Caryophyllales ( ) is a diverse and heterogeneous

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

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 with well-known members including

cacti 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, ...

carnation ''Dianthus caryophyllus'' ( ), commonly known as carnation or clove pink, is a species of ''Dianthus'' native to the Mediterranean Basin, Mediterranean region. Its exact natural range is uncertain due to extensive cultivation over the last 2,00 ...

beets The beetroot (British English) or beet (North American English) is the taproot portion of a ''Beta vulgaris'' subsp. ''vulgaris'' plant in the Conditiva Group. The plant is a root vegetable also known as the table beet, garden beet, dinner beet ...

quinoa Quinoa (''Chenopodium quinoa''; , from Quechuan languages, Quechua ' or ') is a flowering plant in the Amaranthaceae, amaranth family. It is a herbaceous annual plant grown as a crop primarily for its edible seeds; the seeds are high in prote ...

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 ...

amaranth ''Amaranthus'' is a cosmopolitan distribution, cosmopolitan group of more than 50 species which make up the genus of annual plant, annual or short-lived perennial plants collectively known as amaranths. Some names include "prostrate pigweed" an ...

s, pigfaces and ice plants, oraches and saltbushes, goosefoots, sundews,

Venus flytrap The Venus flytrap (''Dionaea muscipula'') is a carnivorous plant native to the temperate and subtropical wetlands of North Carolina and South Carolina, on the East Coast of the United States. Although various modern hybrids have been created ...

s, monkey cup pitcher plants,

Malabar spinach ''Basella alba'' is an edible perennial vine in the family Basellaceae. It is found in tropical Asia and Africa where it is widely used as a leaf vegetable. It is native to the Indian subcontinent, Southeast Asia and New Guinea. It is naturaliz ...

bougainvillea ''Bougainvillea'' ( , ) is a genus of thorny ornamental vines, bushes, and trees belonging to the family, Nyctaginaceae. They are native to Brazil, Bolivia, Paraguay, Peru, and Argentina. There are between 4 and 22 species in the genus. The i ...

s, four o'clock flowers,

buckwheat Buckwheat (''Fagopyrum esculentum'') or common buckwheat is a flowering plant in the knotweed family Polygonaceae cultivated for its grain-like seeds and as a cover crop. Buckwheat originated around the 6th millennium BCE in the region of what ...

knotweed Knotweed is a common name for plants in several genera in the family Polygonaceae. Knotweed may refer to: * ''Fallopia'' * ''Persicaria'' * ''Polygonum'' * ''Reynoutria'' ** ''Reynoutria japonica'' or Japanese knotweed, a highly invasive species in ...

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 plant is a herbaceous perennial that grows from short, thick rhizomes. ...

sorrel Sorrel (''Rumex acetosa''), also called common sorrel or garden sorrel, is a perennial herbaceous plant in the family Polygonaceae. Other names for sorrel include spinach dock and narrow-leaved dock ("dock" being a common name for the genus ''Ru ...

portulaca ''Portulaca'' () is a genus of flowering plants in the family Portulacaceae, and is the type genus of the family. With over 100 species, it is found in the tropics and warm temperate regions. Portulacas are also known as the purslanes. Common p ...

jojoba Jojoba (; botanical name: ''Simmondsia chinensis'')also commonly called goat nut, deer nut, pignut, wild hazel, quinine nut, coffeeberry, and gray box bushis an evergreen, dioecious shrub native to the Southwestern United States and northern M ...

, and

tamarisk The genus ''Tamarix'' (tamarisk, salt cedar, taray) is composed of about 50–60 species of flowering plants in the family Tamaricaceae, native to drier areas of Eurasia and Africa. The generic name originated in Latin and may refer to the Tamb ...

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 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, ...

. 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 o ...

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

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

and

Molluginaceae The Molluginaceae are a family (biology), 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 ...

. Noncore families, such as Nepenthaceae, instead produce anthocyanins. In its modern definition, the order encompasses a whole new group of families (formerly included in the order

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: * ord ...

) that never synthesize betalains, among which several families are carnivorous (like Nepenthaceae and

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, the vast majority being in the sundew genus '' Drosera''. The family also contains the wel ...

). According to

molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleot ...

calculations, the lineage that led to Caryophyllales split from other plants about 111 million years ago.

Description

The members of Caryophyllales include about 6% of

eudicot 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 ...

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), ...

. This order is part of the

core eudicot 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 ...

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 within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the ...

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

Taxonomy and Phylogeny

As with all

taxa In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...

, 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 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, and occasionally small mammals and birds. They have adapted to grow in waterlo ...

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

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

. 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 (: locules) or loculus (; : loculi) 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'' usually refers to a chamber within an ovary ...

