Thebetalain pigments are unique in plants of this order and occur in all its core families with the exception ofCaryophyllaceae andMolluginaceae.[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 orderPolygonales) that never synthesize betalains, among which several families are carnivorous (likeNepenthaceae andDroseraceae).
According tomolecular clock calculations, the lineage that led to Caryophyllales split from other plants about 111 million years ago.[5]
The members of Caryophyllales include about 6% ofeudicotspecies.[6] This order is part of thecore eudicots.[7] Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species[8] The monophyly of the Caryophyllales has been supported byDNA sequences, cytochrome c sequence data and heritable characters such as anther wall development and vessel-elements with simple perforations.[9]
As with alltaxa, 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 manycarnivorous plants.[citation needed]
Systematists were undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asteridclade.[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 oflocules 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]
Cactaceae native to the middle region of South America, at Marsh Botanical Garden. Cactaceae are a plant family, under the order Caryophyllales.
Cactaceaeː Gymnocalycium Matoensea at Yale's Marsh Botanical Garden.
26 of the 38 families were included in the originalAPG system (1998) and the remaining 12 were added during later revisions: three in theAPG II system (2003), five in theAPG III system (2009), and four in theAPG IV system (2016). Families added during APG revisions are so noted above.
The earlierCronquist 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 Caryophyllalessensu APG should rather be compared to subclassCaryophyllidaesensu Cronquist.[citation needed]
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.[citation needed]
New to the order (sensuAPG) are the Asteropeiaceae and Physenaceae, each containing a single genus, and two genera from Cronquist's orderNepenthales.[citation needed]
^abcdJuan, R.; Pastor, J.; Alaiz, M.; Vioque, J. (1 September 2007). "Electrophoretic characterization ofAmaranthus 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.