Caryophyllales

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Ordo: Caryophyllales
Familiae: Achatocarpaceae – Aizoaceae – Amaranthaceae – Anacampserotaceae – Ancistrocladaceae – Asteropeiaceae – Barbeuiaceae – Basellaceae – Cactaceae – Caryophyllaceae – Corbichoniaceae – Didiereaceae – Dioncophyllaceae – Droseraceae – Drosophyllaceae – Frankeniaceae – Gisekiaceae – Halophytaceae – Kewaceae – Limeaceae – Lophiocarpaceae – Macarthuriaceae – Microteaceae – Molluginaceae – Montiaceae – Nepenthaceae – Nyctaginaceae – Petiveriaceae – Physenaceae – Phytolaccaceae – Plumbaginaceae – Polygonaceae – Portulacaceae – Rhabdodendraceae – Sarcobataceae – Simmondsiaceae – Stegnospermataceae – Talinaceae – Tamaricaceae
Incertae sedis paleogenera: †Gomphrenipollis

Name

Caryophyllales Juss. ex Bercht. & J. Presl Prir. Rostlin 239. (1820)
Synonyms

Heterotypic
Aizoales
Amaranthales
Ancistrocladales
Atriplicales
Cactales Juss. ex Bercht. & J. Presl, Přir. Rostlin: 238. 1820.
Type genus: Cactus L., nom. rej.
Chenopodiales
Dioncophyllales
Droserales
Nepenthales
Nyctaginales
Opuntiales
Petiveriales
Physenales
Phytolaccales
Plumbaginales
Polygonales
Portulacales
Rhabdodendrales
Scleranthales
Silenales
Simmondsiales
Stellariales
Tamaricales

References

Jussieu, A.L. de 1820. O Prirozenosti Rostlin 239. 1820
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 Reference page.
Borsch, T., Hernández-Ledesma, P., Berendsohn, W.G., Flores Olvera, M.H., Ochoterena, H., Zuloaga, F.O., Mering, S. v. & Kilian, N. 2015. An integrative and dynamic approach for monographing species-rich plant groups – Building the global synthesis of the angiosperm order Caryophyllales. Perspectives in Plant Ecology, Evolution and Systematics 17(4): 284–300. DOI: 10.1016/j.ppees.2015.05.003 PDF Reference page.
Cameron, K.M., Wurdack, K.J. & Jobson, R.W. 2002. Molecular evidence for the common origin of snap-traps among carnivorous plants, American Journal of Botany 89: 1503–1509. DOI: 10.3732/ajb.89.9.1503 Full Text PDF. Reference page.
Christenhusz, M.J.M., Brockington, S.F., Christin, P.A. & Sage, R.F. 2014. On the disintegration of Molluginaceae: a new genus and family (Kewa, Kewaceae) segregated from Hypertelis, and placement of Macarthuria in Macarthuriaceae. Phytotaxa 181(4): 238–242. DOI: 10.11646/phytotaxa.181.4.4 Full text PDF from ResearchGate Reference page.
Cuénoud, P. 2002. Introduction to expanded Caryophyllales. Pp. 1-4, in Kubitzki, K. (ed.), The Families and Genera of Vascular Plants. V. Flowering Plants. Dicotyledons. Malvales, Capparales and Non-betalain Caryophyllales. Springer, Berlin.
Hernández-Ledesma, P., Berendsohn, W. G., Borsch, T., Mering, S. v., Akhani, H., Arias, S., Castañeda-Noa, I., Eggli, U., Eriksson, R., Flores-Olvera, H., Fuentes-Bazán, S., Kadereit, G., Klak, C., Korotkova, N., Nyffeler R., Ocampo G., Ochoterena, H., Oxelman, B., Rabeler, R. K., Sanchez, A., Schlumpberger, B. O. & Uotila, P. 2015. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45(3): 281–383. DOI: 10.3372/wi.45.45301 Open access Reference page.
Nyffeler, R. & Eggli, U. 2010. Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59(1): 227–240. Abstract. Full text (PDF). Reference page.
Schäferhoff, B., Müller, K.F. & Borsch, T. 2010. Caryophyllales phylogenetics: disentangling Phytolaccaceae and Molluginaceae and description of Microteaceae as a new isolated family. Willdenowia 39(2): 209-228. DOI: 10.3372/wi.39.39201 Full text PDF Reference page.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.
Thulin, M., Moore, A.J., El-Seedi, H., Larsson, A., Christin, P.A. & Edwards, E.J. 2016. Phylogeny and generic delimitation in Molluginaceae, new pigment data in Caryophyllales, and the new family Corbichoniaceae. Taxon 65(4): 775–793. DOI: 10.12705/654.6 Full text PDF from ResearchGate Reference page.
Tropicos.org 2014. Caryophyllales. Missouri Botanical Garden. Published on the internet. Accessed: 2014 Jan. 29.

Vernacular names
العربية: قرنفليات
azərbaycanca: Qərənfilçiçəklilər
беларуская: Гваздзікакветныя
català: Cariofil·lal
čeština: hvozdíkotvaré
dansk: Nellike-ordenen
Deutsch: Nelkenartige
español: Cariofilales
eesti: Nelgilaadsed
فارسی: میخک‌سانان
עברית: ציפורנאים
hrvatski: Klinčićolike
magyar: Szegfűvirágúak
italiano: Cariofillali
日本語: ナデシコ目
한국어: 석죽목
kurdî: Koma mêxikan
lietuvių: Gvazdikiečiai
latviešu: Neļķu rinda
македонски: Каранфиловидни
polski: Goździkowce, Śródłożne
русский: Гвоздичноцветные
slovenčina: klinčekotvaré
svenska: Nejlikordningen
ไทย: อันดับคาร์เนชัน
Türkçe: Karanfilgiller
українська: Гвоздикоцвіті
Tiếng Việt: Bộ Cẩm chướng
中文: 石竹目

Caryophyllales (/ˌkærioʊfɪˈleɪliːz/ KARR-ee-oh-fil-AY-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 families with the exception of Caryophyllaceae and Molluginaceae. [3]

Description

The members of Caryophyllales include about 6% of eudicot species.[4] This order is part of the core eudicots.[5] Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species[6] 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.[7]
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.[7] 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.' "[8]

Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae.[7] These two suborders were formerly (and sometimes still are) recognized as two orders, Polygonales and Caryophyllales.[7]
Cactaceae native to the middle region of South America, at Marsh Botanical Garden. Cactaceae are a plant family, under the order Caryophyllales.
APG IV

Kewaceae, Macarthuriaceae, Microteaceae, and Petiveriaceae were added in APG IV. [9]
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

Cactaceaeː Gymnocalycium Matoensea at Yale's Marsh Botanical Garden.
APG
Carnegiea gigantea
Sweet William Dwarf from the family Caryophyllaceae
A flower of Dianthus

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
Chenopodium album

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

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.
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. pp. 1–4. ISBN 978-3-642-07680-0.
"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.
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.