The Chenopodioideae are a subfamily of the

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 ...

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, making it ...

in the APG III system, which is largely based on

molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

, but were included - together with other subfamilies - in family

Chenopodiaceae 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, making i ...

in the

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) ...

. Food species comprise

Spinach Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to central and western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common edible vegetable consumed either f ...

(''Spinacia oleracea''),

Good King Henry ''Blitum bonus-henricus'' ( syn. ''Chenopodium bonus-henricus''), also called Good-King-Henry, poor-man's asparagus, perennial goosefoot, Lincolnshire spinach, Markery, English mercury, or mercury goosefoot, is a species of goosefoot which is na ...

(''Blitum bonus-henricus''), several ''

Chenopodium ''Chenopodium'' is a genus of numerous species of perennial or annual herbaceous flowering plants known as the goosefoots, which occur almost anywhere in the world. It is placed in the family Amaranthaceae in the APG II system; older classifica ...

'' species (

Quinoa Quinoa (''Chenopodium quinoa''; , from Quechua ' or ') is a flowering plant in the amaranth family. It is a herbaceous annual plant grown as a crop primarily for its edible seeds; the seeds are rich in protein, dietary fiber, B vitamins, a ...

Kañiwa ''Chenopodium pallidicaule'', known as ''cañihua'', ''canihua'' or ''cañahua'' (from Quechua ''qañiwa, qañawa or qañawi'') and also kaniwa, is a species of goosefoot, similar in character and uses to the closely related ''quinoa'' ''(Chenopo ...

, Fat Hen),

Orache ''Atriplex'' () is a plant genus of about 250 species, known by the common names of saltbush and orache (; also spelled orach). It belongs to the subfamily Chenopodioideae of the family Amaranthaceae ''s.l.''. The genus is quite variable and ...

(''Atriplex spp.''), and

Epazote ''Dysphania ambrosioides'', formerly ''Chenopodium ambrosioides'', known as Jesuit's tea, Mexican-tea, ''payqu'' ''(paico)'', ''epazote'', ''mastruz'', or ''herba sanctæ Mariæ'', is an annual or short-lived perennial herb native to Central A ...

(''Dysphania ambrosioides''). The name is Greek for

goosefoot ''Chenopodium'' is a genus of numerous species of perennial or annual herbaceous flowering plants known as the goosefoots, which occur almost anywhere in the world. It is placed in the family Amaranthaceae in the APG II system; older classific ...

, the common name of a genus of plants having small greenish flowers.

Description

The Chenopodioideae are annual or perennial herbs, subshrubs, shrub or small trees. The leaves are usually alternate and flat. The flowers are often unisexual. Many species are monoecious or have mixed inflorescences of bisexual and unisexual flowers. Some species are

dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...

, like ''

Spinacia ''Spinacia'' is a flowering plant genus in the subfamily Chenopodioideae of the family Amaranthaceae. The most common member is spinach. Description The species in genus ''Spinacia'' are annual or biennial herbs. Plants are always glabrous. ...

'', '' Grayia'', ''

Exomis The exomis ( grc, ἐξωμίς from ''exo'' "outside", and ''omos'' "shoulder") was a Greek tunic used by the workers and the light infantry. The tunic largely replaced the older chitoniskos (or short chiton) as the main tunic of the hoplites ...

'', and '' Atriplex''. In several species of tribe Atripliceae, the female flowers are without perianth, but enclosed by two bracts. The species with a perianth have up to five

tepal 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 very ...

s. The seed is horizontal or vertical, with annular or horseshoe-shaped embryo.

Distribution

The subfamily Chenopodioideae is distributed worldwide, but originates from

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...

Systematics

The genera of this subfamily were formerly classified in family

in the

. According to Fuentes-Bazan et al. (2012) and based on molecular genetic research, the subfamily comprises 4 tribes and includes about 26 genera: * Tribus Anserineae Dumort. (Syn. Spinacieae), with 2 genera: ** ''

'' L.: with 3 species in Western Asia and North Africa: ***

(''Spinacia oleracea'') ** ''

Blitum ''Blitum'' is a genus of flowering plants in the amaranth family Amaranthaceae, subfamily Chenopodioideae. It is closely related to genus ''Spinacia''. Its 12 species were traditionally placed in the genera '' Chenopodium'', ''Monolepis'', or ''S ...

'', with 12 species nearly worldwide, for example: *** ''

Blitum capitatum Strawberry blite (''Blitum capitatum'', syn. ''Chenopodium capitatum'') is an edible annual plant, also known as blite goosefoot, strawberry goosefoot, strawberry spinach, Indian paint, and Indian ink. It is native to most of North America thr ...

