Schübler
Gustav Schübler was a German naturalist, and the founder of applied meteorology in Germany.In 1817 Schübler became professor of botany, natural history and agricultural chemistry at the University of Tübingen, Germany...

) Thell.
Albert Thellung
-Works:*Illustrierte Flora von Mittel-Europa 1906*Die Gattung Lepidium R. Br. : eine monographische Studie 1906 *Die Entstehung der Kulturpflanzen 1930 *Flora der Schweiz - Exkursionsflora, 1923...

Triticum dicoccum
Emmer
Emmer wheat , also known as farro especially in Italy, is a low yielding, awned wheat. It was one of the first crops domesticated in the Near East...

Schrank
Franz Paula von Schrank
Franz von Paula Schrank was a German botanist and entomologist.Schrank was the first director of the botanical gardens in Munich from 1809 to 1832.Shrank was the first author to use the genus name Triops...

 ex Schübler
Gustav Schübler
Gustav Schübler was a German naturalist, and the founder of applied meteorology in Germany.In 1817 Schübler became professor of botany, natural history and agricultural chemistry at the University of Tübingen, Germany...

BAu Triticum ispahanicum Heslot Triticum ispahanicum Heslot
BAu Triticum turgidum L. subsp. paleocolchicum Á.
Áskell Löve
Áskell Löve was an Icelandic systematic botanist, particularly active in the Arctic.Löve was born in Reykjavik. He studied botany at Lund University, Sweden, from 1937. He received his PhD in 1942 in botany and a D.Sc. degree in genetics the year after...

 & D. Löve
Doris Löve
Doris Benta Maria Löve, née Wahlén was a Swedish systematic botanist, particularly active in the Arctic.Doris Löve was born in Kristianstad, Sweden. She studied botany at Lund University from 1937. She married her fellow student and colleague, the Icelander Áskell Löve. She received her PhD in...

Triticum karamyschevii Nevski
Tetraploid (4x), Domesticated, Free-threshing
Durum or macaroni wheat BAu Triticum turgidum L. subsp. durum (Desf.) Husn. Triticum durum
Durum
Durum wheat or macaroni wheat is the only tetraploid species of wheat of commercial importance that is widely cultivated today...

Desf.
Rivet or cone wheat BAu Triticum turgidum L. subsp. turgidum Triticum turgidum L.
Polish wheat BAu Triticum turgidum L. subsp. polonicum (L.) Thell.
Albert Thellung
-Works:*Illustrierte Flora von Mittel-Europa 1906*Die Gattung Lepidium R. Br. : eine monographische Studie 1906 *Die Entstehung der Kulturpflanzen 1930 *Flora der Schweiz - Exkursionsflora, 1923...

Triticum polonicum
Triticum polonicum
Triticum polonicum, also known as polish wheat, is an ordinary variant of wheat. It's a tetraploid species with 28 chromosomes. It can be found in small areas of the Mediterranean region, Ethiopia, Russia and in other regions of Asia. It was first described by Carl Linnaeus in 1762....

L.
Khorasan wheat BAu Triticum turgidum L. subsp. turanicum (Jakubz.) Á.
Áskell Löve
Áskell Löve was an Icelandic systematic botanist, particularly active in the Arctic.Löve was born in Reykjavik. He studied botany at Lund University, Sweden, from 1937. He received his PhD in 1942 in botany and a D.Sc. degree in genetics the year after...

 & D. Löve
Doris Löve
Doris Benta Maria Löve, née Wahlén was a Swedish systematic botanist, particularly active in the Arctic.Doris Löve was born in Kristianstad, Sweden. She studied botany at Lund University from 1937. She married her fellow student and colleague, the Icelander Áskell Löve. She received her PhD in...

Triticum turanicum Jakubz.
Persian wheat BAu Triticum turgidum L. subsp. carthlicum (Nevski) Á.
Áskell Löve
Áskell Löve was an Icelandic systematic botanist, particularly active in the Arctic.Löve was born in Reykjavik. He studied botany at Lund University, Sweden, from 1937. He received his PhD in 1942 in botany and a D.Sc. degree in genetics the year after...

 & D. Löve
Doris Löve
Doris Benta Maria Löve, née Wahlén was a Swedish systematic botanist, particularly active in the Arctic.Doris Löve was born in Kristianstad, Sweden. She studied botany at Lund University from 1937. She married her fellow student and colleague, the Icelander Áskell Löve. She received her PhD in...

Triticum carthlicum Nevski in Kom.
Tetraploid (4x) - timopheevi group
Wild, Hulled
GAu Triticum timopheevii (Zhuk.) Zhuk. subsp. armeniacum (Jakubz.) Slageren Triticum araraticum Jakubz.
Domesticated, Hulled
GAu Triticum timopheevii (Zhuk.) Zhuk. subsp. timopheevii Triticum timopheevii
Triticum timopheevii
Timopheev's Wheat is a tetraploid wheat that has both cultivated and wild forms. The domesticated form is restricted to western Georgia, while the wild form Timopheev's Wheat (Triticum timopheevii, also called Zanduri Wheat) is a tetraploid wheat that has both cultivated and wild forms. The...

