View source for Chenopodioideae

{{short description|Subfamily of flowering plants}}
{{Automatic taxobox
| image = Chenopodium berlandieri NPS-1.jpg
| image_caption = ''[[Chenopodium berlandieri]]''  
| taxon = Chenopodioideae
| authority = Burnett
| subdivision_ranks = Genera
| subdivision = About 26 genera, see text
}}

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 [[goosefoot]] family, in the [[Cronquist system]].

Food species comprise [[spinach]] (''Spinacia oleracea''), [[Good King Henry]] (''Blitum bonus-henricus''), several ''[[Chenopodium]]'' species ([[quinoa]], [[Chenopodium pallidicaule|kañiwa]], [[Chenopodium album|fat hen]]), [[Atriplex|orache]] (''Atriplex'' spp.), and [[Dysphania ambrosioides|epazote]] (''Dysphania ambrosioides''). The name is Greek for goosefoot, 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]], like ''[[Spinacia]]'', ''[[Grayia (plant)|Grayia]]'', ''[[Exomis microphylla]]'', 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]]s. The seed is horizontal or vertical, with annular or horseshoe-shaped embryo.

== Distribution ==
The subfamily Chenopodioideae is distributed worldwide, but originates from [[Eurasia]].

== Systematics ==
[[File:Blitum bonus-henricus.jpg|thumb|Good King Henry (''[[Blitum bonus-henricus]]''), tribe Anserineae]]
[[File:Rhagodia spinescens 1.jpg|thumb|''[[Chenopodium spinescens]]'', tribe Atripliceae]]
[[File:Grayia spinosa 3.jpg|thumb|''[[Grayia spinosa]]'', tribe Atripliceae]]
[[File:Lipandra polysperma RF.jpg|thumbnail|''[[Lipandra polysperma]]'', tribe Atripliceae]]
[[File:Krascheninnikovia lanata 3.jpg|thumb|''[[Krascheninnikovia lanata]]'', tribe Axyrideae]]
[[File:Starr 080415-3985 Chenopodium carinatum.jpg|thumbnail|''[[Dysphania carinata]]'', tribe Dysphanieae]]
The genera of this subfamily were formerly classified in family [[Chenopodiaceae]] in the [[Cronquist system]].

