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| − | '''Amaranthaceae''' is a family of [[flowering plants]] known as the [[amaranth]] family. It now includes the former goosefoot family Chenopodiaceae, and contains about 165 genera and 2,040 species.<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press }}</ref><ref name="APW" /> making it the most species-rich lineage within the [[flowering plant]] [[order (biology)|order]] of [[Caryophyllales]]. | + | '''Amaranthaceae''' is a family of [[flowering plants]] known as the [[amaranth]] family. It now includes the former goosefoot family Chenopodiaceae, and contains a high diversity of edible plants, from supermarket staples to nutritious "weeds". |
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| − | == Description ==
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| − | === Vegetative characters ===
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| − | [[File:Illustration Bassia laniflora0 clean.JPG|thumb|left|''[[Bassia laniflora]]'' (illustration), [[Camphorosmoideae]]]]
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| − | Most of these species are [[Annual plant|annual]] or [[perennial]] [[herb]]s or [[subshrub]]s, some are [[shrub]]s; very few species are [[vine]]s or [[tree]]s. Some species are [[succulent]]. Many species have stems with thickened nodes. The wood of the perennial stem has a typical "anomalous" [[secondary growth]], only in subfamily [[Polycnemoideae]] is secondary growth normal.<ref name="APW" />
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| − | The [[Leaf|leaves]] are mostly alternate, sometimes opposite. They never possess [[stipule]]s. The simple leaves are flat or terete, their shape is extremely variable, with entire or toothed margins. In some species, the leaves are reduced to minute scales. In most cases, neither basal or terminal aggregations of leaves occur.<ref name="APW" />
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| − | [[File:Nitrophila occidentalis 7.jpg|thumb|left|Flower of ''[[Nitrophila occidentalis]]'', [[Polycnemoideae]]]]
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| − | [[File:Halothamnus Pollen 1.JPG|thumb|Pollen grains of ''[[Halothamnus glaucus]]'']]
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| − | === Inflorescence and flowers ===
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| − | [[File:Silver Cockscomb (Celosia argentea) in Tirunelveli.jpg|right|thumb|Flower of Silver Cockscomb ''[[Celosia argentea]]'' in [[Tirunelveli]], India]]
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| − | The [[flower]]s are solitary or aggregated in [[Cyme (botany)|cymes]], [[Raceme|spikes]], or [[panicle]]s and typically perfect (bisexual) and [[actinomorphic]]. Some species have unisexual flowers. [[Bract]]s and [[bracteole]]s are either herbaceous or scarious. Flowers are regular with a herbaceous or scarious perianth of (one to) mostly five (rarely to eight) [[tepal]]s, often joined. One to five [[stamen]]s are opposite to tepals or alternating, inserting from a hypogynous disc, which may have appendages (pseudo[[staminodes]]) in some species. The [[anther]]s have two or four pollen sacs (locules). In tribe Caroxyloneae, antheres have vesicular appendages. The [[pollen]] grains are spherical with many pores (pantoporate), with pore numbers from a few to 250 (in ''[[Froelichia]]'').<ref name="Müller2005" /> One to three (rarely six) [[carpel]]s are fused to a superior [[ovary (plants)|ovary]] with one (rarely two) basal ovule.<ref name="APW" />
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| − | === Fruits and seeds ===
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| − | The [[Diaspore (botany)|diaspore]]s are [[seed]]s or [[fruit]]s ([[utricle (fruit)|utricles]]), more often the perianth persists and is modified in fruit for means of dispersal. Sometimes even bracts and bracteoles may belong to the diaspore. More rarely the fruit is a circumscissile [[Capsule (fruit)|capsule]] or a [[berry (botany)|berry]]. The horizontal or vertical seed often has a thickened or woody seed coat. The green or white embryo is either spirally (and without [[perisperm]]) or annular (rarely straight).<ref name="APW" />
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| − | === Chromosome number ===
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| − | The basic [[chromosome]] number is (rarely 6) mostly 8-9 (rarely 17).<ref name="APW" />