'' - Strawberry Blite (Syn. ''Chenopodium capitatum'') *** '' Blitum bonus-henricus'' - Good king henry (Syn. ''Chenopodium bonus-henricus'') *** '' Blitum virgatum'' - leafy goosefoot (Syn. ''Chenopodium foliosum'') * Tribus

Atripliceae Atripliceae are a tribe of the subfamily Chenopodioideae belonging to the plant family Amaranthaceae. '' Atriplex'' is the largest genus of the tribe. Species of Atripiceae are ecologically important in steppe and semi-desert climates. Distrib ...

C. A. Mey. (Syn. Chenopodieae Dumort.): Fuentes-Bazan et al. (2012) include here also ''Chenopodium'' and related genera, as ''Chenopodiastrum'', ''Lipandra'' and ''Oxybasis''. **'' Archiatriplex'' G.L.Chu, with only one species: ***''Archiatriplex nanpinensis'' G.L.Chu: endemitic in the Chinese province Sichuan. ** '' Atriplex'' L. - saltbush, orache (Syn.: ''Blackiella, Cremnophyton, Haloxanthium, Neopreissia, Obione, Pachypharynx, Senniella, Theleophyton''), with about 300 species worldwide ** '' Baolia'' H.W.Kung & G.L.Chu, with only one species: *** ''Baolia bracteata'' H.W.Kung & G.L.Chu, endemitic in the Chinese province Gansu. ** '' Chenopodiastrum'' S. Fuentes, Uotila & Borsch: with 5 species, for example: *** '' Chenopodiastrum hybridum'' (L.) S. Fuentes, Uotila & Borsch (Syn. ''Chenopodium hybridum'' L.) *** ''

Chenopodiastrum murale ''Chenopodiastrum murale'', (Syn. ''Chenopodium murale'') is a species of plant in the family Amaranthaceae known by the common names nettle-leaved goosefoot, Australian-spinach, salt-green, and sowbane. This plant is native to Europe and parts o ...

'' (L.) S. Fuentes, Uotila & Borsch - Sowbane, nettle-leaf goosefoot (Syn. ''Chenopodium murale'' L.) *** ''

Chenopodiastrum simplex ''Chenopodiastrum'' is a genus of herbaceous flowering plants in the family Amaranthaceae. The genus was formally described in 2012. The 5 species occur in Eurasia, North Africa, and North America. Description The species in genus ''Chenopod ...

'' (Torrey) S.Fuentes, Uotila & Borsch - Maple-leaf goosefoot (Syn.: ''Chenopodium simplex'' (Torrey) Raf.) ** ''

'' L. - goosefoot ('' sensu stricto'', incl. ''Einadia'' Raf. and ''Rhagodia'' R.Br.): with about 90 species worldwide. ** ''

'' Fenzl ex Moq., with only one species: *** ''Exomis microphylla'' (Thunb.) Aellen: a subshrub in southern and western Africa growing in gardens and hedges. ** ''

Extriplex ''Extriplex'' is a plant genus in the subfamily Chenopodioideae of the family Amaranthaceae. It has been described in 2010 and comprises two species, that were formerly included in genus '' Atriplex''. They are restricted to the California Flori ...

'' E.H.Zacharias, with 2 species in western North America: *** ''

Extriplex californica ''Extriplex californica'' is a plant species known by the common name California saltbush or California orache. Formerly, it was included in genus '' Atriplex''. It is native to coastal California and Baja California, where it grows in areas with ...

'' (Moq.) E.H.Zacharias - California saltbush, California orache (Syn.: ''Atriplex californica'' Moq.) *** ''

Extriplex joaquinana ''Extriplex joaquinana'' is a species known by the common name San Joaquin saltbush. It was formerly included in genus '' Atriplex''. Distribution It is endemic to California, where it grows in alkaline soils in the Sacramento-San Joaquin River ...

'' (A.Nelson) E.H.Zacharias - San Joaquin saltbush, San Joaquin orach (Syn.: ''Atriplex joaquinana'' A.Nelson) ** '' Grayia'' Hook. & Arn. - siltbush, hopsage (Syn. ''Zuckia'' Standl.), with 4 shrubby species in western North America, for example: *** ''

Grayia spinosa ''Grayia'' may refer to: * ''Grayia'' (snake), a genus of African water snakes * ''Grayia'' (plant), a genus of desert shrubs in the family Amaranthaceae {{Genus disambiguation ...

'' (Hook.) Moq. - spiny hopsage ** ''

Halimione ''Halimione'' is a plant genus from the subfamily Chenopodioideae of the family Amaranthaceae. It is a sister genus of ''Atriplex'' and is included in that genus by Plants of the World Online. Description The species in genus ''Halimione'' are ...