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,

and Caryophyllales. Wigginsia Pauciareolata

Angiosperm Phylogeny Group (APG)

As of the

APG IV system The APG IV system of flowering plant classification is the fourth version of a modern, mostly molecular-based, system of plant taxonomy for flowering plants (angiosperms) being developed by the Angiosperm Phylogeny Group (APG). It was publish ...

of classification (2016) the order Caryophyllales contains 38 families: * family

Achatocarpaceae The Achatocarpaceae are a family of woody flowering plants consisting of two genera and 11 known species, and has been recognized by most taxonomists. The family is found from the southwestern United States south to tropical and subtropical Sout ...

* family

Aizoaceae The Aizoaceae (), or fig-marigold family, is a large family of dicotyledonous flowering plants containing 135 genera and about 1,800 species. Several genera are commonly known as 'ice plants' or 'carpet weeds'. The Aizoaceae are also referred to a ...

* 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, maki ...

* 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 Caryophy ...

(added in APG III) * family

Ancistrocladaceae ''Ancistrocladus'' is a genus of woody lianas in the Monotypic taxon, 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. Classific ...

* family Asteropeiaceae * 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 ...

(added in APG II) * family

Basellaceae Basellaceae is a family (biology), 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: * ''An ...

* family

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, ...

* family

Didiereaceae Didiereaceae is a family of flowering plants found in continental Africa and Madagascar. It contains 20 species classified in three subfamilies and six genera. Species of the family are succulent plants, growing in sub-arid to arid habitats. Seve ...

* family

Dioncophyllaceae The Dioncophyllaceae are a 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 carnivorous plant ...

* family

* family Drosophyllaceae * family

Frankeniaceae ''Frankenia'' (sea heath) is the only genus in the Frankeniaceae family of flowering plants. Other genera have been recognized within the family, such as ''Anthobryum'', ''Hypericopsis'' and ''Niederleinia'', but molecular phylogenetic studies h ...

* 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 ...

(added in APG II) * 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 solitary f ...

(added in APG II) * family

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, ...

(added in APG IV) * family

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

(added in APG III) * 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 ...

(added in APG III) * family

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 ...

(added in APG IV) * family

Microteaceae ''Microtea'', the jumby peppers, are a genus of flowering plants in the family Microteaceae, native to the Caribbean islands, Central America, and tropical South America. Species Ten species are currently accepted. *''Microtea bahiensis'' *''M ...

(added in APG IV) * 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 ...

(added in APG III) * 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 distinctive fr ...

* family

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 A species () is often defined as the largest group of organisms in whi ...

(added in APG IV) * family Physenaceae * family

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

* family

Plumbaginaceae Plumbaginaceae is a family (biology), 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 plant, perennial h ...

* family

Polygonaceae The Polygonaceae are a Family (biology), family of flowering plants known Common name, informally as the knotweed family or smartweed—buckwheat family in the United States. The Botanical name, name is Basionym, based on the genus ''Polygonum'', ...

* 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 of flowering plant in the family Rhabdodendraceae. It comprises three species of Tropics, tropical South American trees. It is placed in its own family (biology), family, Rhabdodendraceae, which has only been recogni ...

* family

Sarcobataceae ''Sarcobatus'' is a North American genus of two species of flowering plants, formerly considered to be a single species. Common names for ''S. vermiculatus'' include greasewood, seepwood, and saltbush. Traditionally, ''Sarcobatus'' has been ...

* 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, o ...

* 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 Talinaceae is a family of two genera and 28 species of flowering plants comprising shrubs, lianas, and herbaceous species native to the Americas, Africa and Madagascar. The family is newly recognized through research by the Angiosperm Phylogeny ...

(added in APG III) * 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 ...

26 of the 38 families were included in the original

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 ...

(1998) and the remaining 12 were added during later revisions: three in 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 ...

(2003), five in 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 f ...

(2009), and four in the

(2016). Families added during APG revisions are so noted above.

Cronquist

The earlier

Cronquist system The Cronquist system is a list of systems of plant taxonomy, taxonomic classification system of angiosperms, flowering plants. It was developed by Arthur Cronquist in a series of monographs and texts, including ''The Evolution and Classification of ...

(1981) recognised the order with 12 families: * 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 Nepenthales (Nepenthales Bercht. & J.Presl) is an order of carnivorous flowering plants in the Cronquist system of plant classification. Taxonomy The order Nepenthales as of 2018 is a clade within Caryophyllales and contains the following f ...

Earlier circumscriptions

Earlier systems, such as the

Wettstein system A list of systems of plant taxonomy, 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 Syn ...

, 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 Eichler, meaning "with the seed in the center", referring to the free (central) placentation. It was used in the Engler system and the Wettstein system) for an order of fl ...

References

External links

Tree of Life
Characteristics and Phylogenetic Relationships {{Authority control Superasterids Angiosperm orders