{{Short description|none}} {{DISPLAYTITLE:List of C<sub>4</sub> plants}} [[File:A_farmer_in_the_foreground_examining_the_stalks_and_looking_at_some_ears_of_corn_on_some_very_tall_plants,_Ontario_(20691586748).jpg|thumb|right|alt=A farmer among high maize plants|Maize (''Zea mays'', Poaceae) is the most widely cultivated {{C4}} plant.<ref name="Sage2016"/>]]
In botany, {{C4}} carbon fixation is one of three known methods of photosynthesis used by plants. {{C4}} plants increase their photosynthetic efficiency by reducing or suppressing photorespiration, which mainly occurs under low atmospheric {{CO2}} concentration, high light, high temperature, drought, and salinity.<ref name="SageSage2012"/><ref name="ChristinOsborne2014"/> There are roughly 8,100 known {{C4}} species, which belong to at least 61 distinct evolutionary lineages in 19 families (as per APG IV classification<ref name="APGIV"/>) of flowering plants.<ref name="Sage2016"/> Among these are important crops such as maize, sorghum and sugarcane, but also weeds and invasive plants.<ref name="Sage2016"/> Although only 3% of flowering plant species use {{C4}} carbon fixation, they account for 23% of global primary production.<ref name="Kellogg2013"/> The repeated, convergent {{C4}} evolution from {{C3}} ancestors has spurred hopes to bio-engineer the {{C4}} pathway into {{C3}} crops such as rice.<ref name="Sage2016"/><ref name="Kellogg2013"/>
{{C4}} photosynthesis probably first evolved 30–35 million years ago in the Oligocene, and further origins occurred since, most of them in the last 15 million years. {{C4}} plants are mainly found in tropical and warm-temperate regions, predominantly in open grasslands where they are often dominant. While most are graminoids, other growth forms such as forbs, vines, shrubs, and even some trees and aquatic plants are also known among {{C4}} plants.<ref name="Sage2016"/>
{{C4}} plants are usually identified by their higher <sup>13</sup>C/<sup>12</sup>C isotopic ratio compared to {{C3}} plants or their typical leaf anatomy.<ref name="Kellogg2013"/> The distribution of {{C4}} lineages among plants has been determined through phylogenetics and was considered well known {{as of|2016|lc=y}}. Monocots – mainly grasses (Poaceae) and sedges (Cyperaceae) – account for around 80% of {{C4}} species, but they are also found in the eudicots.<ref name="Sage2016"/> Moreover, almost all {{C4}} plants are herbaceus, with the notable exception of some woody species from the ''Euphorbia'' genus, such as the tree ''Euphorbia olowaluana''.<ref>{{Cite web |last=Elliott |first=Liam |date=2020-05-22 |title=To C4 or not to C4 if you're a tree? Some possible answers |url=https://botany.one/2020/05/to-c4-or-not-to-c4-if-youre-a-tree-some-possible-answers/ |access-date=2024-04-30 |website=Botany One |language=en-US}}</ref> The reason behind {{C4}} metabolism extreme rarity in trees is debated: hypotheses vary from a possible reduction in photosynthetic quantum yield under dense canopy conditions, coupled with an increased metabolic energy consumption (inherent to {{C4}} metabolism itself), to less efficient sunflecks utilization.<ref>{{Cite journal |url=https://academic.oup.com/jxb/article/71/16/4629/5837361 |access-date=2024-05-01 |journal=Journal of Experimental Botany |doi=10.1093/jxb/eraa234 |pmc=7410182 |pmid=32409834 |title=Why is C4 photosynthesis so rare in trees? |date=2020 |last1=Young |first1=Sophie N R. |last2=Sack |first2=Lawren |last3=Sporck-Koehler |first3=Margaret J. |last4=Lundgren |first4=Marjorie R. |volume=71 |issue=16 |pages=4629–4638 }}</ref>