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| − | === Phytochemistry ===
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| − | Widespread in the Amaranthaceae is the occurrence of [[betalain]] pigments. The former Chenopodiaceae often contain [[isoflavonoid]]s.<ref name="APW" />
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| − | In phytochemical research, several [[methylenedioxyflavonols]], [[saponins]], [[triterpenoids]], [[ecdysteroids]], and specific root-located [[carbohydrates]] have been found.<ref name="Müller2005" />
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| − | === Photosynthesis pathway ===
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| − | With around 800 species that are [[C4 carbon fixation|C<sub>4</sub>-plants]], the Amaranthaceae represent the largest group with this [[photosynthesis]] pathway among the eudicots (1600 C<sub>4</sub>-species).<ref name="Kadereit2003" /> Within the family, several types of C<sub>4</sub>-photosynthesis occur, and about 17 different types of leaf anatomy are realized. Therefore, this photosynthesis pathway seems to have been developed about 15 times independently during the evolution of the family. About two-thirds of the C<sub>4</sub>-species belong to the former Chenopodiaceae. The first occurrence of C<sub>4</sub>-photosynthesis dates from the early [[miocene]], about 24 million years ago, but in some groups, this photosynthesis pathway has evolved much later, about 6 (or less) million years ago.<ref name="Kadereit2003" />
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| − | The multiple origin of C<sub>4</sub>-photosynthesis in the Amaranthaceae is regarded as an evolutionary response to inexorably decreasing atmospheric CO2 levels, coupled with a more recent permanent shortage in water supply in combination with high temperatures. These species with their higher water use efficiency had a selective advantage and were able to spread out into dry (arid) habitats.<ref name="Kadereit2003" />
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| − | == Distribution ==
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| − | This is a widespread and [[cosmopolitan family]] from the tropics to cool temperate regions. The Amaranthaceae (sensu stricto) are predominantly tropical, whereas the former Chenopodiaceae have their centers of diversity in dry temperate and warm temperate areas.<ref name="Müller2005" /> Many of the species are [[halophyte]]s, tolerating [[sodium chloride|salty]] soils, or grow in dry steppes or semideserts.
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| − | == Economic importance ==
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| | Some species, such as [[spinach]] (''[[Spinacia oleracea]]'') or forms of [[Beta (plant)|beet]] (''[[Beta vulgaris]]'') ([[beetroot]], [[chard]]), are used as [[vegetable]]s. Forms of ''Beta vulgaris'' include fodder beet (''[[Mangelwurzel]]'') and [[sugar beet]]. The seeds of ''[[Amaranthus]]'', [[lamb's quarters]] (''Chenopodium berlandieri''), [[quinoa]] (''Chenopodium quinoa'') and [[kañiwa]] (''Chenopodium pallidicaule'') are edible and are used as [[pseudocereals]]. | | Some species, such as [[spinach]] (''[[Spinacia oleracea]]'') or forms of [[Beta (plant)|beet]] (''[[Beta vulgaris]]'') ([[beetroot]], [[chard]]), are used as [[vegetable]]s. Forms of ''Beta vulgaris'' include fodder beet (''[[Mangelwurzel]]'') and [[sugar beet]]. The seeds of ''[[Amaranthus]]'', [[lamb's quarters]] (''Chenopodium berlandieri''), [[quinoa]] (''Chenopodium quinoa'') and [[kañiwa]] (''Chenopodium pallidicaule'') are edible and are used as [[pseudocereals]]. |
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| | ''[[Dysphania ambrosioides]]'' (epazote) and ''Dysphania anthelmintica'' are used as [[medicinal herbs]]. | | ''[[Dysphania ambrosioides]]'' (epazote) and ''Dysphania anthelmintica'' are used as [[medicinal herbs]]. |
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| − | Several species are used indirectly as a source of [[soda ash]], such as [[Salicornia]] (see [[Glasswort]]).
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| − | A number of species are popular garden [[ornamental plant]]s, especially species from ''[[Alternanthera]]'', ''[[Amaranthus]]'', ''[[Celosia]]'', and ''[[Iresine]]''.
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| − | Some species are considered [[weed]]s, e.g., redroot pigweed (''[[Amaranthus retroflexus]]'') and alligatorweed (''[[Alternanthera philoxeroides]]'').
