Caryophyllales ( ) is a diverse and heterogeneous

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number 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 t ...

s that includes the

cacti 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 ...

, carnations,

amaranth ''Amaranthus'' is a cosmopolitan genus of annual or short-lived perennial plants collectively known as amaranths. Some amaranth species are cultivated as leaf vegetables, pseudocereals, and ornamental plants. Catkin-like cymes of densely p ...

s, ice plants, beets, and many

carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants still generate some of their energy from photosynthesis ...

s. Many members are

succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...

, having fleshy stems or

leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...

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

Caryophyllaceae Caryophyllaceae, commonly called the pink family or carnation family, is a family of flowering plants. It is included in the dicotyledon order Caryophyllales in the APG III system, alongside 33 other families, including Amaranthaceae, Cactacea ...

and Molluginaceae.

Description

The members of Caryophyllales include about 6% of

eudicot The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...

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 appropriat ...

. This order is part of the

core eudicot The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...

s. 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

s. Systematists were undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

. 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

locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...

s 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,

Macarthuriaceae Macarthuriaceae is a family of plants in the order Caryophyllales and consists of a single genus, '' Macarthuria''. Description Macarthuriaceae are rigid or wiry, rush-like herbs or subshrubs with green stems and reduced leaves. The small flower ...

, Microteaceae, and

Petiveriaceae Petiveriaceae is a family of flowering plants formerly included as subfamily Rivinoideae in Phytolaccaceae. The family comprises nine genera, with about 20 known species In biology, a species is the basic unit of Taxonomy (biology), classi ...

were added in APG IV.

APG III

As circumscribed by the

APG III system The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a fu ...

(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 The Aizoaceae, or fig-marigold family, is a large family of dicotyledonous flowering plants containing 135 genera and about 1800 species. They are commonly known as ice plants or carpet weeds. They are often called vygies in South Africa and N ...

* family

Amaranthaceae Amaranthaceae is a family of flowering plants commonly known as the amaranth family, in reference to its type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making i ...

* family Anacampserotaceae * family Ancistrocladaceae * family

Asteropeiaceae :''"Asteropeia" may also refer to a figure in Greek mythology, see Antinoe'' ''Asteropeia'' is a genus of flowering plants. The genus contains 8 known species of shrubs and small trees, all endemic to Madagascar. It is the sole genus in fami ...

* family

Barbeuiaceae ''Barbeuia madagascariensis'' is a liana found only on the island of Madagascar. ''Barbeuia'' has occasionally been placed in its own family, Barbeuiaceae. The APG II system of 2003, for instance, recognizes such a family and assigns it to the ...

* family Basellaceae * 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 Gre ...

* family

* family Didiereaceae * family Dioncophyllaceae * family

Droseraceae Droseraceae is a family of carnivorous flowering plants, also known as the sundew family. It consists of approximately 180 species in three extant genera. Representatives of the Droseraceae are found on all continents except Antarctica. Desc ...

* family

Drosophyllaceae ''Drosophyllum'' ( , rarely ) is a genus of carnivorous plants containing the single species ''Drosophyllum lusitanicum'', commonly known as Portuguese sundew or dewy pine. In appearance, it is similar to the related genus '' Drosera'' (the sun ...

* 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 Kewaceae * family

Limeaceae ''Limeum'' is a genus of flowering plants. It includes 25 species. ''Limeum'' has traditionally been recognized as belonging to the Molluginaceae family, but is now treated as the sole genus in the family Limeaceae. The family is newly recognize ...

* family

Lophiocarpaceae The Lophiocarpaceae are a family of flowering plants comprising mostly succulent subshrubs and herbaceous species native to tropical to southern sub-Saharan Africa to western India. It includes the genera '' Corbichonia'' and '' Lophiocarpus''.An ...

* family

* family Microteaceae * family Molluginaceae * family Montiaceae * family Nepenthaceae * family Nyctaginaceae * family

* family

Physenaceae ''Physena'' is the sole genus of the flowering plant family Physenaceae. It contains two species of shrubs and small trees which are endemic to Madagascar. The APG II system, of 2003 (unchanged from the APG system, of 1998), does recognize this ...

* family

Phytolaccaceae Phytolaccaceae is a family of flowering plants. Though almost universally recognized by taxonomists, its circumscription has varied. It is also known as the Pokeweed family. The APG II system, of 2003 (unchanged from the APG system, of 1998), ...

* family

Plumbaginaceae Plumbaginaceae is a family of flowering plants, with a cosmopolitan distribution. The family is sometimes referred to as the leadwort family or the plumbago family. Most species in this family are perennial herbaceous plants, but a few grow ...

* family

Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus ''Polygonum'', and was first used by Antoine Laurent de Jussieu in ...

* family Portulacaceae * family Rhabdodendraceae * family Sarcobataceae * family Simmondsiaceae * family

Stegnospermataceae ''Stegnosperma'' is a genus of flowering plants, consisting of three species of woody plants, native to the Caribbean, Central America, and the Sonoran Desert. These are shrubs or lianas, with anomalous secondary thickening in mature stems, by su ...

* 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

, carnations,

spinach Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to central and western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common edible vegetable consumed either f ...

, beet,

rhubarb Rhubarb is the fleshy, edible stalks ( petioles) of species and hybrids (culinary rhubarb) of '' Rheum'' in the family Polygonaceae, which are cooked and used for food. The whole plant – a herbaceous perennial growing from short, thick rhi ...

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 ...

s, venus fly traps, and

bougainvillea ''Bougainvillea'' ( , ) is a genus of thorny ornamental vines, bushes, and trees belonging to the four o' clock family, Nyctaginaceae. It is native to eastern South America, found from Brazil, west to Peru, and south to southern Argentina. ...

. Recent molecular and biochemical evidence has resolved additional well-supported

clades A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English te ...

within the Caryophyllales. * order Caryophyllales ** family Achatocarpaceae ** family

** family

Pupalia lappacea (Forest Burr) W IMG 1562

** family Anacampserotaceae (added in APG III) ** family Ancistrocladaceae ** family

** family

** family Basellaceae ** family

** family

** family Didiereaceae ** family Dioncophyllaceae ** family

** family

** family Frankeniaceae ** family

** family

(added in APG III) ** family

(added in APG III) ** family Molluginaceae Glinus oppositifolius (Bitter cumin) W2 IMG 0462

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

** family

** family Portulacaceae ** family Rhabdodendraceae ** family Sarcobataceae ** family Simmondsiaceae ** family

** 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 Caryophyllidae is a botanical name at the rank of subclass. At the moment there is no complete consensus about what orders it includes, except that it presumably contains the order Caryophyllales. Note that this is only a naming difficulty: w ...

''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 Nepenthales (Nepenthales Bercht. & J.Presl) is an order of carnivorous flowering plants in the Cronquist system of plant classification. Cronquist system The order was placed in the subclass Dilleniidae, which in the 1981 version of this s ...

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

References

External links

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