The following list presents known {{C4}} lineages by family, based on the overview by Sage (2016).<ref name="Sage2016"/> They correspond to single species or clades thought to have acquired the {{C4}} pathway independently. In some lineages that also include {{C3}} and {{C3}}–{{C4}} intermediate species, the {{C4}} pathway may have evolved more than once.<ref name="Sage2016"/>
[[File:Blepharis_attenuata_1.jpg|thumb|right|alt=Spiny plant with pale violet flowers|''Blepharis attenuata'' (Acanthaceae) grows in deserts.]] [[File:Atriplex_confertifolia_(5063217522).jpg|thumb|right|alt=Dense bush with yellow flowers in dry habitat|Shadscale (''Atriplex confertifolia'', Amaranthaceae) is a halophytic shrub common in steppes of western North America.<ref name="Sage2016"/>]] [[File:Saxaul-baum.JPG|thumb|right|alt=Bush in very dry desert habitat|Black saxaul (''Haloxylon ammodendron'', Amaranthaceae) provided fuel for caravans following the Silk Road in Central Asia.<ref name="Sage2016"/>]] [[File:Starr 040323-0095 Cleome gynandra.jpg|thumb|right|alt=Plant with inflorescence of white flowers|''Cleome gynandra'' (Cleomaceae) has been a {{C4}} model plant.<ref name="Sage2016"/>]] [[File:Cyperus_papyrus_(Kafue_River).jpg|thumb|right|alt=Dense papyrus stand on river bank|Papyrus (''Cyperus papyrus'', Cyperaceae) has been of major cultural importance.<ref name="Sage2016"/>]] [[File:Chamaesyce maculata (L.) Small (AM AK360262-3).jpg|thumb|right|alt=Prostrate purple plants growing on sidewalk |Spotted spurge (''Euphorbia maculata'', Euphorbiaceae) commonly grows in sidewalk cracks in North America.<ref name="YangBerry2011"/>]] [[File:Egeria_densa_iceland.JPG|thumb|right|alt=Bright green foliage of aquatic plant|The aquatic ''Egeria densa'' (Hydrocharitaceae) uses the {{C4}} pathway under high temperature and light intensity.]] [[File:Caña de Azucar.jpg|thumb|right|alt=Cut sugarcane|Sugarcane (''Saccharum officinarum'', Poaceae) is grown for sugar and bioethanol.<ref name="Sage2016"/>]] [[File:20150610Portulaca_oleracea3.jpg|thumb|right|alt=Prostrate plant with shiny, fleshy leaves|Purslane (''Portulaca oleracea'', Portulacaceae), a weed and ancient vegetable, uses both {{C4}} and CAM photosynthesis.<ref name="Sage2016"/><ref name="KochKennedy1982"/>]] [[File:Chloris_gayana_habit4_(7069868575).jpg|thumb|right|alt=Grass with inflorescence in front of blue sky|Rhodes grass (''Chloris gayana'', Poaceae) is a major forage grass in tropical areas.<ref name="Sage2016"/>]] [[File:Starr_030612-0067_Tribulus_terrestris.jpg|thumb|alt=Spiny fruits developing on plant|The spiny fruits of puncture vine (''Tribulus terrestris'', Zygophyllaceae) may even puncture tyres.<ref name="Sage2016"/>]]
==Acanthaceae==
The large acanthus family Acanthaceae includes one genus with {{C4}} species, found in dry habitats from Africa to Asia.<ref name="FisherMcDade2015"/>
*''Blepharis'' – 15 {{C4}} species, 1–4 origins
==Aizoaceae==
While many species in the ice plant family Aizoaceae use crassulacean acid metabolism (CAM), one subfamily with drought-tolerant and halophytic plants includes {{C4}} species:<ref name="BohleyJoos2015"/>
*Sesuvioideae – 30 {{C4}} species, 1–6 origins
==Amaranthaceae==