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| − | Many species may cause [[Allergic rhinitis|pollen allergies]].<ref>[http://www.pollenlibrary.com/FAMILY/CHENOPODIACEAE/ List of allergic plants in Chenopodiaceae family at pollenlibrary.com]</ref>
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| | == Systematics == | | == Systematics == |
| | [[File:Cladogramm Amaranthaceae.jpg|thumb|left|280px|Cladogram of Amaranthaceae ''s.l.'', modified and simplified, based on phylogenetic research of Müller & Borsch 2005, Kadereit ''et al.'' 2006, Sanchez del-Pino'' et al.'' 2009]] | | [[File:Cladogramm Amaranthaceae.jpg|thumb|left|280px|Cladogram of Amaranthaceae ''s.l.'', modified and simplified, based on phylogenetic research of Müller & Borsch 2005, Kadereit ''et al.'' 2006, Sanchez del-Pino'' et al.'' 2009]] |
| − | [[File:A splendens.jpg|thumb|''[[Achyranthes splendens]]'', Amaranthoideae]]
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| − | [[File:Gomphrena arborescens-1.jpg|thumb|''[[Gomphrena arborescens]]'', Gomphrenoideae]]
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| − | [[File:Suikerbiet bloeiwijze Beta vulgaris.jpg|thumb|Sugar beet, ''[[Beta vulgaris]]'' subsp. ''vulgaris'' 'altissima', Betoideae]]
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| − | [[File:Grayia spinosa 1.jpg|thumb|''[[Grayia spinosa]]'', Chenopodioideae]]
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| − | [[File:Arthrocnemum perenne.jpg|thumb|''[[Sarcocornia perennis]]'', Salicornioideae]]
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| − | [[File:Salsola kali flowers.jpg|thumb|''[[Kali turgidum]]'' (Syn. ''Salsola kali'' subsp. ''kali''), Salsoloideae]]
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| − | [[File:Suaeda nigra 2.jpg|thumb|''[[Suaeda nigra]]'', Suaedoideae]]
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| − | In the [[APG II system]] of 2003 (unchanged from the [[APG system]], of 1998), the family is placed in the order [[Caryophyllales]]. It includes the plants formerly treated as the family [[Chenopodiaceae]]. The monophyly of this new, broadly defined Amaranthaceae has been strongly supported by both morphological and [[phylogenetic]] analyses.<ref name="Judd" />
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| − | The family Amaranthaceae was first published in 1789 by [[Antoine Laurent de Jussieu]] in ''Genera Plantarum'', p. 87–88. The first publication of family Chenopodiaceae was in 1799 by [[Étienne Pierre Ventenat]] in ''Tableau du Regne Vegetal'', 2, p. 253. The older name has priority and is now the valid scientific name of the extended Amaranthaceae (''s.l.'' = ''sensu lato'').
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| − | Many recent publications still refer to the family name Chenopodiaceae.<ref name="Akhani" /><ref name="Kadereit2006Beto" /><ref name="Kadereit2006Salic" /><ref name="Kapralov" /><ref name="Kadereit2010" /><ref name="Kadereit2011" /> Phylogenetic research revealed the important impact of the subfamily [[Polycnemoideae]] on the classification (see Cladogram): If Polycnemoideae are considered being part of Chenopodiaceae, then Amaranthaceae (''s.str.'' = ''sensu stricto'') have to be included, too, and the name of the extended family is Amaranthaceae. If Polycnemoideae would be separated as an own family, Chenopodiaceae and Amaranthaceae (''s.str.'') would form two distinct [[monophyletic]] groups and could be treated as two separate families.
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| − | Amaranthaceae Juss. (''s.l.'') include the former families Achyranthaceae Raf., Atriplicaceae Durande, Betaceae Burnett, Blitaceae T.Post & Kuntze, Celosiaceae Martynov, Chenopodiaceae Vent., ''nom. cons.', Corispermaceae Link, Deeringiaceae J.Agardh, Dysphaniaceae (Pax) Pax, ''nom. cons.'', Gomphrenaceae Raf., Polycnemaceae Menge, Salicorniaceae Martynov, Salsolaceae Menge, and Spinaciaceae Menge.
