Aizoaceae

{{Short description|Family of dicotyledonous flowering plants}} {{Automatic taxobox | image = Mesembryanthemum crystallinum.JPG | image_caption = ''Mesembryanthemum crystallinum'' | taxon = Aizoaceae | authority = Martinov | type_genus = ''Aizoon'' | type_genus_authority = L. | subdivision_ranks = Genera | subdivision = See text | synonyms = * Ficoidaceae * Mesembryanthemaceae * Tetragoniaceae Lindl. | synonyms_ref = <ref name = "POWO">{{cite POWO |id=30004741-2 |title=''Aizoaceae'' Martinov |access-date=2022-09-08}}</ref> }}

The '''Aizoaceae''' ({{IPAc-en|ˌ|eɪ|z|oʊ|'|eɪ|s|i|.|iː|,_|-|ˌ|aɪ}}), or '''fig-marigold family''', are a large family of dicotyledonous flowering plants containing 135 genera and about 1,800 species.<ref>{{cite book |title=Illustrated Handbook of Succulent Plants |last=Hartmann|first=HEK |publisher=Springer |year=2001 |location=Berlin, Germany}}</ref> 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 ==

[[File:Mesembryanthemum guerichianum seedling IMG 8167-cropped.jpg|thumb|''Mesembryanthemum guerichianum'' seedling, showing the epidermal bladder cells that inspired the name "ice plant"]] thumb|right| Pronunciation of the South African colloquial name, '''Vygie''', for Aizoaceae 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.<ref name="SSP">{{cite web|url=http://succulent-plant.com/families/aizoaceae.html|title=The Living Stone Page|publisher=The Succulent Plant Page|access-date=23 June 2017}}</ref> Glistening epidermal bladder cells give the family its common name "ice plants".<ref name=":0">{{Cite web|url=http://www.mobot.org/MOBOT/research/APweb/|title=Angiosperm Phylogeny Website|website=www.mobot.org|access-date=2017-12-20}}</ref>

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.<ref>{{cite web |author1=Watson, L. |author2=Dallwitz, M.J. |date=1992 |title=The families of flowering plants: descriptions, illustrations, identification, and information retrieval |version=20 June 2017 |url=http://delta-intkey.com/angio/www/aizoacea.htm |access-date=9 July 2017}}</ref> The seed capsules have one to numerous seeds per cell and are often hygrochastic, dispersing seeds by "jet action" when wet.<ref name=":0" />

== 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.<ref name="Mesembryanthemaceae">{{cite journal |author1=Chesselet, P. |author2=Smith, G.F. |author3=Burgoyne, P.M. |author4=Klak, C. |author5=Hammer, S.A. |author6=Hartmann, H.E.K. |author7=Kurzweil, H. |author8=van Jaarsveld, E.J. |author9=van Wyk, B.-E. |author10=Leistner, O.A. |display-authors=3| title=Seed Plants of Southern Africa | journal=Strelitzia | year=2000| volume=10| pages=360–410}}</ref> 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.<ref>{{Cite journal|last1=Born|first1=J.|last2=Linder|first2=H. P.|last3=Desmet|first3=P.|date=2007|title=The Greater Cape Floristic Region|jstor=4125143|journal=Journal of Biogeography|volume=34|issue=1|pages=147–162|doi=10.1111/j.1365-2699.2006.01595.x}}</ref> A few species are found in Australia and the Central Pacific area.<ref name="WOS">{{cite web |title=Browsing: Aizoaceae |url=http://worldofsucculents.com/?families=aizoaceae |publisher=World of Succulents |access-date=23 June 2017}}</ref>

