Miracle wheat (Triticum turgidum var. mirabile).
During 10,000 years of cultivation, numerous forms of wheat, many of them hybrids, have developed under a combination of artificial and natural 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. Information on the cultivation and uses of wheat is at the main wheat page.
Aegilops and Triticum
Spike and spikelets of Aegilops tauschii
The genus Triticum includes the wild and domesticated species usually thought of as wheat.
In the 1950s growing awareness of the genetic similarity of the wild goatgrasses (Aegilops) led botanists such as Bowden 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 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 closely related 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, 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.
Important characters in wheat
Ploidy level
As with many grasses, polyploidy is common in wheat. There are two wild diploid (non-polyploid) 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
are sometimes found. Hexaploid wheats developed 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, 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).
- A – 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. tauschii, 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
- GAm T. timopheevi
- BAuD, T. aestivum
Domestication
There are four wild species, all growing in rocky habitats in the fertile crescent of the Near East. All the other species are domesticated.
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 Pre-Pottery Neolithic B 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.
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 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).
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 classification 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
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. The term landrace is applied to informal, farmer-maintained populations of crop plants.
Naming
Botanical names for wheat are generally expected to follow an existing classification, such as those listed as current at the Wheat Classification Tables Site.
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 on the Wheat page.
A general rule 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 ) | Traditional (Dorofeev et al. 1979 ) |
| Diploid (2x), Wild, Hulled | |||
| Wild einkorn | Am | Triticum monococcum L. subsp. aegilopoides (Link) Thell. | 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. | Triticum dicoccoides (Körn. ex Asch. & Graebner) Schweinf. |
| Tetraploid (4x), Domesticated, Hulled | |||
| Emmer | BAu | Triticum turgidum L. subsp. dicoccum (Schrank ex Schübl.) Thell. | Triticum dicoccum Schrank ex Schübler |
| BAu | Triticum ispahanicum Heslot | Triticum ispahanicum Heslot | |
| BAu | Triticum turgidum L. subsp. paleocolchicum Á. & D. Löve | Triticum karamyschevii Nevski | |
| Tetraploid (4x), Domesticated, Free-threshing | |||
| Durum or macaroni wheat | BAu | Triticum turgidum L. subsp. durum (Desf.) Husn. | Triticum durum Desf. |
| Rivet, cone or English wheat | BAu | Triticum turgidum L. subsp. turgidum | Triticum turgidum L. |
| Polish wheat | BAu | Triticum turgidum L. subsp. polonicum (L.) Thell. | Triticum polonicum L. |
| Khorasan wheat | BAu | Triticum turgidum L. subsp. turanicum (Jakubz.) Á. & D. Löve | Triticum turanicum Jakubz. |
| Persian wheat | BAu | Triticum turgidum L. subsp. carthlicum (Nevski) Á. & D. Löve | Triticum carthlicum Nevski in Kom. |
| Tetraploid (4x) – timopheevi group | |||
| Wild, Hulled | |||
| GAm | Triticum timopheevii (Zhuk.) Zhuk. subsp. armeniacum (Jakubz.) Slageren | Triticum araraticum Jakubz. | |
| Domesticated, Hulled | |||
| GAm | Triticum timopheevii (Zhuk.) Zhuk. subsp. timopheevii | Triticum timopheevii (Zhuk.) Zhuk. | |
| Hexaploid (6x), Domesticated, Hulled | |||
| Spelt wheat | BAuD | Triticum aestivum L. subsp. spelta (L.) Thell. | 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 L. |
| Club wheat | BAuD | Triticum aestivum L. subsp. compactum (Host) Mackey | Triticum compactum Host |
| Indian dwarf or shot wheat | BAuD | Triticum aestivum L. subsp. sphaerococcum (Percival) Mackey | Triticum sphaerococcum 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 ex Schübler is also known as Triticum dicoccon Schrank.
- 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)
