List of C4 plants

C4 plants use the C4 carbon fixation pathway to increase their photosynthetic efficiency by reducing or suppressing photorespiration, which mainly occurs under low atmospheric CO
2 concentration, high light, high temperature, drought, and salinity.[2][3] 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[4]) of flowering plants.[1] Among these are important crops such as maize, sorghum and sugarcane, but also weeds and invasive plants.[1] Although only 3% of flowering plant species use C4 carbon fixation, they account for 23% of global primary production.[5] The repeated, convergent C4 evolution from C3 ancestors has spurred hopes to bio-engineer the C4 pathway into C3 crops such as rice.[1][5]

Maize (Zea mays, Poaceae) is the most widely cultivated C4 plant.[1]

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.[1]

C4 plants are usually identified by their higher 13C/12C isotopic ratio compared to C3 plants or their typical leaf anatomy.[5] 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.[1]

The following list presents known C4 lineages by family, based on the overview by Sage (2016).[1] 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.[1]

Blepharis attenuata (Acanthaceae) grows in deserts.
Shadscale (Atriplex confertifolia, Amaranthaceae) is a halophytic shrub common in steppes of western North America.[1]
Black saxaul (Haloxylon ammodendron, Amaranthaceae) provided fuel for caravans following the Silk Road in Central Asia.[1]
Cleome gynandra (Cleomaceae) has been a C4 model plant.[1]
Papyrus (Cyperus papyrus, Cyperaceae) has been of major cultural importance.[1]
Spotted spurge (Euphorbia maculata, Euphorbiaceae) commonly grows in sidewalk cracks in North America.[6]
The aquatic Egeria densa (Hydrocharitaceae) uses the C4 pathway under high temperature and light intensity.
Sugarcane (Saccharum officinarum, Poaceae) is grown for sugar and bioethanol.[1]
Purslane (Portulaca oleracea, Portulacaceae), a weed and ancient vegetable, uses both C4 and CAM photosynthesis.[1][7]
Rhodes grass (Chloris gayana, Poaceae) is a major forage grass in tropical areas.[1]
The spiny fruits of puncture vine (Tribulus terrestris, Zygophyllaceae) may even punctuate tyres.[1]

Acanthaceae

The large acanthus family Acanthaceae includes one genus with C4 species, found in dry habitats from Africa to Asia.[8]

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:[9]

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.[1] Suaeda aralocaspica and species of the genus Bienertia use a particular, single-cell type of C4 carbon fixation.[1][10]

Asteraceae

The composite family Asteraceae contains three C4 lineages, in two different tribes of subfamily Asteroideae.[1][18] They include the model genus Flaveria with closely related C3, C4, and intermediate species.[1]

Boraginaceae

The borage family Boraginaceae contains one widespread C4 genus, Euploca, which has also been treated as part of a distinct family Heliotropiaceae.[22]

  • Euploca (also includes C3–C4 intermediates[23]) – 130 C4 species, 1–3 origins[1]

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.[1][24][25]

Caryophyllaceae

In the carnation family Caryophyllaceae, the C4 pathway evolved once, in a clade within the polyphyletic genus Polycarpaea.[1][26]

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.[1]

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).[1][6]

Gisekiaceae

Contains a C4 genus with a single species.

  • Gisekia pharnaceoides[27]

Hydrocharitaceae

Includes the only known aquatic C4 plants.[1]

Molluginaceae

The two C4 species within the same genus have acquired the pathway independently.

  • Mollugo – 2 C4 species, 2 origins

Nyctaginaceae

Polygonaceae

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.[1]

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.[1]

Scrophulariaceae
  • Anticharis – 4 C4 species

Zygophyllaceae
gollark: We could provide some kind of encoding for regular stuff in unprintables.
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gollark: The language name should be unprintable too in some weird way.
gollark: esoteric language idea: a regular language but run through gzip.
gollark: crates.io

