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]
| Caryophyllales | |
|---|---|
| |
| Dianthus caryophyllus | |
Scientific classification 
| |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Superasterids |
| Order: | Caryophyllales Juss. ex Bercht. & J.Presl[1] |
| Suborders | |
| Synonyms | |
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
editThe 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
editAs 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
editKewaceae, Macarthuriaceae, Microteaceae, Cactaceae, and Petiveriaceae were added in APG IV. [11]
APG III
editAs 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
editAs 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
editThis 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
editThe 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
editEarlier 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
edit- ^ 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.
External links
edit- Tree of Life Characteristics and Phylogenetic Relationships