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 million years ago. 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.<ref name=":1">{{Cite journal|last1=Valente|first1=Luis M.|last2=Britton|first2=Adam W.|last3=Powell|first3=Martyn P.|last4=Papadopulos|first4=Alexander S. T.|last5=Burgoyne|first5=Priscilla M.|last6=Savolainen|first6=Vincent|date=2014-01-01|title=Correlates of hyperdiversity in southern African ice plants (Aizoaceae)|url= |journal=Botanical Journal of the Linnean Society|volume=174|issue=1|pages=110–129|doi=10.1111/boj.12117|pmid=25821244|issn=0024-4074|pmc=4373134}}</ref> This diversification was roughly contemporaneous with major radiations in two other succulent lineages, Cactaceae and ''Agave.''<ref>{{Cite journal|last1=Arakaki|first1=Mónica|last2=Christin|first2=Pascal-Antoine|last3=Nyffeler|first3=Reto|last4=Lendel|first4=Anita|last5=Eggli|first5=Urs|last6=Ogburn|first6=R. Matthew|last7=Spriggs|first7=Elizabeth|last8=Moore|first8=Michael J.|last9=Edwards|first9=Erika J.|date=2011|title=Contemporaneous and recent radiations of the world's major succulent plant lineages|jstor=25830059|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=108|issue=20|pages=8379–8384|doi=10.1073/pnas.1100628108|pmid=21536881|pmc=3100969|bibcode=2011PNAS..108.8379A|doi-access=free}}</ref>

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.<ref name="BohleyJoos2015">{{cite journal |last1=Bohley |first1=Katharina |last2=Joos |first2=Olga |last3=Hartmann |first3=Heidrun |last4=Sage |first4=Rowan |last5=Liede-Schumann |first5=Sigrid |last6=Kadereit |first6=Gudrun |title=Phylogeny of Sesuvioideae (Aizoaceae) – Biogeography, leaf anatomy and the evolution of {{C4}} photosynthesis |journal=Perspectives in Plant Ecology, Evolution and Systematics |volume=17 |issue=2 |year=2015 |pages=116–130 |issn=1433-8319 |doi=10.1016/j.ppees.2014.12.003}}</ref>

===Subfamily Acrosanthoideae===

Genera:<ref>{{cite web|url=https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomygenuslist?id=3270&type=subfamily |title=GRIN Genera of ''Aizoaceae'' subfam. ''Acrosanthoideae'' |work=Germplasm Resources Information Network |access-date=2022-11-10 }}</ref> * ''Acrosanthes'' Eckl. & Zeyh.

===Subfamily Aizooideae===

Genera:<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2448 |archive-url=https://archive.today/20120719010400/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2448 |url-status=dead |archive-date=2012-07-19 |title=GRIN Genera of ''Aizoaceae'' subfam. ''Aizooideae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 }}</ref> * ''Aizoanthemopsis'' Klak * ''Aizoanthemum'' Dinter ex Friedrich * ''Aizoon'' L. * ''Gunniopsis'' Pax * ''Tetragonia'' L.

===Subfamily Mesembryanthemoideae===

Genera:<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2451 |archive-url=https://archive.today/20120718080426/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2451 |url-status=dead |archive-date=2012-07-18 |title=GRIN Genera of ''Aizoaceae'' subfam. ''Mesembryanthemoideae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 }}</ref> {{div col|colwidth=28em}} * ''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'' {{div col end}}

===Subfamily Ruschioideae===

Genera: {| |- valign=top | ;Tribe Apatesieae<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2452 |title=GRIN Genera of ''Aizoaceae'' tribe ''Apatesieae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 |url-status=dead |archive-url=https://web.archive.org/web/20150924052721/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2452 |archive-date=2015-09-24 }}</ref> * ''Apatesia'' N.E.Br * ''Carpanthea'' N.E.Br * ''Caryotophora'' Leistner, synonym of ''Skiatophytum'' * ''Conicosia'' N.E.Br * ''Hymenogyne'' Haw. * ''Saphesia'' N.E.Br * ''Skiatophytum'' L.Bolus | ;Tribe Dorotheantheae<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2453 |archive-url=https://archive.today/20120724033836/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2453 |url-status=dead |archive-date=2012-07-24 |title=GRIN Genera of ''Aizoaceae'' tribe ''Dorotheantheae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 }}</ref> * ''Aethephyllum'' N.E.Br, synonym of ''Cleretum'' * ''Cleretum'' N.E.Br * ''Dorotheanthus'' Schwantes, synonym of ''Cleretum''

|} ;Tribe Ruschieae<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2454 |archive-url=https://archive.today/20120720223417/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2454 |url-status=dead |archive-date=2012-07-20 |title=GRIN Genera of ''Aizoaceae'' tribe ''Ruschiae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 }}</ref>

