The betalain pigments are unique in plants of this order and occur in all its core families with the exception of Caryophyllaceae and Molluginaceae.[2] Noncore families, such as Nepenthaceae, instead produce anthocyanins.[3] 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.[4]
The members of Caryophyllales include about 6% of eudicotspecies.[5] This order is part of the core eudicots.[6] Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species[7] 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.[8]
Taxonomy and Phylogeny
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.Script error: No such module "Unsubst".
Systematists were undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid clade.[8] 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.' "[9]
Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae.[8] These two suborders were formerly (and sometimes still are) recognized as two orders, Polygonales and Caryophyllales.[8]
File:Wigginsia Pauciareolata.JPGCactaceae native to the middle region of South America, at Marsh Botanical Garden. Cactaceae are a plant family, under the order Caryophyllales.
Angiosperm Phylogeny Group (APG)
As of the APG IV system of classification (2016) the order Caryophyllales contains 38 families:[10]
26 of the 38 families were included in the original APG system (1998) and the remaining 12 were added during later revisions: three in the APG II system (2003), five in the APG III system (2009), and four in the APG IV system (2016). Families added during APG revisions are so noted above.
The earlier Cronquist system (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 Caryophyllidaesensu Cronquist.Script error: No such module "Unsubst".
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.Script error: No such module "Unsubst".
New to the order (sensu APG) are the Asteropeiaceae and Physenaceae, each containing a single genus, and two genera from Cronquist's order Nepenthales.Script error: No such module "Unsubst".
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.Script error: No such module "Unsubst".
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