The amaranth family Amaranthaceae (including the former goosefoot family Chenopodiaceae) contains around 800 known {{C4}} species, which belong to 14 distinct lineages in seven subfamilies. This makes Amaranthaceae the family with most {{C4}} species and lineages among the eudicots.<ref name="Sage2016"/> ''Suaeda aralocaspica'' and species of the genus ''Bienertia'' use a particular, single-cell type of {{C4}} carbon fixation.<ref name="Sage2016"/><ref name="SchützeFreitag2003"/>
*''Aerva'' (Amaranthoideae) – 4 {{C4}} species<ref name="SageSage2007"/> *''Alternanthera'' (Gomphrenoideae) – 17 {{C4}} species (also includes {{C3}}–{{C4}} intermediates)<ref name="SageSage2007"/> *''Amaranthus'' (Amaranthoideae) – 90 {{C4}} species<ref name="SageSage2007"/> *''Atriplex'' (Chenopodioideae) – around 180 {{C4}} species<ref name="KadereitMavrodiev2010"/> *''Bassia''–''Camphorosma'' clade (Camphorosmoideae) – 24 {{C4}} species (also includes one {{C3}}–{{C4}} intermediate), 1–2 origins<ref name="KadereitFreitag2011"/> *''Bienertia'' (Suaedoideae) – 3 {{C4}} species<ref name="SchützeFreitag2003"/><ref name="KapralovAkhani2006"/> *Caroxyleae (syn. Caroxyloneae, Salsoloideae) – 157 {{C4}} species<ref name="PyankovZiegler2001"/><ref name="AkhaniEdwards2007"/> *Gomphrenoids (Gomphrenoideae) – 138 {{C4}} species<ref name="SageSage2007"/> *Salsoleae (Salsoloideae) – 158 {{C4}} species, 2–4 origins<ref name="PyankovZiegler2001"/><ref name="AkhaniEdwards2007"/> *''Suaeda aralocaspica'' (Suaedoideae)<ref name="SchützeFreitag2003"/> *''Suaeda'' sect. ''Salsina'' – 30 {{C4}} species<ref name="SchützeFreitag2003"/> *''Suaeda'' sect. ''Schoberia'' – 9 {{C4}} species<ref name="SchützeFreitag2003"/> *''Tecticornia'' – (Salicornioideae) 2 {{C4}} species<ref name="VoznesenskayaAkhani2008"/> *''Tidestromia'' (Gomphrenoideae) – 8 {{C4}} species<ref name="SageSage2007"/>
==Asteraceae==
The composite family Asteraceae contains three {{C4}} lineages, in two different tribes of subfamily Asteroideae.<ref name="Sage2016"/><ref name="Kellogg1999"/> They include the model genus ''Flaveria'' with closely related {{C3}}, {{C4}}, and intermediate species.<ref name="Sage2016"/>
*''Flaveria'' (Tageteae) – 7 {{C4}} species, 2–3 origins (also includes {{C3}} and intermediate)<ref name="Sage2016"/><ref name="McKownMoncalvo2005"/><ref name="LyuGowik2015"/> *Coreopsideae – 41 {{C4}} species<ref name="Kellogg1999"/> *''Pectis'' (Tageteae) – 90 {{C4}} species<ref name="Hansen2012"/>
==Boraginaceae== The borage family Boraginaceae contains one widespread {{C4}} genus, ''Euploca'', which has also been treated as part of a distinct family Heliotropiaceae.<ref name="LuebertCecchi2016"/>
*''Euploca'' (also includes {{C3}}–{{C4}} intermediates<ref name="VoganFrohlich2007"/>) – 130 {{C4}} species, 1–3 origins<ref name="Sage2016"/>
==Cleomaceae==
The Cleomaceae, formerly included in the caper family Capparaceae, contains three {{C4}} species in genus ''Cleome''. These three species independently acquired the {{C4}} pathway; the genus also contains numerous {{C3}} as well as {{C3}}–{{C4}} intermediate species.<ref name="Sage2016"/><ref name="MarshallMuhaidat2007"/><ref name="FeodorovaVoznesenskaya2010"/>
*''Cleome angustifolia'' *''C. gynandra'' *''C. oxalidea''
==Caryophyllaceae==
In the carnation family Caryophyllaceae, the {{C4}} pathway evolved once, in a clade within the polyphyletic genus ''Polycarpaea''.<ref name="Sage2016"/><ref name="Kool2012"/>
*''Polycarpaea'' – 20 {{C4}} species
==Cyperaceae==