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| − | The systematics of Amaranthaceae are the subject of intensive recent research. Molecular genetic studies revealed the traditional classification, based on morphological and anatomical characters, often did not reflect the phylogenetical relationships.
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| − | The former Amaranthaceae (in their narrow circumscription) are classified in two subfamilies, [[Amaranthoideae]] and [[Gomphrenoideae]], and contain about 65 genera and 900 species in tropical [[Africa]] and [[North America]]. The Amaranthoideae and some genera of Gomphrenoideae were found to be [[polyphyletic]], so taxonomical changes are needed.<ref name="Sanchez del-Pino 2009" />
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| − | Current studies classified the species of former Chenopodiaceae to eight distinct subfamilies (the research is not yet completed): [[Polycnemoideae]],<ref name="Müller2005" /><ref name="Masson" /> which are regarded as a basal lineage, [[Betoideae]],<ref name="Kadereit2006Beto" /> [[Camphorosmoideae]],<ref name="Kadereit2011" /> [[Chenopodioideae]],<ref name="Kadereit2010" /> [[Corispermoideae]],<ref name="Sukhorukov" /> [[Salicornioideae]],<ref name="Kadereit2006Salic"/> [[Salsoloideae]],<ref name="Akhani" /> and [[Suaedoideae]].<ref name="Schütze2003" />
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| − | In this preliminary classification, the Amaranthaceae ''s.l.'' are divided into 10 subfamilies with approximately 180 genera and 2,500 species.<ref name="Müller2005" />
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| | === Genera === | | === Genera === |
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| − | == References ==
| + | {{Ack-Wikipedia}} |
| − | {{Reflist|2|refs= | |
| − | <ref name="Akhani">
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| − | Hossein Akhani, Gerald Edwards, Eric H. Roalson (2007): Diversification Of The Old World Salsoleae s.l. (Chenopodiaceae): Molecular Phylogenetic Analysis Of Nuclear And Chloroplast Data Sets And A Revised Classification. - ''International Journal of Plant Sciences'', '''168'''(6), p.931–956.</ref>
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| − | <ref name="APW">
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| − | [http://www.mobot.org/MOBOT/Research/APweb/orders/caryophyllalesweb.htm#Amaranthaceae The Amaranthaceae family] at [http://www.mobot.org/MOBOT/Research/APweb/welcome.html APWebsite.]</ref>
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| − | <ref name="Judd">
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| − | Judd et al. (2008). Plant Systematics: A Phylogenetic Approach, Third Edition. Sinauer Associates, Inc. Sunderland, MA</ref>
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| − | <ref name="Kadereit2003">
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| − | Gudrun Kadereit, Thomas Borsch, Kurt Weising, Helmut Freitag (2003): Phylogeny of Amaranthaceae and Chenopodiaceae and the evolution of C<sub>4</sub> photosynthesis. - ''International Journal of Plant Sciences'', Volume '''164''' (6), p.959–986.</ref>
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| − | <ref name="Kadereit2006Beto">
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| − | G. Kadereit, S. Hohmann, J.W. Kadereit (2006): A synopsis of Chenopodiaceae subfam. Betoideae and notes on the taxonomy of Beta. - ''Willdenowia'' '''36''', p.9-19.</ref>
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| − | <ref name="Kadereit2006Salic">
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| − | Gudrun Kadereit, Ladislav Mucina, Helmut Freitag (2006): Phylogeny of Salicornioideae (Chenopodiaceae): diversification, biogeography, and evolutionary trends in leaf and flower morphology. - ''Taxon'' '''55'''(3), p. 617–642.</ref>
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| − | <ref name="Kadereit2010">
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| − | Gudrun Kadereit, Evgeny V. Mavrodiev, Elizabeth H. Zacharias, Alexander P. Sukhorukov (2010): Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 Photosynthesis. - ''American Journal of Botany'' '''97'''(10), p. 1664-1687.</ref>
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| − | <ref name="Kadereit2011">
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| − | Gudrun Kadereit, Helmut Freitag (2011): Molecular phylogeny of Camphorosmeae (Camphorosmoideae, Chenopodiaceae): Implications for biogeography, evolution of C4-photosynthesis and taxonomy. - ''Taxon'' '''60'''(1), p. 51-78</ref>
