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Caryophyllales
	Dianthus caryophyllus
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Superasterids
Order:	Caryophyllales; Juss. ex Bercht. & J.Presl
Suborders
	Caryophyllineae; Polygonineae; Portulacineae
Synonyms
	Centrospermae

Caryophyllales (/ˌkæri.oʊfɪˈleɪliːz/ KARR-ee-oh-fih-LAY-leez)^[2] is a diverse and heterogeneous order of flowering plants that includes the cacti, carnations, amaranths, ice plants, beets, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves. The betalain pigments are unique in plants of this order and occur in all its core families with the exception of Caryophyllaceae and Molluginaceae.^[3] Noncore families, such as Nepenthaceae, instead produce anthocyanins.^[4]

In its modern definition, the order encompasses a whole new group of families (formerly included in the order Polygonales) that never synthesize betalains, among which several families are carnivorous (like Nepenthaceae and Droseraceae).

According to molecular clock calculations, the lineage that led to Caryophyllales split from other plants about 111 million years ago.^[5]

Description

The members of Caryophyllales include about 6% of eudicot species.^[6] This order is part of the core eudicots.^[7] Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species^[8] The monophyly of the Caryophyllales has been supported by DNA sequences, cytochrome c sequence data and heritable characters such as anther wall development and vessel-elements with simple perforations.^[9]

Circumscription

As with all taxa, 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.^[9] 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.' "^[10]

Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae.^[9] These two suborders were formerly (and sometimes still are) recognized as two orders, Polygonales and Caryophyllales.^[9]

APG IV

Kewaceae, Macarthuriaceae, Microteaceae, Cactaceae, and Petiveriaceae were added in APG IV. ^[11]

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.^[1]

family Achatocarpaceae
family Aizoaceae
family Amaranthaceae
family Anacampserotaceae
family Ancistrocladaceae
family Asteropeiaceae
family Barbeuiaceae
family Basellaceae
family Cactaceae
family Caryophyllaceae
family Didiereaceae
family Dioncophyllaceae
family Droseraceae
family Drosophyllaceae
family Frankeniaceae
family Gisekiaceae
family Halophytaceae
family Kewaceae
family Limeaceae
family Lophiocarpaceae
family Macarthuriaceae
family Microteaceae
family Molluginaceae
family Montiaceae
family Nepenthaceae
family Nyctaginaceae
family Petiveriaceae
family Physenaceae
family Phytolaccaceae
family Plumbaginaceae
family Polygonaceae
family Portulacaceae
family Rhabdodendraceae
family Sarcobataceae
family Simmondsiaceae
family Stegnospermataceae
family Talinaceae
family Tamaricaceae

APG II

As circumscribed by the APG II system (2003), this order includes well-known plants like cacti, carnations, spinach, beet, rhubarb, sundews, venus fly traps, 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 from family Amaranthaceae
- family Anacampserotaceae (added in APG III)^[1]
- family Ancistrocladaceae
- family Asteropeiaceae
- family Barbeuiaceae
- family Basellaceae
- family Cactaceae
- family Caryophyllaceae
- family Didiereaceae
- family Dioncophyllaceae
- family Droseraceae
- family Drosophyllaceae
- family Frankeniaceae
- family Gisekiaceae
- family Halophytaceae
- family Limeaceae (added in APG III)^[1]
- family Lophiocarpaceae (added in APG III)^[1]
- family Molluginaceae
  Glinus oppositifolius from family Molluginaceae
- family Montiaceae (added in APG III)^[1]
- family Nepenthaceae
- family Nyctaginaceae
- family Physenaceae
- family Phytolaccaceae
- family Plumbaginaceae
- family Polygonaceae
- family Portulacaceae
- family Rhabdodendraceae
- family Sarcobataceae
- family Simmondsiaceae
- family Stegnospermataceae
- family Talinaceae (added in APG III)^[1]
- family Tamaricaceae

APG

This represents a slight change from the APG system, 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 (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, last edition in 1935, and the Engler system, updated in 1964, had a similar order under the name Centrospermae.

References

^ ^a ^b ^c ^d ^e ^f ^g Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
^ Clarke, Ian; Lee, Helen (2003). Name that Flower: The Identification of Flowering Plants. Melbourne University Publishing. p. 56. ISBN 978-0-522-85060-4.
^ Kubitzki, Klaus; Bayer, Clemens; Cuénoud, Philippe (January 2003). Flowering Plants · Dicotyledons: Malvales, Capparales and Non-betalain Caryophyllales. Springer. pp. 1–4. ISBN 978-3-642-07680-0.
^ Dávila-Lara, Alberto (2021). "Proof of anthocyanins in the carnivorous plant genus Nepenthes". FEBS Open Bio. 11 (9): 2576–2585. doi:10.1002/2211-5463.13255. PMC 8409308. PMID 34289256.
^ Susana Magallón & Amanda Castillo (2009), "Angiosperm diversification through time", American Journal of Botany, 96 (1): 349–365, doi:10.3732/ajb.0800060, PMID 21628193
^ "Caryophyllales". Angiosperm Phylogeny Website.
^ Judd., W.; Campbell, C.; Kellogg, E.; Stevens, P.; Donoghue, M. (2008). Plant Systematics: A Phylogenetic Approach (3rd ed.). W. H. Freeman. ISBN 978-0-87893-407-2.
^ Stephens, P.F. (2020). "Angiosperm Phylogeny Website". Version 14. Retrieved 10 June 2020.
^ ^a ^b ^c ^d Juan, R.; Pastor, J.; Alaiz, M.; Vioque, J. (1 September 2007). "Electrophoretic characterization of Amaranthus L. seed proteins and its systematic implications". Botanical Journal of the Linnean Society. 155 (1): 57–63. doi:10.1111/j.1095-8339.2007.00665.x.
^ Lawrence, G.H.M (1960). Taxonomy of Vascular Plants. Macmillan. p. 660.
^ Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.