Caryophyllales ( ) is a diverse and heterogeneous order of

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s that includes the cacti,

carnation ''Dianthus caryophyllus'' (), commonly known as the carnation or clove pink, is a species of ''Dianthus''. It is likely native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years.Med ...

s, amaranths, ice plants,

beet The beetroot is the taproot portion of a beet plant, usually known in North America as beets while the vegetable is referred to as beetroot in British English, and also known as the table beet, garden beet, red beet, dinner beet or golden beet ...

s, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves. The

betalain Betalains are a class of red and yellow tyrosine-derived pigments found in plants of the order Caryophyllales, where they replace anthocyanin pigments. Betalains also occur in some higher order fungi. They are most often noticeable in the petals ...

pigments are unique in plants of this order and occur in all its families with the exception of Caryophyllaceae and

Molluginaceae The Molluginaceae are a family of flowering plants recognized by several taxonomists. It was previously included in the larger family Aizoaceae. The APG III system of 2009 made no change in the status of the family as compared to the APG II syst ...

Description

The members of Caryophyllales include about 6% of eudicot

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

. This order is part of the core eudicots. Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species The monophyly of the Caryophyllales has been supported by

DNA sequences A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are us ...

, cytochrome c sequence data and heritable characters such as anther wall development and vessel-elements with simple perforations.

Circumscription

As with all

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

, the circumscription of Caryophyllales has changed within various classification systems. All systems recognize a core of families with centrospermous ovules and seeds. More recent treatments have expanded the Caryophyllales to include many carnivorous plants. Systematists were undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid clade. The possible connection between sympetalous angiosperms and Caryophyllales was presaged by Bessey, Hutchinson, and others; as Lawrence relates: "The evidence is reasonably conclusive that the Primulaceae and the Caryophyllaceae have fundamentally the same type of gynecia, and as concluded by Douglas (1936)(and essentially Dickson, 1936) '...the vascular pattern and the presence of locules at the base of the ovary point to the fact that the present much reduced flower of the Primulaceae has descended from an ancestor which was characterized by a plurilocular ovary and axial placentation. This primitive flower might well be found in centrospermal stock as Wernham, Bessy, and Hutchinson have suggested.' " Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae. These two suborders were formerly (and sometimes still are) recognized as two orders,

Polygonales Polygonales was an order of flowering plants recognized by several older systems such as the Wettstein system, last revised in 1935, the Engler system, in its update of 1964, and the Cronquist system, 1981. Its circumscription was typically: * o ...

and Caryophyllales. Wigginsia Pauciareolata

APG IV

Kewaceae ''Kewa'' is a genus of flowering plants, consisting of eight species of succulent sub-woody plants, native to eastern and southern Africa, including Saint Helena and Madagascar. These are small shrubs or herbs that form cushions and have edible, ...

, Macarthuriaceae, Microteaceae, and Petiveriaceae were added in APG IV.

APG III

As circumscribed by the APG III system (2009), this order includes the same families as the APG II system (see below) plus the new families, Limeaceae, Lophiocarpaceae, Montiaceae, Talinaceae, and Anacampserotaceae. * family Achatocarpaceae * family Aizoaceae * family Amaranthaceae * family

Anacampserotaceae The Anacampserotaceae are a family of plants proposed in the February 2010 issue of the journal ''Taxon''. The family was described by Urs Eggli and Reto Nyffeler in their analysis of the polyphyly in the suborder Portulacineae (order Caryophyl ...

* family

Ancistrocladaceae ''Ancistrocladus'' is a genus of woody lianas in the monotypic family ''Ancistrocladaceae''. The branches climb by twining other stems or by scrambling with hooked tips. They are found in the tropics of the Old World. Classification The APG II ...

* family Asteropeiaceae * family Barbeuiaceae * family

Basellaceae Basellaceae is a family of flowering plants in the order Caryophyllales, in the clade core eudicots, according to the Angiosperm Phylogeny Group. The family comprises 19 known species of herbaceous plants in four genera: * '' Anredera'' – 12 s ...

* family

Cactaceae A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Gree ...

* family Caryophyllaceae * family Didiereaceae * family

Dioncophyllaceae The Dioncophyllaceae are a family (biology), family of flowering plants consisting of three species of lianas native to the rainforests of western Africa. Their closest relatives are Ancistrocladaceae. Both families lie within a clade of mostly ...

* family Droseraceae * family Drosophyllaceae * family Frankeniaceae * family

Gisekiaceae ''Gisekia'' is a genus of flowering plants. It is the only genus in the family Gisekiaceae. The family was recognized in the APG II system (2003) and assigned to the order Caryophyllales in the clade core eudicots. This represents a change from t ...

* family

Halophytaceae ''Halophytum ameghinoi'' is a species of herbaceous plant endemic to Patagonia. It is the only species in the genus ''Halophytum''. It is a succulent annual plant, with simple, fleshy, alternate leaves. The plants are monoecious, with solit ...

* family

* family Limeaceae * family Lophiocarpaceae * family Macarthuriaceae * family Microteaceae * family

* family

Montiaceae Montiaceae are a family of flowering plants, comprising about 14 genera with about 230 known species, ranging from small herbaceous plants to shrubs. The family has a cosmopolitan distribution. The family Montiaceae was newly adopted in the APG ...

* family Nepenthaceae * family

Nyctaginaceae Nyctaginaceae, the four o'clock family, is a family of around 33 genera and 290 species of flowering plants, widely distributed in tropical and subtropical regions, with a few representatives in temperate regions. The family has a unique fruit t ...

* family Petiveriaceae * family Physenaceae * family Phytolaccaceae * family Plumbaginaceae * family Polygonaceae * family

Portulacaceae The Portulacaceae are a family of flowering plants, comprising 115 species in a single genus '' Portulaca''. Formerly some 20 genera with about 500 species, were placed there, but it is now restricted to encompass only one genus, the other genera ...