The sedge family Cyperaceae is second only to the grasses in number of {{C4}} species. Prominent {{C4}} sedges include culturally important species such as papyrus (''Cyperus papyrus'') and chufa (''C. esculentus'') but also purple nutsedge (''C. rotundus''), one of the world's major weeds. ''Eleocharis vivipara'' uses {{C3}} carbon fixation in underwater leaves and {{C4}} carbon fixation in aerial leaves.<ref name="Sage2016"/>
*''Bulbostylis'' – 211 {{C4}} species *''Cyperus'' – 757 {{C4}} species *''Eleocharis'' ser. ''Tenuissimae'' – 10 {{C4}} species *''Eleocharis vivipara'' *''Fimbristylis'' – 303 {{C4}} species *''Rhynchospora'' – 40 {{C4}} species
==Euphorbiaceae==
The spurge family Euphorbiaceae contains the largest single {{C4}} lineage among eudicots. The {{C4}} spurges are diverse and widespread; they range from weedy herbs to the only known {{C4}} trees – four species from Hawaii, including ''Euphorbia olowaluana'' (up to 10 m) and ''E. herbstii'' (up to 8 m).<ref name="Sage2016"/><ref name="YangBerry2011"/>
*''Euphorbia'' subgenus ''Chamaesyce'' section ''Anisophyllum'' (also treated as genus ''Chamaesyce'') – 350 {{C4}} species (also including {{C3}} and {{C3}}–{{C4}} intermediate species)
==Gisekiaceae==
Contains a {{C4}} genus with a single species.
*''Gisekia pharnaceoides''<ref name="BissingerKhoshravesh2014"/>
==Hydrocharitaceae==
Includes the only known aquatic {{C4}} plants.<ref name="Sage2016"/>
*''Egeria densa'' *''Hydrilla verticillata''
==Molluginaceae==
The two {{C4}} species within the same genus have acquired the pathway independently.
*''Mollugo'' – 2 {{C4}} species, 2 origins
==Nyctaginaceae==
*''Allionia'' – 2 {{C4}} species *''Boerhavia'' – 42 {{C4}} species
==Polygonaceae==
*''Calligonum'' – 80 {{C4}} species
==Portulacaceae== The single genus of this family forms one {{C4}} lineage. CAM photosynthesis is also known. Common purslane (''Portulaca oleracea'') is a major weed but also a vegetable.<ref name="Sage2016"/> *''Portulaca'' – 100 {{C4}} species, 1–2 origins
==Poaceae==
The grass family includes most of the known {{C4}} species – around 5000. They are only found in subfamilies of the PACMAD clade. Major {{C4}} crops such as maize, sugarcane, sorghum and pearl millet belong in this family. The only known species with {{C3}}, {{C4}} and intermediate variants, ''Alloteropsis semialata'', is a grass.<ref name="Sage2016"/>
*''Aristida'' – 288 {{C4}} species *''Stipagrostis'' – 56 {{C4}} species *Chloridoideae (without Centropodieae) – 1596 {{C4}} species *''Centropodia'' – 4 {{C4}} species *''Eriachne'' – 50 {{C4}} species *Tristachyideae – 87 {{C4}} species *Andropogoneae – 1228 {{C4}} species (incl. maize, sugarcane, sorghum) *''Reynaudia filiformis'' *''Axonopus'' – 90 {{C4}} species *''Paspalum'' – 379 {{C4}} species *''Anthaenantia'' – 4 {{C4}} species *Arthropoginae/''Mesosetum'' clade – 35 {{C4}} species, 1–2 origins *Arthropoginae/''Onchorachis'' clade – 2 {{C4}} species *Arthropoginae/''Colaeteania'' clade – 7 {{C4}} species, 1–2 origins *Anthephorinae – 286 {{C4}} species *''Echinochloa'' – 35 {{C4}} species *''Neurachne''–''Paraneurachne'' – 2 {{C4}} species, 2 origins *Melinidinae–Panicinae–Cenchrinae – 889 {{C4}} species *''Alloteropsis'' – 5 {{C4}} species, 1–2 origins (incl. {{C3}} and intermediate)
==Scrophulariaceae==
*''Anticharis'' – 4 {{C4}} species
==Zygophyllaceae==
*Tribuloideae – 37 {{C4}} species, 1–2 origins *''Tetraena simplex''
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}}
{{DEFAULTSORT:C4 plants}} Category:Photosynthesis Category:Lists of plants