{{div col|colwidth=18em}} * ''Acrodon'' N.E.Br * ''Aloinopsis'' Schwantes * ''Amphibolia'' L.Bolus ex A.G.J.Herre * ''Antegibbaeum'' Schwantes ex C.Weber * ''Antimima'' N.E.Br * ''Arenifera'' Herre, synonym of ''Mesembryanthemum'' * ''Argyroderma'' N.E.Br * ''Astridia'' Dinter * ''Bergeranthus'' Schwantes * ''Bijlia'' N.E.Br, synonym of ''Pleiospilos'' * ''Braunsia'' Schwantes * ''Brianhuntleya'' Chess. et al. * ''Carpobrotus'' N.E.Br * ''× Carruanthophyllum'' {{small|G.D.Rowley}} (''Carruanthus'' × ''Machairophyllum'') * ''Carruanthus'' (Schwantes) Schwantes * ''Cephalophyllum'' N.E.Br * ''Cerochlamys'' N.E.Br * ''Chasmatophyllum'' Dinter & Schwantes * ''Cheiridopsis'' N.E.Br * ''Circandra'' N.E.Br, synonym of ''Erepsia'' * ''Conophytum'' N.E.Br * ''Corpuscularia'' Schwantes, synonym of ''Delosperma'' * ''Cylindrophyllum'' Schwantes * ''Delosperma'' N.E.Br * ''Dicrocaulon'' N.E.Br * ''Didymaotus'' N.E.Br * ''Dinteranthus'' Schwantes * ''Diplosoma'' Schwantes * ''Disphyma'' N.E.Br * ''Dracophilus'' (Schwantes) Dinter & Schwantes * ''Drosanthemum'' Schwantes * ''Eberlanzia'' Schwantes * ''Ebracteola'' Dinter & Schwantes * ''Ectotropis'' N.E.Br, synonym of ''Delosperma'' * ''Enarganthe'' N.E.Br * ''Erepsia'' N.E.Br * ''Esterhuysenia'' L.Bolus, synonym of ''Lampranthus'' * ''Faucaria'' Schwantes * ''Fenestraria'' N.E.Br * ''Frithia'' N.E.Br * ''Gibbaeum'' Haw. ex N.E.Br * ''Glottiphyllum'' Haw. ex N.E.Br * ''Hallianthus'' H.E.K.Hartmann * ''Hereroa'' (Schwantes) Dinter & Schwantes * ''Ihlenfeldtia'' H.E.K.Hartmann, synonym of ''Cheiridopsis'' * ''Imitaria'' N.E.Br, synonym of ''Gibbaeum'' * ''Jacobsenia'' L.Bolus & Schwantes * ''Jensenobotrya'' A.G.J.Herre * ''Jordaaniella'' H.E.K.Hartmann * ''Juttadinteria'' Schwantes * ''Khadia'' N.E.Br * ''Lampranthus'' N.E.Br * ''Lapidaria'' (Dinter & Schwantes) N.E.Br. * ''Leipoldtia'' L.Bolus * ''Lemonanthemum'' Klak * ''Lithops'' N.E.Br * ''Machairophyllum'' Schwantes * ''Malephora'' N.E.Br * ''Malephoropsis'' {{small|Klak}}<ref name = Klak>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</ref> * ''Malotigena'' Niederle * ''Marlothistella'' Schwantes<ref>{{cite web |title=Mesembryanthemaceae - Ruschioideae - Ruschia Group Marlothistella |url=http://biodiversityadvisor.sanbi.org/wp-content/themes/bst/keys/e-Key-20160604/Genera/G_Marlothistella.html |website=biodiversityadvisor.sanbi.org |access-date=22 October 2021 |archive-date=22 October 2021 |archive-url=https://web.archive.org/web/20211022235209/http://biodiversityadvisor.sanbi.org/wp-content/themes/bst/keys/e-Key-20160604/Genera/G_Marlothistella.html |url-status=dead }}</ref> * ''Mestoklema'' N.E.Br. ex Glen * ''Meyerophytum'' Schwantes * ''Mitrophyllum'' Schwantes * ''Monilaria'' (Schwantes) Schwantes * ''Mossia'' N.E.Br * ''Muiria'' N.E.Br * ''Namaquanthus'' L.Bolus * ''Namibia'' (Schwantes) Schwantes * ''Nananthus'' N.E.Br * ''Nelia'' Schwantes * ''Neohenricia'' L.Bolus * ''Octopoma'' N.E.Br * ''Odontophorus'' N.E.Br, synonym of ''Cheiridopsis'' * ''Oophytum'' N.E.Br * ''Orthopterum'' L.Bolus * ''Oscularia'' Schwantes * ''Ottosonderia'' L.Bolus * ''Phiambolia'' Klak * ''Pleiospilos'' N.E.Br * ''Polymita'' N.E.Br, synonym of ''Schlechteranthus'' * ''Psammophora'' Dinter & Schwantes * ''Rabiea'' N.E.Br * ''Rhinephyllum'' N.E.Br * ''Rhombophyllum'' (Schwantes) Schwantes * ''Roosia'' van Jaarsv., synonym of ''Lampranthus'' * ''Ruschia'' Schwantes * ''Ruschianthemum'' Friedrich, synonym of ''Stoeberia'' * ''Ruschianthus'' L.Bolus * ''Sarcozona'' J.M.Black * ''Schlechteranthus'' Schwantes * ''Schwantesia'' Dinter * ''Scopelogena'' L.Bolus * ''Sederbergia'' {{small|Klak}}<ref name = Klak/> * ''Smicrostigma'' N.E.Br * ''Stayneria'' L.Bolus * ''Stoeberia'' Dinter & Schwantes * ''Stomatium'' Schwantes * ''Tanquana'' H.E.K.Hartmann & Liede * ''Titanopsis'' Schwantes * ''Trichodiadema'' Schwantes * ''Vanheerdea'' L.Bolus ex H.E.K.Hartmann * ''Vanzijlia'' L.Bolus * ''Vlokia'' S.A.Hammer * ''Wooleya'' L.Bolus * ''Zeuktophyllum'' N.E.Br {{div col end}}