According to Fuentes-Bazan et al. (2012) and based on molecular genetic research, the subfamily comprises four tribes and includes about twenty-six genera:
* Tribus [[Anserineae]] Dumort. (Syn. Spinacieae), with two genera: 
** ''[[Spinacia]]'' L.: with three species in Western Asia and North Africa:
*** [[Spinach]] (''Spinacia oleracea'')
** ''[[Blitum]]'', with 12 species nearly worldwide, for example: 
*** ''[[Blitum capitatum]]'' – Strawberry Blite (Syn. ''Chenopodium capitatum'') 
*** ''[[Blitum bonus-henricus]]'' – Good King Henry (Syn. ''Chenopodium bonus-henricus'') 
*** ''[[Blitum virgatum]]'' – leafy goosefoot (Syn. ''Chenopodium foliosum'')
* Tribus [[Atripliceae]] 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: endemic in the Chinese province Sichuan.<ref>Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=102465 ''Archiatriplex'' - Online], In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (Hrsg.): ''Flora of China'', Volume 5: ''Ulmaceae through Basellaceae'', Science Press und Missouri Botanical Garden Press, Beijing und St. Louis, {{ISBN|1-930723-27-X}}. p. 360</ref>
** ''[[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, endemic in the Chinese province Gansu.<ref>Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=103468 ''Baolia'' - Online], In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (Hrsg.): ''Flora of China'', Volume 5: ''Ulmaceae through Basellaceae'', Science Press und Missouri Botanical Garden Press, Beijing und St. Louis, {{ISBN|1-930723-27-X}}. p. 375</ref>
** ''[[Chenopodiastrum]]'' S. Fuentes, Uotila & Borsch: with five species, for example:
*** ''[[Chenopodiastrum hybridum]]'' (L.) S. Fuentes, Uotila & Borsch (Syn. ''Chenopodium hybridum'' L.)
*** ''[[Chenopodiastrum murale]]'' (L.) S. Fuentes, Uotila & Borsch – Sowbane, nettle-leaf goosefoot (Syn. ''Chenopodium murale'' L.)
*** ''[[Chenopodiastrum simplex]]'' (Torrey) S.Fuentes, Uotila & Borsch – Maple-leaf goosefoot (Syn.: ''Chenopodium simplex'' (Torrey) Raf.) 
** ''[[Chenopodium]]'' L. – goosefoot (''[[sensu stricto]]'', incl. ''Einadia'' Raf. and ''Rhagodia'' R.Br.<ref name="Fuentes" />): with about 90 species worldwide. 
** ''[[Exomis (plant)|Exomis]]'' Fenzl ex Moq., with only one species:
*** ''Exomis microphylla'' (Thunb.) Aellen: a subshrub in southern and western Africa growing in gardens and hedges.
** ''[[Extriplex]]'' E.H.Zacharias, with two species in western North America:
*** ''[[Extriplex californica]]'' (Moq.) E.H.Zacharias – California saltbush, California orache (Syn.: ''Atriplex californica'' Moq.)<ref name="Zacharias" />
*** ''[[Extriplex joaquinana]]'' (A.Nelson) E.H.Zacharias – San Joaquin saltbush, San Joaquin orach (Syn.: ''Atriplex joaquinana'' A.Nelson)<ref name="Zacharias" />
** ''[[Grayia (plant)|Grayia]]'' Hook. & Arn. – siltbush, hopsage (Syn. ''Zuckia'' Standl.), with four shrubby species in western North America, for example:
*** ''[[Grayia spinosa]]'' (Hook.) Moq. – spiny hopsage<ref name="Zacharias" />
** ''[[Halimione]]'' Aellen – purslane, with three species in Europe and Western Asia, for example:
*** ''[[Halimione portulacoides]]'' (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]]'' 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]]'' Ulbr., with only one species:
*** ''Micromonolepis pusilla'' (Torr. ex S. Watson) Ulbr. – small povertyweed, in western North America
** ''[[Oxybasis]]'' Kar. & Kir.: with five species, for example:
*** ''[[Oxybasis chenopodioides]]'' (L.) S. Fuentes, Uotila & Borsch – low goosefoot (Syn. ''Chenopodium chenopodioides'' (L.) Aellen) 
*** ''[[Oxybasis glauca]]'' (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]]'' (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]]'' E.H.Zacharias (Syn. ''Endolepis'' Torr.), with two annual species in western North America:
*** ''[[Stutzia covillei]]'' (Standl.) E.H.Zacharias (Syn. ''Atriplex covillei'' (Standl.) J. F. Macbr., ''Endolepis covillei'' Standl)<ref name="Zacharias" />
*** ''[[Stutzia dioica]]'' (Nutt.) E.H.Zacharias (Syn. ''Atriplex suckleyi'' (Torrey) Rydberg, ''Endolepis suckleyi'' Torr.)<ref name="Zacharias" />
* Tribus [[Axyrideae]] (Heklau) G. Kadereit & A. Sukhor., with dendritic trichomes. three genera:
** ''[[Axyris]]'' L., with about six species Central Asia, Himalaya and western China, for example:
*** ''[[Axyris amaranthoides]]'' L. – Russian pigweed, upright axyris
** ''[[Ceratocarpus]]'' L., with two species in Europe and West Asia
** ''[[Krascheninnikovia]]'' Gueldenst., with eight species in Eurasia and western North America, for example:
*** ''[[Krascheninnikovia lanata]]'' (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<ref>Sergei L. Mosyakin (2003): [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=108849 ''Cycloloma'' - Online], In: Flora of North America Editorial Committee (Hrsg.): ''Flora of North America North of Mexico'', Volume 4: ''Magnoliophyta: Caryophyllidae, part 1.'', Oxford University Press, New York, {{ISBN|0-19-517389-9}}, p.264-265</ref>
** ''[[Dysphania (plant)|Dysphania]]'' R.Br., with about 42 species worldwide, for example:
*** ''[[Dysphania ambrosioides]]'' – epazote
*** ''[[Dysphania anthelmintica]]'' –  wormseed
** ''[[Neomonolepis]]'' {{small|Sukhor.}}, with one species, ''Neomonolepis spathulata'', from western North America.<ref>Uotila, P., Sukhorukov, A.P., Bobon, N., McDonald, J., Krinitsina, A.A. and Kadereit, G. (2021), Phylogeny, biogeography and systematics of Dysphanieae (Amaranthaceae). ''Taxon'', 70: 526-551. {{doi|10.1002/tax.12458}}</ref>
** ''[[Suckleya]]'' A.Gray, with only one species:
*** ''Suckleya suckleyana'' (Torr.) Rydb., a succulent annual from western North America.
** ''[[Teloxys]]'' Moq.: with only one species:
*** ''Teloxys aristata'' (L.) Moq.<ref>{{GRIN|name = ''Teloxys aristata''| id = 465040 | accessdate = 26 November 2015}}</ref> (Syn.: ''Chenopodium aristatum'' L., ''Dysphania aristata''): from Eastern Europe to temperate Asia, naturalized elsewhere.

==Fossil record==
The oldest [[fossil]] records for Chenopodioideae are [[pollen]] grains recovered from [[Maastrichtian]] sediments of the [[Edmonton Formation]] in [[Canada]].<ref>Assorted angiosperm pollen from the Edmonton Formation (Maestrichtian), Alberta, Canada by Satish K. Srivastava  – Canadian Journal of Botany, 1969, 47(6): 975-989, {{doi|10.1139/b69-138}}</ref>

==References==
<references>
<ref name="Fuentes">Susy Fuentes-Bazan, Guilhem Mansion, Thomas Borsch (2012): ''Towards a species level tree of the globally diverse genus Chenopodium (Chenopodiaceae)''. In: ''Molecular Phylogenetics and Evolution'', '''62'''(1), pp. 359–374, {{issn|1055-7903}}, {{doi|10.1016/j.ympev.2011.10.006}}</ref>
<ref name="Zacharias">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), pp. 839-857. {{doi|10.1600/036364410X539907}}</ref>
</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.&nbsp;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.&nbsp;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.0061906}}

{{Commons category}}
{{Wikispecies|Chenopodioideae}}

== External links ==
* {{NCBI taxid|1307796|Chenopodioideae }}

{{Taxonbar|from=Q1070132}}

[[Category:Chenopodioideae| ]]
[[Category:Caryophyllales subfamilies]]