Aizoaceae

The Aizoaceae Martynov, nom. cons. (fig-marigold family) is a large family of dicotyledonous flowering plants containing 135 genera and about 1800 species.[1] They are commonly known as ice plants or carpet weeds. They are often called vygies in South Africa and New Zealand. Highly succulent species that resemble stones are sometimes called mesembs.

Aizoaceae
Mesembryanthemum crystallinum
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Aizoaceae
Martinov
Type genus
Aizoon
L.
Genera

See text

Description
Mesembryanthemum guerichianum seedling, showing the epidermal bladder cells that inspired the name "ice plant".

The family Aizoaceae is widely recognised by taxonomists. It once went by the botanical name "Ficoidaceae", now disallowed. The APG II system of 2003 (unchanged from the APG system of 1998) also recognizes the family, and assigns it to the order Caryophyllales in the clade core eudicots. The APG II system also classes the former families Mesembryanthemaceae Fenzl, Sesuviaceae Horan. and Tetragoniaceae Link under the family Aizoaceae.

The common Afrikaans name "vygie" meaning "small fig" refers to the fruiting capsule, which resembles the true fig.[2] Glistening epidermal bladder cells give the family its common name "ice plants".[3]

Most species (96%, 1782 species in 132 genera) in this family are endemic to arid or semiarid parts of Southern Africa in the Succulent Karoo.[4] Much of the Aizoaceae's diversity is found in the Greater Cape Floristic Region, which is the most plant-diverse temperate region in the world.[5] A few species are found in Australia and the Central Pacific area.[6]

Most fig-marigolds are herbaceous, rarely somewhat woody, with sympodial growth and stems either erect or prostrate. Leaves are simple, opposite or alternate, and more or less succulent with entire (or rarely toothed) margins. Flowers are perfect in most species (but unisexual in some), actinomorphic, and appear singularly or in few-flowered cymes developing from the leaf axils. Sepals are typically five (3-8) and more or less connate (fused) below. True petals are absent. However, some species have numerous linear petals derived from staminodes.[7] The seed capsules have one to numerous seeds per cell and are often hygrochastic, dispersing seeds by "jet action" when wet.[3]

Evolution

The radiation of the Aizoaceae, specifically the subfamily Ruschioideae, was one of the most recent among the angiosperms, occurring 1.13-6.49 Mya. It is also one of the fastest radiations ever described in the angiosperms, with a diversification rate of about 4.4 species per million years.[8] This diversification was roughly contemporaneous with major radiations in two other succulent lineages, Cactaceae and Agave.[9]

The family includes many species that use crassulacean acid metabolism as pathway for carbon fixation. Some species in the subfamily Sesuvioideae instead use C4 carbon fixation, which might have evolved multiple times in the group.[10]

Uses
Tetragonia tetragonoides ("New Zealand spinach")

Several genera are cultivated. Lithops, or "living stones", are popular as novelty house plants because of their stone-like appearance.

Some species are edible, including:

C. edulis was introduced to California in the early 1900s to stabilize soil along railroad tracks and has become invasive.[13] In southern California, ice plants are sometimes used as firewalls;[14] however, they do burn if not carefully maintained.[15]

Taxonomy
Aptenia cordifolia or rock rose

Because of the hyperdiversity of the Aizoaceae and the young age of the clade, many generic and species boundaries are uncertain.[8]

Carpobrotus edulis, an "ice plant"
Cheiridopsis denticulata
Drosanthemum speciosum
Fenestraria rhopalophylla
Jensenobotrya lossowiana
Lithops bromfieldii
Sesuvium portulacastrum

Subfamily Aizooideae

Genera:[16]

Subfamily Mesembryanthemoideae

Genera:[17]

  • Phyllobolus N.E.Br
  • Prenia N.E.Br
  • Pseudobrownanthus Ihlenf. & Bittrich
  • Psilocaulon N.E.Br
  • Sceletium N.E.Br
  • Synaptophyllum N.E.Br

Subfamily Ruschioideae

Genera:

Tribe Apatesieae[18]
  • Apatesia N.E.Br
  • Carpanthea N.E.Br
  • Caryotophora Leistner
  • Conicosia N.E.Br
  • Hymenogyne Haw.
  • Saphesia N.E.Br
  • Skiatophytum L.Bolus
Tribe Dorotheantheae[19]

* no longer recognised

Tribe Ruschieae[20]

Subfamily Sesuvioideae

This subfamily includes a number of C4 species.[10]

Genera:[21]