'' Aellen - purslane, with 3 species in Europe and Western Asia, for example: *** ''

Halimione portulacoides ''Halimione portulacoides'' or sea purslane (2n=36) is a shrub found in Eurasia. Description The plant grows to in height. It is evergreen, and in northern temperate climates it flowers from July to September. The flowers are monoecious and a ...

'' (L.) Aellen (Syn.: ''Atriplex portulacoides'' L.) - sea purslane ** '' Holmbergia'' Hicken, with only one species: *** ''Holmbergia tweedii'' (Moq.) Speg., a shrub in Bolivia, Paraguay and Argentina. ** ''

Lipandra ''Lipandra polysperma'' (Syn. ''Chenopodium polyspermum''), common name manyseed goosefoot, is the only species of the monotypic plant genus ''Lipandra'' from the subfamily Chenopodioideae of the family Amaranthaceae. Description ''Lipandra p ...

'' Moq.: with only one species: *** ''Lipandra polysperma'' (L.) S. Fuentes, Uotila & Borsch (Syn. ''Chenopodium polyspermum'' L.) - many-seed goosefoot ** '' Manochlamys'' Aellen, with only one species: *** ''Manochlamys albicans'' Aellen: a subshrub in southern Africa, Namibia and Cape province, growing on rocky and sandy slopes, sand dunes and road sides. ** '' Microgynoecium'' Hook.f., with only one species: *** ''Microgynoecium tibeticum'' Hook.f.: in Tibet and Sikkim, growing in alpine meadows and on disturbed sites. ** ''

Micromonolepis ''Micromonolepis pusilla'', (Syn. ''Monolepis pusilla'') is the only species of the genus ''Micromonolepis'' in the flowering plant family Amaranthaceae, known by the common names small povertyweed and red povertyweed. It is native to the Western ...

'' Ulbr., with only one species: *** ''Micromonolepis pusilla'' (Torr. ex S. Watson) Ulbr. - small povertyweed, in western North America ** ''

Oxybasis ''Oxybasis'' is a flowering plant genus from the subfamily Chenopodioideae of the family Amaranthaceae. It was first described in 1841, and newly used since 2012 for five species that were traditionally grouped into genus ''Chenopodium''. They o ...

'' Kar. & Kir.: with 5 species, for example: *** '' Oxybasis chenopodioides'' (L.) S. Fuentes, Uotila & Borsch - low goosefoot (Syn. ''Chenopodium chenopodioides'' (L.) Aellen) *** ''

Oxybasis glauca ''Oxybasis glauca'' (syn. ''Chenopodium glaucum''), common name oak-leaved goosefoot, is a species of goosefoot plant native to Europe. It has been introduced and become an invasive weed in North America. This invader of European origin also appe ...

'' (L.) S. Fuentes, Uotila & Borsch - Oak-leaved goosefoot (Syn. ''Chenopodium glaucum'' L.) *** '' Oxybasis rubra'' (L.) S. Fuentes, Uotila & Borsch - Red Goosefoot (Syn. ''Chenopodium rubrum'' L.) ** ''

Proatriplex ''Proatriplex'' is a monotypic plant genus in the subfamily Chenopodioideae of the family Amaranthaceae, with the only species ''Proatriplex pleiantha'' (syn. ''Atriplex pleiantha''). It is known by the common names four-corners orach and Mancos ...

'' (W.A.Weber) Stutz & G.L.Chu, with only one species: *** ''Proatriplex pleiantha'' (W.A.Weber) Stutz & G.L.Chu, an annual herb from western North America. ** ''

Stutzia ''Stutzia'' is a plant genus in the subfamily Chenopodioideae of the family Amaranthaceae. It was described in 2010, replacing the illegitimate name ''Endolepis''. It comprises two species, that have also been included in the genus '' Atriplex'' ...

'' E.H.Zacharias (Syn. ''Endolepis'' Torr.), with 2 annual species in western North America: *** '' Stutzia covillei'' (Standl.) E.H.Zacharias (Syn. ''Atriplex covillei'' (Standl.) J. F. Macbr., ''Endolepis covillei'' Standl) *** '' Stutzia dioica'' (Nutt.) E.H.Zacharias (Syn. ''Atriplex suckleyi'' (Torrey) Rydberg, ''Endolepis suckleyi'' Torr.) * Tribus Axyrideae (Heklau) G. Kadereit & A. Sukhor., with dendritic trichomes. 3 genera: ** ''

Axyris ''Axyris'', the Russian pigweeds, are a genus of flowering plants in the amaranth family Amaranthaceae, native to temperate parts of Eastern Europe and Asia. The center of genetic diversity is the Altai to northern Tien-Shan mountains. The best ...

'' L., with about 6 species Central Asia, Himalaya and western China, for example: *** ''

Axyris amaranthoides ''Axyris amaranthoides'' is a species of flowering plant, in the family Amaranthaceae, that has been introduced species, introduced to North America. It was introduced into Manitoba in 1886 and has since spread to other provinces in Canada and th ...

