The Betoideae are a small subfamily of the

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

amaranth family,

Amaranthaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type genus '' Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, maki ...

''sensu lato'' (or in Chenopodiaceae ''sensu stricto''). Commonly known members include beet, sugar beet, chard, and mangelwurzel, which all are cultivars of ''

Beta vulgaris ''Beta vulgaris'' (beet) is a species of flowering plant in the subfamily Betoideae of the family Amaranthaceae. Economically, it is the most important crop of the large order Caryophyllales. It has several cultivar groups: the sugar beet, of gre ...

''.

Description

The species of Betoideae are annuals, biennial or perennial herbs, vines ''( Hablitzia)'' or subshrubs. The flowers have 5

tepals A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of ve ...

('' Aphanisma'' only 3) and 5

stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

s ''( Aphanisma'' only one). The fruits of Betoideae are capsules that open with a circumscissile lid. In tribe Beteae, the perianth is basally indurated in fruit, and the stamens a basally inserted to a thickened bulge surrounding the visible part of the ovary. In tribe Hablitzieae, the tepals are not modified in fruit and membranous, and the stamens are basally united in a membranous ring.

Distribution and habitat

Most genera are distributed in Western and Southern Europe, in the Mediterranean and Southwest Asia, but one disjunct genus, '' Aphanisma'', lives at the coasts of California. The species of Betoideae are adapted to different ecological habitats, several growing in coastal habitats, some on rocks and in mountains, one in deciduous forests (''Hablitzia'').

Systematics

Oskar Eberhard Ulbrich described the subfamily Betoideae in 1934 within the plant family

Chenopodiaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type (biology), type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 ...

. He subdivided the taxon into two tribes, Hablitzieae and Beteae, the latter with only one genus, ''

Beta Beta (, ; uppercase , lowercase , or cursive ; or ) is the second letter of the Greek alphabet. In the system of Greek numerals, it has a value of 2. In Ancient Greek, beta represented the voiced bilabial plosive . In Modern Greek, it represe ...

''.

Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

research by Kadereit et al. (2006) confirmed this classification, whereas Romeira et al. (2016) suggest only one tribe. The subfamily is now classified either in family

''sensu lato'', or in Chenopodiaceae ''sensu stricto'', (excluding the subfamily Polycnemoideae, as the other subfamilies of Chenopodiaceae, Betoideae, Camphorosmoideae,

Chenopodioideae The Chenopodioideae are a subfamily of the flowering plant family Amaranthaceae in the APG III system, which is largely based on molecular phylogeny, but were included – together with other subfamilies – in the family Chenopodiaceae, or goose ...

, Corispermoideae, Salicornioideae, Salsoloideae, and Suaedoideae, form a monophyletic group which is distinct from the Amaranthaceae ''s. str.''). Current taxonomic treatments and morphological, physiological and phylogenetic studies seem to prefer Chenopodiaceae ''s. str.'' for reasons of taxonomic stability. The subfamily comprises five genera with about 13-20 species. * Tribus Beteae Moq.: ** ''

'' L., about 7-12 species in West-Europe, Mediterranean, Southwest-Asia, with the important crops

Sugar beet A sugar beet is a plant whose root contains a high concentration of sucrose and that is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together with ...

Chard Chard (; '' Beta vulgaris'' subsp. ''vulgaris'', Cicla Group and Flavescens Group) is a green leafy vegetable. In the cultivars of the Flavescens Group, or Swiss chard, the leaf stalks are large and often prepared separately from the leaf b ...

Beetroot The beetroot (British English) or beet (North American English) is the taproot portion of a ''Beta vulgaris'' subsp. ''vulgaris'' plant in the Conditiva Group. The plant is a root vegetable also known as the table beet, garden beet, dinner bee ...

Mangelwurzel Mangelwurzel or mangold wurzel (from German ''Mangel/Mangold'', "chard" and ''Wurzel'', "root"), also called mangold,Wright, Clifford A. (2001) ''Mediterranean Vegetables: a cook's ABC of vegetables and their preparation in Spain, France, Italy, ...

* Tribus Hablitzieae Ulbr.: ** '' Aphanisma'' Nutt. ex Moq., with one species: *** ''Aphanisma blitoides'' Nutt. ex Moq., an annual plant on Californian beaches ** '' Hablitzia'' M.Bieb., with one species *** ''Hablitzia tamnoides'' M.Bieb., a vine in the forests of the Caucasian floristic region ** '' Oreobliton'' Durieu, with one species *** ''Oreobliton thesioides'' Durieu & Moq., a subshrub in North-Africa, growing on chalk rocks in the Atlas mountains ** '' Patellifolia'' A.J.Scott, Ford-Lloyd & J.T. Williams (Syn. ''Beta ''sect.'' Procumbentes'' Moq.), was confirmed as a different genus. These are perennial procumbent herbs, with 3 species growing in coastal vegetation in Southern-Western Europe, with its center of diversity in the Macaronesian archipelagos: *** '' Patellifolia patellaris'' (Moq.) A.J. Scott & al. (Syn. ''Beta patellaris'' Moq.), on Canary Islands and in the western Mediterranean (Spain, Balearic islands, Sicily, Marocco) *** '' Patellifolia procumbens'' (Chr. Sm.) A.J. Scott & al. (Syn. ''Beta procumbentes'' Chr. Sm.), on Canary Islands *** '' Patellifolia webbiana'' (Moq.) A.J. Scott & al. (Syn. ''Beta webbiana'' Moq.), on Canary Islands *Classification not sure: phylogenetically isolated, formerly included in Betoideae but to be excluded, maybe to be classified as a subfamily of its own. Recent molecular research provides some evidence for the inclusion in subfamily Corispermoideae: ** '' Acroglochin'' Schrad. ex Schult., with one species ***''Acroglochin persicarioides'' (Poiret) Moq.

