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The Aizoaceae (), or fig-marigold family, is a large
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, ...
ous
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s containing 135
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
and about 1,800
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. Several genera are commonly known as 'ice plants' or 'carpet weeds'. The Aizoaceae are also referred to as ''vygies'' in
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. Some of the unusual
Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...
n genera—such as ''
Conophytum ''Conophytum'' is a genus of South African and Namibian succulent plants that belong to the family Aizoaceae. The name is derived from the Latin ''conus'' (cone) and Greek ''phytum'' (plant). The plants are also known as knopies (buttons in A ...
'', ''
Lithops ''Lithops'' is a genus of succulent plants in the ice plant family, Aizoaceae. Members of the genus are native to southern Africa. They avoid being eaten by herbivores with their camouflage as small stones, and are often known as pebble plants ...
'', ''
Titanopsis ''Titanopsis'' is a genus of about 10 species of succulent plants of the family Aizoaceae, indigenous to the arid regions of South Africa and Namibia. The name ''Titanopsis'' is derived from the Greek (god), ''Titans, Titan'', the sun, and ''ops ...
'' and ''
Pleiospilos ''Pleiospilos'' is a genus of succulent flowering plants of the family Aizoaceae. It is endemic to the Cape Provinces of South Africa. The name is derived from the Greek ''pleios'' "many" and ''spìlos'' "spot". The plants are also known as ''kwa ...
'' (among others)—resemble gemstones, rocks or pebbles, and are sometimes referred to as 'living stones' or 'mesembs' (short for
mesembryanthemum ''Mesembryanthemum'' is a genus of flowering plants in the family Aizoaceae, indigenous to southern Africa. As with many members of that family, it is characterized by long-lasting flower heads. Flowers of ''Mesembryanthemum'' protect their game ...
s).


Description

The family Aizoaceae is widely recognised by taxonomists. It once went by the botanical name "Ficoidaceae", now disallowed. The
APG II system The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly Molecular phylogenetics, molecular-based, list of systems of plant taxonomy, system of plant taxonomy that ...
of 2003 (unchanged from the
APG system The APG system (Angiosperm Phylogeny Group system) of plant classification is the first version of a modern, mostly molecular-based, system of plant taxonomy. Published in 1998 by the Angiosperm Phylogeny Group, it was replaced by the improved ...
of 1998) also recognizes the family, and assigns it to the order
Caryophyllales Caryophyllales ( ) is a diverse and heterogeneous order of flowering plants with well-known members including cacti, carnations, beets, quinoa, spinach, amaranths, pigfaces and ice plants, oraches and saltbushes, goosefoots, sundews, Venu ...
in the clade
core eudicots The eudicots or eudicotyledons are flowering plants that have two seed leaves (cotyledons) upon germination. The term derives from ''dicotyledon'' (etymologically, ''eu'' = true; ''di'' = two; ''cotyledon'' = seed leaf). Historically, authors h ...
. 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. Glistening epidermal bladder cells give the family its common name "ice plants". Most fig-marigolds are
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of ...
, rarely somewhat woody, with
sympodial In botany, sympodial growth is a bifurcating branching pattern where one branch develops more strongly than the other, resulting in the stronger branches forming the primary shoot and the weaker branches appearing laterally. A sympodium, als ...
growth and stems either erect or prostrate.
Leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
are simple, opposite or alternate, and more or less succulent with entire (or rarely toothed) margins.
Flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s are perfect in most species (but unisexual in some), actinomorphic, and appear singularly or in few-flowered cymes developing from the leaf axils.
Sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s are typically five (3–8) and more or less connate (fused) below. True
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s are absent. However, some species have numerous linear petals derived from staminodes. The seed capsules have one to numerous seeds per cell and are often hygrochastic, dispersing seeds by "jet action" when wet.


Distribution

Most species (96%, 1782 species in 132 genera) in this family are endemic to arid or semiarid parts of Southern
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
in the
Succulent Karoo The Succulent Karoo is an terrestrial ecoregion, ecoregion defined by the World Wide Fund for Nature to include regions of deserts and xeric shrublands, desert in South Africa and Namibia, and a biodiversity hotspot. The geographic area chosen ...
. Much of the Aizoaceae's diversity is found in the Greater
Cape Floristic Region The Cape Floral Region is a floristic region located near the southern tip of South Africa. It is the only floristic region of the Cape Floristic Kingdom, and includes only one floristic province, known as the Cape Floristic Province. The Cap ...
, which is the most plant-diverse temperate region in the world. A few species are found in
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
and the Central Pacific area.


