List of C4 plants

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. 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) of flowering plants. Among these are important crops such as maize, sorghum and sugarcane, but also weeds and invasive plants. Although only 3% of flowering plant species use C4 carbon fixation, they account for 23% of global primary production. The repeated, convergent C4 evolution from C3 ancestors has spurred hopes to bio-engineer the C4 pathway into C3 crops such as rice. 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. C4 plants are usually identified by their higher 13C/12C isotopic ratio compared to C3 plants or their typical leaf anatomy. The distribution of C4 lineages among plants has been determined through phylogenetics and was considered well known as of 2016. Monocots – mainly grasses (Poaceae) and sedges (Cyperaceae) – account for around 80% of C4 species, but they are also found in the eudicots. 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. 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. The following list presents known C4 lineages by family, based on the overview by Sage (2016). 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.

Acanthaceae The large acanthus family Acanthaceae includes one genus with C4 species, found in dry habitats from Africa to Asia.

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:

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. Suaeda aralocaspica and species of the genus Bienertia use a particular, single-cell type of C4 carbon fixation.

Aerva (Amaranthoideae) – 4 C4 species Alternanthera (Gomphrenoideae) – 17 C4 species (also includes C3–C4 intermediates) Amaranthus (Amaranthoideae) – 90 C4 species Atriplex (Chenopodioideae) – around 180 C4 species Bassia–Camphorosma clade (Camphorosmoideae) – 24 C4 species (also includes one C3–C4 intermediate), 1–2 origins Bienertia (Suaedoideae) – 3 C4 species Caroxyleae (syn. Caroxyloneae, Salsoloideae) – 157 C4 species Gomphrenoids (Gomphrenoideae) – 138 C4 species Salsoleae (Salsoloideae) – 158 C4 species, 2–4 origins Suaeda aralocaspica (Suaedoideae) Suaeda sect. Salsina – 30 C4 species Suaeda sect. Schoberia – 9 C4 species Tecticornia – (Salicornioideae) 2 C4 species Tidestromia (Gomphrenoideae) – 8 C4 species

Asteraceae The composite family Asteraceae contains three C4 lineages, in two different tribes of subfamily Asteroideae. They include the model genus Flaveria with closely related C3, C4, and intermediate species.

Flaveria (Tageteae) – 7 C4 species, 2–3 origins (also includes C3 and intermediate) Coreopsideae – 41 C4 species Pectis (Tageteae) – 90 C4 species

Boraginaceae The borage family Boraginaceae contains one widespread C4 genus, Euploca, which has also been treated as part of a distinct family Heliotropiaceae.

Euploca (also includes C3–C4 intermediates) – 130 C4 species, 1–3 origins

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.

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.

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.

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).

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

Hydrocharitaceae Includes the only known aquatic C4 plants.

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

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