(Zhuk.) Zhuk.
Hexaploid (6x), Domesticated, Hulled
Spelt wheat BAuD Triticum aestivum L. subsp. spelta (L.) Thell.
Albert Thellung
-Works:*Illustrierte Flora von Mittel-Europa 1906*Die Gattung Lepidium R. Br. : eine monographische Studie 1906 *Die Entstehung der Kulturpflanzen 1930 *Flora der Schweiz - Exkursionsflora, 1923...

Triticum spelta L.
BAuD Triticum aestivum L. subsp. macha (Dekapr. & A. M. Menabde) Mackey Triticum macha Dekapr. & Menabde
BAuD Triticum vavilovii Jakubz. Triticum vavilovii (Tumanian) Jakubz.
Hexaploid (6x), Domesticated, Free-threshing
Common or bread wheat BAuD Triticum aestivum L. subsp. aestivum Triticum aestivum
Common wheat
Common wheat, Triticum aestivum, is a cultivated wheat species.-Nomenclature and taxonomy of the variety and its cultivars:-Evolution:...

L.
Club wheat BAuD Triticum aestivum L. subsp. compactum (Host) Mackey Triticum compactum
Common wheat
Common wheat, Triticum aestivum, is a cultivated wheat species.-Nomenclature and taxonomy of the variety and its cultivars:-Evolution:...

Host
Indian dwarf or shot wheat BAuD Triticum aestivum L. subsp. sphaerococcum (Percival) Mackey Triticum sphaerococcum
Common wheat
Common wheat, Triticum aestivum, is a cultivated wheat species.-Nomenclature and taxonomy of the variety and its cultivars:-Evolution:...

Percival


Note: Blank common name indicates that no common name is in use in the English language.

Explanatory notes on selected names

  • Triticum boeoticum Boiss. is sometimes divided into two subspecies:
    • T. boeoticum Boiss. subsp. thaoudar (Reut. ex Hausskn.) E. Schiem. - with two grains in each spikelet, distributed to east of fertile crescent.
    • T. boeoticum Boiss. subsp. boeoticum - one grain in each spikelet, in Balkans.

  • Triticum dicoccum Schrank
    Franz Paula von Schrank
    Franz von Paula Schrank was a German botanist and entomologist.Schrank was the first director of the botanical gardens in Munich from 1809 to 1832.Shrank was the first author to use the genus name Triops...

     ex Schübler
    Gustav Schübler
    Gustav Schübler was a German naturalist, and the founder of applied meteorology in Germany.In 1817 Schübler became professor of botany, natural history and agricultural chemistry at the University of Tübingen, Germany...

    is also known as Triticum dicoccon Schrank
    Franz Paula von Schrank
    Franz von Paula Schrank was a German botanist and entomologist.Schrank was the first director of the botanical gardens in Munich from 1809 to 1832.Shrank was the first author to use the genus name Triops...

    .

  • Triticum aethiopicum Jakubz. is a variant form of T. durum found in Ethiopia. It is not usually regarded as a separate species.

  • Triticum karamyschevii Nevsky was previously known as Triticum paleocolchicum A. M. Menabde.

Artificial species and mutants

Russian botanists have given botanical names to hybrids developed during genetical experiments. As these only occur in the laboratory environment, it is questionable whether botanical names (rather than lab. numbers) are justified. Botanical names have also been given to rare mutant forms. Examples include:
  • Triticum × borisovii Zhebrak - (T. aestivum × T. timopheevi)
  • Triticum × fungicidum Zhuk. - Hexaploid, artificial cross (T. carthlicum × T. timopheevi)
  • Triticum jakubzineri Udaczin & Schachm.
  • Triticum militinae Zhuk. & Migush. - mutant form of T. timopheevi.
  • Triticum petropavlovskyi Udaczin & Migush.
  • Triticum sinskajae A.A.Filatenko & U.K.Kurkiev - mutant, free-threshing form of T. monococcum.
  • Triticum × timococcum Kostov
  • Triticum timonovum Heslot - Hexaploid, artificial cross.
  • Triticum zhukovskyi Menabde & Ericzjan (T. timopheevi × T. monococcum)

Germplasm


Taxonomy


Genetics


Morphology


See also

  • Cultivated plant taxonomy
    Cultivated plant taxonomy
    Cultivated plant taxonomy is the study of the theory and practice of the science that identifies, describes, classifies, and names cultigens—those plants whose origin or selection is primarily due to intentional human activity...

  • List of Canadian Heritage Wheat Varieties
The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
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