Evolution

The radiation of the Aizoaceae, specifically the subfamily Ruschioideae, was one of the most recent among the
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...
, 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. This diversification was roughly contemporaneous with major radiations in two other succulent lineages,
Cactaceae A cactus (: cacti, cactuses, or less commonly, cactus) is a member of the plant family Cactaceae (), a family of the order Caryophyllales comprising about 127 genera with some 1,750 known species. The word ''cactus'' derives, through Latin, ...
and ''
Agave ''Agave'' (; ; ) is a genus of monocots native to the arid regions of the Americas. The genus is primarily known for its succulent and xerophytic species that typically form large Rosette (botany), rosettes of strong, fleshy leaves. Many plan ...
.'' The family includes many species that use
crassulacean acid metabolism Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions that allows a plant to photosynthesize during the day, but only exchange gases at night. ...
as pathway for
carbon fixation Biological carbon fixation, or сarbon assimilation, is the Biological process, process by which living organisms convert Total inorganic carbon, inorganic carbon (particularly carbon dioxide, ) to Organic compound, organic compounds. These o ...
. Some species in the subfamily Sesuvioideae instead use carbon fixation, which might have evolved multiple times in the group.


Taxonomy

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


Subfamily Acrosanthoideae

Genera: * '' Acrosanthes'' Eckl. & Zeyh.


Subfamily Aizooideae

Genera: * '' Aizoanthemopsis'' Klak * '' Aizoanthemum'' Dinter ex Friedrich * ''
Aizoon ''Aizoon'' or ''Aizoön'' is a genus of flowering plants in the iceplant family, Aizoaceae. Description No raphides occur in at least the leaves and stalks of at least 3 species. Distribution Apart from '' A. canariense'', which is native to Ma ...
'' L. * '' Gunniopsis'' Pax * '' Tetragonia'' L.


Subfamily Mesembryanthemoideae

Genera:


Subfamily Ruschioideae

Genera: ;Tribe Ruschieae * ''
Acrodon ''Acrodon'' is a genus of Aizoaceae, ice plants from South Africa. It comprises five species, mostly endangered and all restricted to the southern Cape regions of the Western Cape and Eastern Cape Provinces, South Africa. Description Species of ...
'' 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 ''Carpobrotus'', commonly known as pigface, ice plant, sour fig, Hottentot fig, and clawberry is a genus of ground-creeping plants with succulent leaves and large daisy-like flowers. The name comes from the Ancient Greek ' "fruit" and ' "edible", ...
'' N.E.Br * '' × Carruanthophyllum'' (''Carruanthus'' × ''Machairophyllum'') * '' Carruanthus'' (Schwantes) Schwantes * '' Cephalophyllum'' N.E.Br * '' Cerochlamys'' N.E.Br * '' Chasmatophyllum'' Dinter & Schwantes * '' Cheiridopsis'' N.E.Br * '' Circandra'' N.E.Br * ''
Conophytum ''Conophytum'' is a genus of South African and Namibian succulent plants that belong to the family Aizoaceae. The name is derived from the Latin ''conus'' (cone) and Greek ''phytum'' (plant). The plants are also known as knopies (buttons in A ...
'' N.E.Br * ''Corpuscularia'' Schwantes, synonym of ''Delosperma'' * '' Cylindrophyllum'' Schwantes * ''
Delosperma ''Delosperma'' ('delos'=evident, 'sperma'=seed) is a genus of around 170 species of succulent plants, formerly included in ''Mesembryanthemum'' in the family Aizoaceae. It was defined by English botanist N. E. Brown in 1925. The genus is common ...
'' 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 * '' Faucaria'' Schwantes * ''
Fenestraria ''Fenestraria'' (known as babies' toes) is a (possibly monotypic) genus of succulent plants in the family Aizoaceae, native to the Namaqualand in Namibia. Description The only species currently recognised in this genus is ''Fenestraria rh ...
'' 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 ''Lithops'' is a genus of succulent plants in the ice plant family, Aizoaceae. Members of the genus are native to southern Africa. They avoid being eaten by herbivores with their camouflage as small stones, and are often known as pebble plants ...
'' N.E.Br * ''
Machairophyllum ''Machairophyllum'' is a genus of plants in the family Aizoaceae. Species , Plants of the World Online accepted the following species: *''Machairophyllum albidum'' Schwantes *''Machairophyllum bijlii'' (N.E.Br.) L.Bolus *''Machairophyllum brevif ...
'' Schwantes * '' Malephora'' N.E.Br * '' Malotigena'' Niederle * '' Marlothistella'' Schwantes * ''
Mestoklema ''Mestoklema'' is a genus of flowering plants belonging to the family Aizoaceae. Its native range is Southern Africa. Species Species: *''Mestoklema albanicum'' *''Mestoklema arboriforme'' *''Mestoklema copiosum'' *''Mestoklema elatum'' *' ...
'' N.E.Br. ex Glen * '' Meyerophytum'' Schwantes * '' Mitrophyllum'' Schwantes * '' Monilaria'' (Schwantes) Schwantes * '' Mossia'' N.E.Br * '' Muiria'' N.E.Br * '' Namaquanthus'' L.Bolus * ''
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
'' (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 ''Pleiospilos'' is a genus of succulent flowering plants of the family Aizoaceae. It is endemic to the Cape Provinces of South Africa. The name is derived from the Greek ''pleios'' "many" and ''spìlos'' "spot". The plants are also known as ''kwa ...
'' 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. * '' Ruschia'' Schwantes * ''Ruschianthemum'' Friedrich, synonym of ''Stoeberia'' * '' Ruschianthus'' L.Bolus * '' Sarcozona'' J.M.Black * '' Schlechteranthus'' Schwantes * '' Schwantesia'' Dinter * '' Scopelogena'' L.Bolus * '' Smicrostigma'' N.E.Br * '' Stayneria'' L.Bolus * '' Stoeberia'' Dinter & Schwantes * '' Stomatium'' Schwantes * '' Tanquana'' H.E.K.Hartmann & Liede * ''
Titanopsis ''Titanopsis'' is a genus of about 10 species of succulent plants of the family Aizoaceae, indigenous to the arid regions of South Africa and Namibia. The name ''Titanopsis'' is derived from the Greek (god), ''Titans, Titan'', the sun, and ''ops ...
'' 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