'' L.- Russian pigweed, upright axyris ** ''

Ceratocarpus ''Ceratocarpus'' is a genus of flowering plants in the 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 ...

'' L., with 2 species in Europe and West Asia ** ''

Krascheninnikovia ''Krascheninnikovia'' is a genus of flowering plants in the subfamily Chenopodioideae of the family Amaranthaceae known as winterfat, so-called because it is a nutritious livestock forage. They are known from Eurasia and western North America. Th ...

'' Gueldenst., with 8 species in Eurasia and North America, for example: *** ''

Krascheninnikovia lanata ''Krascheninnikovia'' is a genus of flowering plants in the subfamily Chenopodioideae of the family Amaranthaceae known as winterfat, so-called because it is a nutritious livestock forage. They are known from Eurasia and western North America. Th ...

'' (Pursh) A.Meeuse & A.Smit - winterfat * Tribus Dysphanieae: ** '' Cycloloma'' Moq. (Syn.: Cyclolepis Moquin-Tandon) with only one species: *** ''Cycloloma atriplicifolium'' (Sprengel) J.M.Coulter: widespread in Canada, USA and northern Mexico ** '' Dysphania'' R.Br., with about 42 species worldwide, for example: *** ''

Dysphania ambrosioides ''Dysphania ambrosioides'', formerly ''Chenopodium ambrosioides'', known as Jesuit's tea, Mexican-tea, ''payqu'' ''(paico)'', ''epazote'', ''mastruz'', or ''herba sanctæ Mariæ'', is an annual or short-lived perennial herb native to Central ...

'' - epazote *** ''

Dysphania anthelmintica ''Dysphania'' may refer to: * ''Dysphania'' (moth), a Lepidoptera animal genus * ''Dysphania'' (plant), an Amaranthaceae plant genus {{Genus disambiguation ...

'' - wormseed ** ''

Suckleya ''Suckleya'' is a monotypic genus of flowering plant belonging to the family Amaranthaceae. It only contains one known species, Suckleya suckleyana (Torr.) Rydb. Its native range is western central Canada (within the province of Alberta) to weste ...

'' A.Gray, with only one species: *** ''Suckleya suckleyana'' (Torr.) Rydb., a succulent annual from western North America. ** ''

Teloxys ''Teloxys'' is a genus of flowering plants belonging to the 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 goosef ...

'' Moq.: with only one species: *** ''Teloxys aristata'' (L.) Moq. (Syn.: ''Chenopodium aristatum'' L., ''Dysphania aristata''): from Eastern Europe to temperate Asia, naturalized elsewhere.

Fossil record

The oldest

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

records for Chenopodioideae are

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...

grains recovered from

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...

sediments of the Edmonton Formation in

Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...

.Assorted angiosperm pollen from the Edmonton Formation (Maestrichtian), Alberta, Canada by Satish K. Srivastava – Canadian Journal of Botany, 1969, 47(6): 975-989, https://doi.org/10.1139/b69-138

References

* Susy Fuentes-Bazan, Pertti Uotila, Thomas Borsch (2012): ''A novel phylogeny-based generic classification for Chenopodium sensu lato, and a tribal rearrangement of Chenopodioideae (Chenopodiaceae).'' In: ''Willdenowia.'' Vol. 42, No. 1, p. 5-24. * Gudrun Kadereit, Evgeny V. Mavrodiev, Elizabeth H. Zacharias & Alexander P. Sukhorukov: ''Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae) (2010): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 Photosynthesis''. - In: ''American Journal of Botany'' 97(10): p. 1664-1687. (chapters description, distribution and systematics) * A.P. Sukhorukov, M. Zhang (2013): Fruit and seed Anatomy of Chenopodium and related genera (Chenopodioideae, Chenopodiaceae/Amaranthaceae): Implications for evolution and taxonomy. - PLOS ONE. Vol. 8, № 4. e61906. doi:10.1371/journal.pone.006190
online
Susy Fuentes-Bazan, Guilhem Mansion, Thomas Borsch: ''Towards a species level tree of the globally diverse genus Chenopodium (Chenopodiaceae)''. In: ''Molecular Phylogenetics and Evolution'', online 22 October 2011, ISSN 1055-7903

Elizabeth H. Zacharias, Bruce G. Baldwin (2010): ''A Molecular Phylogeny of North American Atripliceae (Chenopodiaceae), with Implications for Floral and Photosynthetic Pathway Evolution''. In: ''Systematic Botany'' 35(4), p.839-857.

External links

* {{Taxonbar, from=Q1070132 Chenopodioideae, Caryophyllales subfamilies