Footnotes
  1. Hartmann, HEK (2001). Illustrated Handbook of Succulent Plants. Berlin, Germany: Springer.
  2. "The Living Stone Page". The Succulent Plant Page. Retrieved 23 June 2017.
  3. "Angiosperm Phylogeny Website". www.mobot.org. Retrieved 2017-12-20.
  4. Chesselet, P., Smith, G.F., Burgoyne, P.M., Klak, C., Hammer, S.A., Hartmann, H.E.K., Kurzweil, H., van Jaarsveld, E.J., van Wyk, B-E. & Leistner, O.A (2000). "Seed Plants of Southern Africa". Strelitzia. 10: 360–410.CS1 maint: uses authors parameter (link)
  5. Born, J.; Linder, H. P.; Desmet, P. (2007). "The Greater Cape Floristic Region". Journal of Biogeography. 34 (1): 147–162. doi:10.1111/j.1365-2699.2006.01595.x. JSTOR 4125143.
  6. "Browsing: Aizoaceae". World of Succulents. Retrieved 23 June 2017.
  7. Watson, L., and Dallwitz, M.J. (1992). "The families of flowering plants: descriptions, illustrations, identification, and information retrieval". 20 June 2017. Retrieved 9 July 2017.CS1 maint: uses authors parameter (link)
  8. Valente, Luis M.; Britton, Adam W.; Powell, Martyn P.; Papadopulos, Alexander S. T.; Burgoyne, Priscilla M.; Savolainen, Vincent (2014-01-01). "Correlates of hyperdiversity in southern African ice plants (Aizoaceae)". Botanical Journal of the Linnean Society. 174 (1): 110–129. doi:10.1111/boj.12117. ISSN 0024-4074. PMC 4373134. PMID 25821244.
  9. Arakaki, Mónica; Christin, Pascal-Antoine; Nyffeler, Reto; Lendel, Anita; Eggli, Urs; Ogburn, R. Matthew; Spriggs, Elizabeth; Moore, Michael J.; Edwards, Erika J. (2011). "Contemporaneous and recent radiations of the world's major succulent plant lineages". Proceedings of the National Academy of Sciences of the United States of America. 108 (20): 8379–8384. doi:10.1073/pnas.1100628108. JSTOR 25830059. PMC 3100969. PMID 21536881.
  10. Bohley, Katharina; Joos, Olga; Hartmann, Heidrun; Sage, Rowan; Liede-Schumann, Sigrid; Kadereit, Gudrun (2015). "Phylogeny of Sesuvioideae (Aizoaceae) – Biogeography, leaf anatomy and the evolution of C4 photosynthesis". Perspectives in Plant Ecology, Evolution and Systematics. 17 (2): 116–130. doi:10.1016/j.ppees.2014.12.003. ISSN 1433-8319.
  11. Facciola. S. (1990). Cornucopia - A Source Book of Edible Plants. Kampong Publications. ISBN 0-9628087-0-9.
  12. Low. T. (1989). Wild Food Plants of Australia. Angus and Robertson. ISBN 0-207-14383-8.
  13. "Invasive Plants of California's Wildland". California Invasive Plant Council. Retrieved 23 June 2017.
  14. "Fire Safe Landscaping". Cal Fire. Archived from the original on 2 July 2017. Retrieved 23 June 2017.
  15. Baldwin, Debra Lee. "Firewise Landscaping with Succulents - How succulents saved a Rancho Santa Fe home from wildfire". Retrieved 23 June 2017.
  16. "GRIN Genera of Aizoaceae subfam. Aizooideae". Germplasm Resources Information Network. Archived from the original on 2012-07-19. Retrieved 2011-03-04.
  17. "GRIN Genera of Aizoaceae subfam. Mesembryanthemoideae". Germplasm Resources Information Network. Archived from the original on 2012-07-18. Retrieved 2011-03-04.
  18. "GRIN Genera of Aizoaceae tribe Apatesieae". Germplasm Resources Information Network. Archived from the original on 2015-09-24. Retrieved 2011-03-04.
  19. "GRIN Genera of Aizoaceae tribe Dorotheantheae". Germplasm Resources Information Network. Archived from the original on 2012-07-24. Retrieved 2011-03-04.
  20. "GRIN Genera of Aizoaceae tribe Ruschiae". Germplasm Resources Information Network. Archived from the original on 2012-07-20. Retrieved 2011-03-04.
  21. "GRIN Genera of Aizoaceae subfam. Sesuvioideae". Germplasm Resources Information Network. Archived from the original on 2012-07-18. Retrieved 2011-03-04.
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References
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