* family

Rhabdodendraceae ''Rhabdodendron'' is a genus comprising two or three species of tropical South American trees. ''Rhabdodendron'' is placed in its own family Family (from la, familia) is a group of people related either by consanguinity (by recognized bir ...

* family Sarcobataceae * family

Simmondsiaceae Simmondsiaceae or the jojoba family is a family of flowering plants. The family is not recognized by all taxonomic systems, the single species, ''Simmondsia chinensis'', often being treated as belonging to family Buxaceae. The APG II system, of ...

* family Stegnospermataceae * family Talinaceae * family

Tamaricaceae The Tamaricaceae, the tamarisk family, are a family of plants native to drier areas of Europe, Asia, and Africa. It contains four genera: '' Tamarix'' (with 73 species), ''Reaumuria'' (25 species), ''Myricaria'' (13 species), and '' Myrtama'' (a ...

APG II

As circumscribed by 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-based, system of plant taxonomy that was published in April 2003 by the Angiosperm Phylogeny Gr ...

(2003), this order includes well-known plants like cacti,

s, spinach,

, rhubarb,

sundew ''Drosera'', which is commonly known as the sundews, is one of the largest genera of carnivorous plants, with at least 194 species. 2 volumes. These members of the family Droseraceae lure, capture, and digest insects using stalked mucilaginou ...

venus fly trap The Venus flytrap (''Dionaea muscipula'') is a carnivorous plant native to subtropical wetlands on the East Coast of the United States in North Carolina and South Carolina. It catches its prey—chiefly insects and arachnids—with a trapping ...

s, and bougainvillea. Recent molecular and biochemical evidence has resolved additional well-supported clades within the Caryophyllales. * order Caryophyllales ** family Achatocarpaceae ** family Aizoaceae ** family Amaranthaceae Pupalia lappacea (Forest Burr) W IMG 1562

Pupalia lappacea (Forest Burr) W IMG 1562

** family

(added in APG III) ** family

** family Asteropeiaceae ** family Barbeuiaceae ** family

** family

** family Caryophyllaceae ** family Didiereaceae ** family

** family Droseraceae ** family Drosophyllaceae ** family Frankeniaceae ** family

** family

** family Limeaceae (added in APG III) ** family Lophiocarpaceae (added in APG III) ** family

Glinus oppositifolius (Bitter cumin) W2 IMG 0462

** family

(added in APG III) ** family Nepenthaceae ** family

** family Physenaceae ** family Phytolaccaceae ** family Plumbaginaceae ** family Polygonaceae ** family

** family

** family Sarcobataceae ** family

** family Stegnospermataceae ** family Talinaceae (added in APG III) ** family

APG

This represents a slight change 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 A ...

, of 1998 * order Caryophyllales *: family Achatocarpaceae *: family Aizoaceae *: family Amaranthaceae *: family Ancistrocladaceae *: family Asteropeiaceae *: family Basellaceae *: family Cactaceae *: family Caryophyllaceae *: family Didiereaceae *: family Dioncophyllaceae *: family Droseraceae *: family Drosophyllaceae *: family Frankeniaceae *: family Molluginaceae *: family Nepenthaceae *: family Nyctaginaceae *: family Physenaceae *: family Phytolaccaceae *: family Plumbaginaceae *: family Polygonaceae *: family Portulacaceae *: family Rhabdodendraceae *: family Sarcobataceae *: family Simmondsiaceae *: family Stegnospermataceae *: family Tamaricaceae

Cronquist

The

Cronquist system The Cronquist system is a taxonomic classification system of flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of Flowering Plants'' (1968; 2nd edition, 1988) ...

(1981) also recognised the order, with this circumscription: * order Caryophyllales *: family Achatocarpaceae *: family Aizoaceae *: family Amaranthaceae *: family Basellaceae *: family Cactaceae *: family Caryophyllaceae *: family Chenopodiaceae *: family Didiereaceae *: family Nyctaginaceae *: family Phytolaccaceae *: family Portulacaceae *: family Molluginaceae The difference with the order as recognized by APG lies in the first place in the concept of "order". The APG favours much larger orders and families, and the order Caryophyllales ''sensu'' APG should rather be compared to subclass Caryophyllidae ''sensu'' Cronquist. A part of the difference lies with what families are recognized. The plants in the Stegnospermataceae and Barbeuiaceae were included in Cronquist's Phytolaccaceae. The Chenopodiaceae (still recognized by Cronquist) are included in Amaranthaceae by APG. New to the order (''sensu ''APG) are the Asteropeiaceae and Physenaceae, each containing a single genus, and two genera from Cronquist's order Nepenthales.

Earlier circumscriptions

Earlier systems, such as the

Wettstein system A system of plant taxonomy, the Wettstein system recognised the following main groups, according to Richard Wettstein's ''Handbuch der Systematischen Botanik'' (1901–1924). 3rd edition (1924) Outline Synopsis * Flagellatae p. 65 * ...

, last edition in 1935, and the

Engler system One of the prime systems of plant taxonomy, the Engler system was devised by Adolf Engler (1844–1930), and is featured in two major taxonomic texts he authored or co-authored. His influence is reflected in the use of the terms "Engler School" and ...

, updated in 1964, had a similar order under the name

Centrospermae ''Centrospermae'' is a descriptive botanical name, published in 1878 by August W. Eichler, Eichler, meaning "with the seed in the center", referring to the Placentation#In_plants, free (central) placentation. It was used in the Engler system and ...

References

External links

Tree of Life
Characteristics and Phylogenetic Relationships {{Taxonbar, from=Q21808 Angiosperm orders