Subfamily Sesuvioideae

This subfamily includes a number of species. Genera: * ''Anisostigma'' Schinz, synonym of '' Tetragonia'' * ''
Sesuvium ''Sesuvium'' is a genus of flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'se ...
'' L. * ''
Trianthema ''Trianthema'' is a genus of flowering plants in the ice plant family, Aizoaceae. Members of the genus are annuals or perennials generally characterized by fleshy, opposite, unequal, smooth-margined leaves, a prostrate growth form, flowers with ...
'' L. * '' Tribulocarpus'' S.Moore * '' Zaleya'' Burm.f.


Unplaced genera

Include; * '' Hammeria'' * '' Peersia''


Uses

Several genera are cultivated. ''
Lithops ''Lithops'' is a genus of succulent plants in the ice plant family, Aizoaceae. Members of the genus are native to southern Africa. They avoid being eaten by herbivores with their camouflage as small stones, and are often known as pebble plants ...
'', or "living stones", are popular as novelty house plants because of their stone-like appearance. Some species are edible, including: * ''
Carpobrotus edulis ''Carpobrotus edulis'' is a creeping plant with succulent leaves in the genus '' Carpobrotus'', native to South Africa. Its common names include hottentot-fig, sour fig, ice plant, highway ice plant, or vygie. Description ''Carpobrotus eduli ...
'' (Hottentot fig, highway ice plant) has edible leaves and fruit. * ''
Mesembryanthemum crystallinum ''Mesembryanthemum crystallinum'' (the common, 'crystalline' or 'crystallinum' iceplant, or simply 'iceplant') is a species of Annual plant, annual/perennial, Succulent plant, succulent flowering plant in the ''Mesembryanthemum'' genus of the Aiz ...
'' has edible leaves. * ''
Tetragonia tetragonoides ''Tetragonia tetragonioides'', commonly called New Zealand spinach, warrigal greens and other local names, is a flowering plant in the fig-marigold family (Aizoaceae). It is often cultivated as a leafy vegetable. It is a widespread species, nat ...
'' ("New Zealand spinach") is grown as a garden plant in somewhat dry climates and used as an alternative to
spinach Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to Central Asia, Central and Western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common vegetable consumed eit ...
in upscale
salad A salad is a dish consisting of mixed ingredients, frequently vegetables. They are typically served chilled or at room temperature, though some can be served warm. Condiments called '' salad dressings'', which exist in a variety of flavors, a ...
s. '' C. edulis'' was introduced to California in the early 1900s to stabilize soil along railroad tracks and has become invasive. In southern California, ice plants are sometimes used as firewalls; however, they do burn if not carefully maintained.


References


Further reading

* *


External links


Aizoaceae
in L. Watson and M.J. Dallwitz (1992 onwards)
''The families of flowering plants''

NCBI Taxonomy Browser

P. Chesselet (2004 onwards). Interactive Mesembs2

Plants of southern Africa (2005 onwards). SANBI




Flowers in Israel

i
BoDD – Botanical Dermatology Database
{{Authority control Caryophyllales families Flora of South Africa Fynbos Succulent plants