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| − | <ref name="Kapralov">
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| − | Maxim V. Kapralov, Hossein Akhani, Elena V. Voznesenskaya, Gerald Edwards, Vincent Franceschi, Eric H. Roalson (2006): Phylogenetic Relationships in the Salicornioideae / Suaedoideae / Salsoloideae s.l. (Chenopodiaceae) Clade and a Clarification of the Phylogenetic Position of Bienertia and Alexandra Using Multiple DNA Sequence Datasets. - ''Systematic Botany''.</ref>
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| − | <ref name="Masson">
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| − | Rüdiger Masson & Gudrun Kadereit (2013): Phylogeny of Polycnemoideae (Amaranthaceae): Implications for biogeography, character evolution and taxonomy. ''Taxon'' 62 (1): 100-111. [http://www.ingentaconnect.com/content/iapt/tax/2013/00000062/00000001/art00009]</ref>
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| − | <ref name="Müller2005">
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| − | 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.</ref>
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| − | <ref name="Sanchez del-Pino 2009">
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| − | Ivonne Sánchez del-Pino, Thomas Borsch, Timothy J. Motle (2009): trnL-F and rpl16 Sequence Data and Dense Taxon Sampling Reveal Monophyly of Unilocular Anthered Gomphrenoideae (Amaranthaceae) and an Improved Picture of Their Internal Relationships. - ''Systematic Botany'', Volume '''34''' (1), p. 57-67. {{DOI|10.1600/036364409787602401}}</ref>
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| − | <ref name="Schütze2003">
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| − | Peter Schütze, Helmut Freitag, Kurt Weising (2003): An integrated molecular and morphological study of the subfamily Suaedoideae Ulbr. (Chenopodiaceae). - ''Plant Systematics and Evolution'', Volume '''239''', p. 257-286. abstract: {{DOI|10.1007/s00606-003-0013-2|}}</ref>
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| − | <ref name="Sukhorukov">
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| − | Alexander P. Sukhorukov (2007): Fruit anatomy and its taxonomic significance in Corispermum (Corispermoideae, Chenopodiaceae). – ''Willdenowia'' '''37''', {{ISSN|0511-9618}}, p.63-87, {{doi|10.3372/wi.37.37103}}</ref>
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| − | ==External links==
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| − | {{commons category|Amaranthaceae}}
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| − | {{wikispecies|Amaranthaceae}}
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| − | *[http://www.mobot.org/MOBOT/Research/APweb/orders/caryophyllalesweb.htm#Amaranthaceae The Amaranthaceae family] at [http://www.mobot.org/MOBOT/Research/APweb/welcome.html APWebsite.]
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| − | * [http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?48 Genera of ''Amaranthaceae''] at [[Germplasm Resources Information Network|Germplasm Resources Information Network (USDA)]]
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| − | * {{ITIS|id=20714|taxon=Amaranthaceae}}
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| − | * {{NCBI|3563|Amaranthaceae }}
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| − | * [http://www.tropicos.org/Name/42000073 Amaranthaceae] at Tropicos
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| − | * [http://www.iucnredlist.org/search IUCN link: Amaranthaceae threatened species]
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| − | * Stanley L. Welsh, Clifford W. Crompton & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10185 Chenopodiaceae] in Flora of North America
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| − | * Kenneth R. Robertson & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10031 Amaranthaceae] in Flora of North America
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| − | * Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10185 Chenopodiaceae] in Flora of China
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| − | * Bojian Bao, Thomas Borsch & Steven E. Clemants (2003): [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10031 Amaranthaceae] in Flora of China
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| − | {{taxonbar}}
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| | [[Category:Amaranthaceae| ]] | | [[Category:Amaranthaceae| ]] |
| | [[Category:Caryophyllales families]] | | [[Category:Caryophyllales families]] |
