Biology:Aizoaceae

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The Aizoaceae (/ˌeɪzoʊˈeɪsi.iː, -ˌaɪ/), or fig-marigold family, are a large family of dicotyledonous flowering plants containing 135 genera and about 1,800 species.[1] Several genera are commonly named ice plants or carpet weeds. The Aizoaceae are also referred to as vygies in South Africa. Some of the unusual Southern African genera—such as Conophytum, Lithops, Titanopsis, and Pleiospilos (among others)—resemble gemstones, rocks, or pebbles, and are sometimes called living stones or mesembs (short for mesembryanthemums).

Description

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

File:LL-Q14196 (afr)-Oesjaar-vygie.wav 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 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.[4] The seed capsules have one to numerous seeds per cell and are often hygrochastic, dispersing seeds by "jet action" when wet.[3]

Distribution

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.[5] 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.[6] A few species are found in Australia and the Central Pacific area.[7]

Carpobrotus is found as an introduced species on the western coast of the United States, New Zealand, the Mediterranean coast of Europe and the southern coast of Brazil.

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]

Taxonomy

Mesembryanthemum cordifolium 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 Acrosanthoideae

Genera:[11]

Subfamily Aizooideae

Genera:[12]

Subfamily Mesembryanthemoideae

Genera:[13]

  • Aptenia N.E.Br, synonym of Mesembryanthemum
  • Aridaria N.E.Br, synonym of Mesembryanthemum
  • Aspazoma N.E.Br, synonym of Mesembryanthemum
  • Brownanthus Schwantes, synonym of Mesembryanthemum
  • Dactylopsis N.E.Br, synonym of Mesembryanthemum
  • Mesembryanthemum L.
  • Phyllobolus N.E.Br, synonym of Mesembryanthemum
  • Prenia N.E.Br, synonym of Mesembryanthemum
  • Psilocaulon N.E.Br, synonym of Mesembryanthemum
  • Synaptophyllum N.E.Br, synonym of Mesembryanthemum

Subfamily Ruschioideae

Genera:

Tribe Apatesieae[14]
Tribe Dorotheantheae[15]
  • Aethephyllum N.E.Br, synonym of Cleretum
  • Cleretum N.E.Br
  • Dorotheanthus Schwantes, synonym of Cleretum
Tribe Ruschieae[16]

Subfamily Sesuvioideae

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

Genera:[19]

Tribe Anisostigmateae[20]
Tribe Sesuvieae

Unplaced genera

Include;

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.[23] In southern California, ice plants are sometimes used as firewalls;[24] however, they do burn if not carefully maintained.[25]

References

  1. ↑ Hartmann, HEK (2001). Illustrated Handbook of Succulent Plants. Berlin, Germany: Springer. 
  2. ↑ "The Living Stone Page". The Succulent Plant Page. http://succulent-plant.com/families/aizoaceae.html. 
  3. ↑ 3.0 3.1 "Angiosperm Phylogeny Website". http://www.mobot.org/MOBOT/research/APweb/. 
  4. ↑ "The families of flowering plants: descriptions, illustrations, identification, and information retrieval". 1992. http://delta-intkey.com/angio/www/aizoacea.htm. 
  5. ↑ Chesselet, P.; Smith, G.F.; Burgoyne, P.M. et al. (2000). "Seed Plants of Southern Africa". Strelitzia 10: 360–410. 
  6. ↑ 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. 
  7. ↑ "Browsing: Aizoaceae". World of Succulents. http://worldofsucculents.com/?families=aizoaceae. 
  8. ↑ 8.0 8.1 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. PMID 25821244. 
  9. ↑ Arakaki, Mónica; Christin, Pascal-Antoine; Nyffeler, Reto; Lendel, Anita; Eggli, Urs; Ogburn, R. Matthew; Spriggs, Elizabeth; Moore, Michael J. et al. (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. PMID 21536881. Bibcode: 2011PNAS..108.8379A. 
  10. ↑ 10.0 10.1 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. ↑ "GRIN Genera of Aizoaceae subfam. Acrosanthoideae". Germplasm Resources Information Network. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomygenuslist?id=3270&type=subfamily. 
  12. ↑ "GRIN Genera of Aizoaceae subfam. Aizooideae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2448. 
  13. ↑ "GRIN Genera of Aizoaceae subfam. Mesembryanthemoideae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2451. 
  14. ↑ "GRIN Genera of Aizoaceae tribe Apatesieae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2452. 
  15. ↑ "GRIN Genera of Aizoaceae tribe Dorotheantheae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2453. 
  16. ↑ "GRIN Genera of Aizoaceae tribe Ruschiae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2454. 
  17. ↑ 17.0 17.1 Klak, C., Hanáček, P. and Bruyns, P.V. (2024), Phylogeny and reclassification of Lampranthus (Ruschieae, Aizoaceae) in southern Africa. Taxon, 73: 818-853. https://doi.org/10.1002/tax.13166
  18. ↑ "Mesembryanthemaceae - Ruschioideae - Ruschia Group Marlothistella". http://biodiversityadvisor.sanbi.org/wp-content/themes/bst/keys/e-Key-20160604/Genera/G_Marlothistella.html. 
  19. ↑ "GRIN Genera of Aizoaceae subfam. Sesuvioideae". Germplasm Resources Information Network. http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2449. 
  20. ↑ Klak, C., Hanáček2, P. and Bruyns, P.V. (2017), Disentangling the Aizooideae: New generic concepts and a new subfamily in Aizoaceae. Taxon, 66: 1147-1170. https://doi.org/10.12705/665.9
  21. ↑ 21.0 21.1 21.2 Facciola. S. (1990). Cornucopia - A Source Book of Edible Plants. Kampong Publications. ISBN 0-9628087-0-9. 
  22. ↑ Low. T. (1989). Wild Food Plants of Australia. Angus and Robertson. ISBN 0-207-14383-8. 
  23. ↑ "Invasive Plants of California's Wildland". California Invasive Plant Council. http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=25&surveynumber=182.php. 
  24. ↑ "Fire Safe Landscaping". Cal Fire. http://www.readyforwildfire.org/Fire-Safe-Landscaping. 
  25. ↑ Baldwin, Debra Lee. "Firewise Landscaping with Succulents - How succulents saved a Rancho Santa Fe home from wildfire". http://debraleebaldwin.com/firewise-landscaping-with-succulents. 

Further reading

  • Volker Bittrich; Heidrun E. K. Hartman (1988). "The Aizoaceae—a new approach". Bot. J. Linn. Soc. 97 (3): 239–254. doi:10.1111/j.1095-8339.1988.tb01581.x. 
  • Cornelia Klak; Angeline Khunou; Gail Reeves; Terry Hedderson (2003). "A phylogenetic hypothesis for the Aizoaceae (Caryophyllales) based on four plastid DNA regions". Am. J. Bot. 90 (10): 1433–1445. doi:10.3732/ajb.90.10.1433. PMID 21659095. 

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