Phylogenetics

The subfamily Betoideae is regarded as a

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

taxon, if '' Acroglochin'' is excluded. The age of the subfamily seems to be relatively old, originating during the Early

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

Glacial Maximum, estimated 48.6-35.4 million years ago They early diversified into genera about 32.5 million years ago. The extant genera show narrow distributions in distant geographic regions, which may have resulted from speciation by isolation and following extinction events. The areals of '' Aphanisma'' in California and '' Oreobliton'' in North-Africa are interpreted as remnants from a

Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...

n ancestor, the disjunction circa 15.4-9.2 million years ago. The differentiation between ''Beta'' and ''Patellifolia'' probably occurred early in the Late Oligocene. Both lineages tolerate aridity and highly saline soils, so they were able to survive dramatic aridity events in the past that led to the extinction of other more vulnerable lineages in the subfamily.

Uses

Beet (''Beta vulgaris'') has an immense economic importance as sugar crop (

sugar beet A sugar beet is a plant whose root contains a high concentration of sucrose and that is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together with ...

), and a great importance as a

vegetable Vegetables are edible parts of plants that are consumed by humans or other animals as food. This original meaning is still commonly used, and is applied to plants collectively to refer to all edible plant matter, including edible flower, flo ...

(

chard Chard (; '' Beta vulgaris'' subsp. ''vulgaris'', Cicla Group and Flavescens Group) is a green leafy vegetable. In the cultivars of the Flavescens Group, or Swiss chard, the leaf stalks are large and often prepared separately from the leaf b ...

beetroot The beetroot (British English) or beet (North American English) is the taproot portion of a ''Beta vulgaris'' subsp. ''vulgaris'' plant in the Conditiva Group. The plant is a root vegetable also known as the table beet, garden beet, dinner bee ...

), and as fodder plant (

mangelwurzel Mangelwurzel or mangold wurzel (from German ''Mangel/Mangold'', "chard" and ''Wurzel'', "root"), also called mangold,Wright, Clifford A. (2001) ''Mediterranean Vegetables: a cook's ABC of vegetables and their preparation in Spain, France, Italy, ...

). This species is also used as

medicinal plant Medicinal plants, also called medicinal herbs, have been discovered and used in traditional medicine practices since prehistoric times. Plants synthesize hundreds of chemical compounds for various functions, including Plant defense against h ...

ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...

dye Juan de Guillebon, better known by his stage name DyE, is a French musician. He is known for the music video of the single "Fantasy Fantasy is a genre of speculative fiction that involves supernatural or Magic (supernatural), magical ele ...

and as

renewable resource A renewable resource (also known as a flow resource) is a natural resource which will replenish to replace the portion depleted by usage and consumption, either through natural reproduction or other recurring processes in a finite amount of t ...

. It is the crop species with the highest economic value in the order

Caryophyllales Caryophyllales ( ) is a diverse and heterogeneous order of flowering plants with well-known members including cacti, carnations, beets, quinoa, spinach, amaranths, pigfaces and ice plants, oraches and saltbushes, goosefoots, sundews, Venu ...

. Therefore, the other members of Betoideae, especially ''

'' and '' Patellifolia'', are interesting as

crop wild relative A crop wild relative (CWR) is a wild plant closely related to a domesticated plant. It may be a wild ancestor of the domesticated (cultivated) plant or another closely related taxon. Overview The wild relatives of crop plants constitute an i ...

References

The family Amaranthaceae
a

/ref> Hernández-Ledesma P., Berendsohn W. G., Borsch Th., Mering S. von, Akhani H., Arias S., Castañeda-Noa I., Eggli U., Eriksson R., Flores-Olvera H., Fuentes-Bazán S., Kadereit G., Klak C., Korotkova N., Nyffeler R., Ocampo G., Ochoterena H., Oxelman B., Rabeler R. K., Sanchez A., Schlumpberger B. O. & Uotila P. 2015. "A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales". — Willdenowia 45: 281–383. Hohmann, S., Kadereit, J.w., & Kadereit, G. (2006). "Understanding Mediterranean-Californian disjunctions: molecular evidence". ''Taxon'' 55 (1): 67–78. Kadereit, G., Hohmann, S. & Kadereit, J.W. (2006). "A synopsis of Chenopodiaceae subfam. Betoideae and notes on the taxonomy of ''Beta''". - Willdenowia 36, p. 9-19. Kai Müller, Thomas Borsch (2005). "Phylogenetics of Amaranthaceae using matK/trnK sequence data – evidence from parsimony, likelihood and Bayesian approaches". - ''Annals of the Missouri Botanical Garden'', 92: p.66-102. Romeiras, M.M., Vieira, A., Silva, D.N., Moura, M., Santos-Guerra, A., Batista, D., Duarte, M.C., & Paulo, O.S. (2016). "Evolutionary and Biogeographic Insights on the Macaronesian ''Beta-Patellifolia'' Species (Amaranthaceae) from a Time-Scaled Molecular Phylogeny." PLoS One. 2016; 11(3): e0152456. Ulbrich, O.E. (1934). "Chenopodiaceae". - In: Engler, A. & Prantl, K.A.E. (eds.): "Die natürlichen Pflanzenfamilien", ed.2. Vol. 16c: p.379-584, Duncker & Humblot, Berlin , p. 455.

External links

* *
Betoideae at GRIN
{{Authority control Amaranthaceae Caryophyllales subfamilies Taxa named by Oskar Eberhard Ulbrich