===Subfamily Sesuvioideae=== This subfamily includes a number of {{C4}} species.<ref name="BohleyJoos2015"/>

Genera:<ref>{{cite web|url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2449 |archive-url=https://archive.today/20120718193929/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2449 |url-status=dead |archive-date=2012-07-18 |title=GRIN Genera of ''Aizoaceae'' subfam. ''Sesuvioideae'' |work=Germplasm Resources Information Network |access-date=2011-03-04 }}</ref> ;Tribe Anisostigmateae<ref>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</ref> * ''Anisostigma'' Schinz * ''Tribulocarpus'' S.Moore

;Tribe Sesuvieae * ''Sesuvium'' L. * ''Trianthema'' L. * ''Zaleya'' Burm.f.

===Unplaced genera=== Include; * ''Hammeria'' {{small|Burgoyne}} * ''Peersia'' {{small|L.Bolus}}

==Uses==

thumb|right|''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: * ''Carpobrotus edulis'' (Hottentot fig, highway ice plant) has edible leaves and fruit.<ref name=PFAF183>{{cite book |author=Facciola. S. |title=Cornucopia - A Source Book of Edible Plants |publisher=Kampong Publications |date=1990 |isbn=0-9628087-0-9}}</ref> * ''Mesembryanthemum crystallinum'' has edible leaves.<ref name=PFAF183/> * ''Tetragonia tetragonoides'' ("New Zealand spinach") is grown as a garden plant in somewhat dry climates and used as an alternative to spinach in upscale salads.<ref name=PFAF183/><ref name=PFAF194>{{cite book |author=Low. T. |title=Wild Food Plants of Australia |publisher=Angus and Robertson |date=1989|isbn=0-207-14383-8}}</ref>

''C. edulis'' was introduced to California in the early 1900s to stabilize soil along railroad tracks and has become invasive.<ref name=CAL-IPC>{{cite web |title= Invasive Plants of California's Wildland |url=http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=25&surveynumber=182.php |publisher=California Invasive Plant Council |access-date=23 June 2017}}</ref> In southern California, ice plants are sometimes used as firewalls;<ref>{{cite web |title=Fire Safe Landscaping |url=http://www.readyforwildfire.org/Fire-Safe-Landscaping |publisher=Cal Fire |access-date=23 June 2017 |archive-url=https://web.archive.org/web/20170702113714/http://readyforwildfire.org/Fire-Safe-Landscaping/ |archive-date=2 July 2017 |url-status=dead }}</ref> however, they do burn if not carefully maintained.<ref>{{cite web |last=Baldwin |first=Debra Lee |title=Firewise Landscaping with Succulents - How succulents saved a Rancho Santa Fe home from wildfire |url=http://debraleebaldwin.com/firewise-landscaping-with-succulents |access-date=23 June 2017}}</ref>{{Clear}}

==References== {{Reflist}}

==Further reading== * {{Cite journal|author1=Volker Bittrich |author2-link=H.E.K.Hartmann |author2=Heidrun E. K. Hartman |title= The Aizoaceae—a new approach |journal= Bot. J. Linn. Soc. |volume=97 |pages=239–254 |year=1988 |doi= 10.1111/j.1095-8339.1988.tb01581.x|issue=3}} * {{Cite journal|author1=Cornelia Klak |author2=Angeline Khunou |author3=Gail Reeves |author4=Terry Hedderson | title = A phylogenetic hypothesis for the Aizoaceae (Caryophyllales) based on four plastid DNA regions | journal = Am. J. Bot. | year = 2003| volume=90| pages=1433–1445 | doi = 10.3732/ajb.90.10.1433|pmid=21659095 | issue = 10 | doi-access=free }}

==External links== {{Commons category}} * [https://www.delta-intkey.com/angio/www/aizoacea.htm Aizoaceae] in L. Watson and M.J. Dallwitz (1992 onwards). [https://web.archive.org/web/20070103200438/http://delta-intkey.com/angio/ ''The families of flowering plants''] * [https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=3542&lvl=3&p=mapview&p=has_linkout&p=blast_url&p=genome_blast&lin=f&keep=1&srchmode=1&unlock NCBI Taxonomy Browser] * [https://web.archive.org/web/20070221160857/http://www.flora.sanbi.org/its_page?comID=3 P. Chesselet (2004 onwards). Interactive Mesembs2] * [https://web.archive.org/web/20110727234332/http://posa.sanbi.org/flora/results.php?taxon=famno=162&showdetail=F Plants of southern Africa (2005 onwards). SANBI] * [https://web.archive.org/web/20070926233924/http://www.museums.org.za/bio/plants/aizoaceae/index.htm Aizoaceae of South Africa] * [http://www.flowersinisrael.com/FamAizoaceae.htm Family Aizoaceae] Flowers in Israel * [https://www.botanical-dermatology-database.info/BotDermFolder/AIZO.html Aizoaceae] in [https://www.botanical-dermatology-database.info/ BoDD – Botanical Dermatology Database]

Category:Aizoaceae Category:Caryophyllales families Category:Flora of South Africa Category:Fynbos Category:Succulent plants