References
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  2. Sage, R.F.; Sage, T.L.; Kocacinar, F. (2012). "Photorespiration and the evolution of C4 photosynthesis". Annual Review of Plant Biology. 63 (1): 19–47. doi:10.1146/annurev-arplant-042811-105511. ISSN 1543-5008. PMID 22404472.
  3. Christin, P.-A.; Osborne, C.P. (2014). "The evolutionary ecology of C4 plants" (PDF). New Phytologist. 204 (4): 765–781. doi:10.1111/nph.13033. ISSN 0028-646X.
  4. Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.
  5. Kellogg, E.A. (2013). "C4 photosynthesis". Current Biology. 23 (14): R594–R599. doi:10.1016/j.cub.2013.04.066. ISSN 0960-9822. PMID 23885869.
  6. Yang, Y.; Berry, P.E. (2011). "Phylogenetics of the Chamaesyce clade (Euphorbia, Euphorbiaceae): Reticulate evolution and long-distance dispersal in a prominent C4 lineage" (PDF). American Journal of Botany. 98 (9): 1486–1503. doi:10.3732/ajb.1000496. ISSN 0002-9122. PMID 21875975.
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  12. Kadereit, G.; Mavrodiev, E.V.; Zacharias, E.H.; Sukhorukov, A.P. (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): 1664–1687. doi:10.3732/ajb.1000169. ISSN 0002-9122. PMID 21616801.
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  14. Kapralov, M.V.; Akhani, H.; Voznesenskaya, E.V.; Edwards, G.; Franceschi, V.; Roalson, E.H. (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. 31 (3): 571–585. doi:10.1600/036364406778388674. ISSN 0363-6445.
  15. Pyankov, V.; Ziegler, H.; Kuz'min, A.; Edwards, G. (2001). "Origin and evolution of C4 photosynthesis in the tribe Salsoleae (Chenopodiaceae) based on anatomical and biochemical types in leaves and cotyledons". Plant Systematics and Evolution. 230 (1–2): 43–74. doi:10.1007/s006060170004. ISSN 0378-2697.
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  19. McKown, A.D.; Moncalvo, J.-M.; Dengler, N.G. (2005). "Phylogeny of Flaveria (Asteraceae) and inference of C4 photosynthesis evolution". American Journal of Botany. 92 (11): 1911–1928. doi:10.3732/ajb.92.11.1911. ISSN 0002-9122. PMID 21646108.
  20. Lyu, M.-J.A.; Gowik, U.; Kelly, S.; et al. (2015). "RNA-Seq based phylogeny recapitulates previous phylogeny of the genus Flaveria (Asteraceae) with some modifications". BMC Evolutionary Biology. 15 (1). doi:10.1186/s12862-015-0399-9. ISSN 1471-2148. PMC 4472175. PMID 26084484.
  21. Hansen, D.R. (2012). The molecular phylogeny of Pectis L. (Tageteae, Asteraceae), with implications for taxonomy, biogeography, and the evolution of C4 photosynthesis (Doctoral dissertation). University of Texas, Austin. Retrieved 2018-05-27.
  22. Luebert, F.; Cecchi, L.; Frohlich, M.W.; Gottschling, M.; Guilliams, C.M.; Hasenstab-Lehman, K.E.; Hilger, H.H.; Miller, J.S.; Mittelbach, M.; Nazaire, M.; Nepi, M.; Nocentini, D.; Ober, D.; Olmstead, R.G.; Selvi, F.; Simpson, M.G.; Sutorý, K.; Valdés, B.; Walden, G.K.; Weigend, M. (2016). "Familial classification of the Boraginales". Taxon. 65 (3): 502–522. doi:10.12705/653.5. ISSN 0040-0262. Retrieved 16 June 2018.
  23. Vogan, P.J.; Frohlich, M.W.; Sage, R.F. (2007). "The functional significance of C3–C4 intermediate traits in Heliotropium L. (Boraginaceae): gas exchange perspectives". Plant, Cell & Environment. 30 (10): 1337–1345. doi:10.1111/j.1365-3040.2007.01706.x. ISSN 0140-7791. PMID 17727423.
  24. Marshall, D.M.; Muhaidat, R.; Brown, N.J.; et al. (2007). "Cleome, a genus closely related to Arabidopsis, contains species spanning a developmental progression from C3 to C4 photosynthesis". The Plant Journal. 51 (5): 886–896. doi:10.1111/j.1365-313X.2007.03188.x. ISSN 0960-7412.
  25. Feodorova, T.A.; Voznesenskaya, E.V.; Edwards, G.E.; Roalson, E.H. (2010). "Biogeographic patterns of diversification and the origins of C4 in Cleome (Cleomaceae)" (PDF). Systematic Botany. 35 (4): 811–826. doi:10.1600/036364410X539880. ISSN 0363-6445. Retrieved 16 June 2016.
  26. Kool, A. (2012). Desert plants and deserted islands: systematics and ethnobotany in Caryophyllaceae (PhD thesis). Uppsala University. pp. 35–36. ISBN 978-91-554-8471-2. Retrieved 23 June 2018.
  27. Bissinger, K.; Khoshravesh, R.; Kotrade, J.P.; et al. (2014). "Gisekia (Gisekiaceae): Phylogenetic relationships, biogeography, and ecophysiology of a poorly known C4 lineage in the Caryophyllales". American Journal of Botany. 101 (3): 499–509. doi:10.3732/ajb.1300279. ISSN 0002-9122.
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