Caryophyllales

Caryophyllales (/ˌkærioʊfɪˈleɪliːz/ KARR-ee-oh-fil-AY-leez)[2] is an 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.

Caryophyllales
Dianthus caryophyllus L (Clove pink)
Dianthus caryophyllus
Scientific classification
Kingdom: Plantae
Clade: Angiosperms
Clade: Eudicots
Clade: Superasterids
Order: Caryophyllales
Juss. ex Bercht. & J.Presl[1]
Suborders

Caryophyllineae
Polygonineae

Synonyms

Centrospermae

Description

The members of Caryophyllales include about 6% of eudicot species.[3] This order is part of the core eudicots.[4] Currently, the Caryophyllales contains 33 families, 692 genera and 11,155 species.[5] 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.[6]

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.

Although the monophyly of the order has been strongly supported, their placement is still uncertain. Systematists are undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid clade.[6] 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.' "[7]

Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae.[6] These two suborders were formerly (and sometimes still are) recognized as two orders, Polygonales and Caryophyllales.[6]

Wigginsia Pauciareolata
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. [8]

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]

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.

Pupalia lappacea (Forest Burr) W IMG 1562
Pupalia lappacea Forest Burr from family Amaranthaceae
Glinus oppositifolius (Bitter cumin) W2 IMG 0462
Glinus oppositifolius from family Molluginaceae
Cactaceaeː Gymnocalycium Matoensea
Cactaceaeː Gymnocalycium Matoensea at Yale's Marsh Botanical Garden.

APG

Carnegiea gigantea top
Carnegiea gigantea
Spring Flowers
Sweet William Dwarf from the family Caryophyllaceae
Flower dianthus
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

Melganzenvoet bloeiwijze Chenopodium album
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

  1. ^ 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. Archived from the original (PDF) on 2017-05-25. Retrieved 2013-07-06.
  2. ^ Clarke, Ian; Lee, Helen (2003). Name that Flower: The Identification of Flowering Plants. Melbourne University Publishing. p. 56. ISBN 978-0-522-85060-4.
  3. ^ "Caryophyllales". Angiosperm Phylogeny Website.
  4. ^ 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.
  5. ^ Stephens, P.F. (2001). "Angiosperm Phylogeny Website. Version 8, June 2007".
  6. ^ 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.
  7. ^ Lawrence, G.H.M (1960). Taxonomy of Vascular Plants. Macmillan. p. 660.
  8. ^ 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

Aizoaceae

The Aizoaceae Martynov, nom. cons. (fig-marigold family) is a large family of dicotyledonous flowering plants containing 135 genera and about 1800 species. They are commonly known as ice plants or carpet weeds. They are often called vygies in South Africa and New Zealand. Highly succulent species that resemble stones are sometimes called mesembs.

Amaranthaceae

Amaranthaceae is a family of flowering plants commonly known as the amaranth family, in reference to its type genus Amaranthus. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making it the most species-rich lineage within its parent order, Caryophyllales.

Cacteae

Cacteae is a tribe of plants belonging to the family Cactaceae found mainly in North America especially Mexico. As of August 2018, the internal classification of the family Cactaceae remained uncertain and subject to change. A classification incorporating many of the insights from the molecular studies was produced by Nyffeler and Eggli in 2010.

Cactoideae

The Cactoideae are the largest subfamily of the cactus family, Cactaceae. Around 80% of cactus species belong to this subfamily. As of August 2018, the internal classification of the family Cactaceae remained uncertain and subject to change. A classification incorporating many of the insights from the molecular studies was produced by Nyffeler and Eggli in 2010.

Caryophyllaceae

Caryophyllaceae, commonly called the pink family or carnation family, is a family of flowering plants. It is included in the dicotyledon order Caryophyllales in the APG III system, alongside 33 other families, including Amaranthaceae, Cactaceae, and Polygonaceae. It is a large family, with 81 genera and about 2,625 known species.This cosmopolitan family of mostly herbaceous plants is best represented in temperate climates, with a few species growing on tropical mountains. Some of the more commonly known members include pinks and carnations (Dianthus), and firepink and campions (Lychnis and Silene). Many species are grown as ornamental plants, and some species are widespread weeds. Most species grow in the Mediterranean and bordering regions of Europe and Asia. The number of genera and species in the Southern Hemisphere is rather small, although the family does contain Antarctic pearlwort (Colobanthus quitensis), the world's southernmost dicot, which is one of only two flowering plants found in Antarctica.The name comes from Caryophyllus, an obsolete synonym of Dianthus.

Cerastium nigrescens

Cerastium nigrescens, commonly known as the Shetland mouse-ear, Shetland mouse-eared chickweed or Edmondston's chickweed, is an endemic plant found in Shetland, Scotland.

It was first recorded in 1837 by botanist Thomas Edmondston, who was 12 at the time. For a long time it was synonymised with Arctic Mouse-ear Cerastium arcticum but it is now widely regarded as a separate species. Although reported from two other sites in the 19th century, it currently grows only on two serpentine hills on the island of Unst (see Keen of Hamar).

The numbers of C. nigrescens can vary dramatically from year to year, for reasons that are unclear (probably due to a varying rates of seedling germination and survival), but the underlying trend seems stable, and there has been no change in its distribution.

Mature plants may be not much more than a single shoot with one flower or can be a fist-sized cushion with as many as 40 flowers. Flowers look disproportionately large compared with the size of the plant.

Limonium

Limonium is a genus of 120 flowering plant species. Members are also known as sea-lavender, statice, caspia or marsh-rosemary. Despite their common names, species are not related to the lavenders or to rosemary. They are instead in Plumbaginaceae, the plumbago or leadwort family.

The generic name is from the Latin līmōnion, used by Pliny for a wild plant and is ultimately derived from the Ancient Greek leimon (λειμών, ‘meadow’).

Montiaceae

Montiaceae are a family of flowering plants, comprising about 14 genera with about 230 known species, ranging from herbaceous plants to shrubs. The family has a cosmopolitan distribution.

The family Montiaceae was newly adopted in the APG III system and includes members of the Caryophyllales formerly listed in Portulacaceae.

Nepenthes parvula

Nepenthes parvula is a tropical pitcher plant native to the Cape York Peninsula of Queensland, Australia.The specific epithet parvula refers to the diminutive size of mature plants.

Nepenthes rowaniae

Nepenthes rowaniae (; after Ellis Rowan, Australian naturalist and illustrator) is a species of pitcher plant endemic to the Cape York Peninsula, Australia. It is closely related to N. mirabilis and was once considered an extreme form of this species.

Nepenthes tenax

Nepenthes tenax (; from Latin: tenax "tenacious") is a lowland species of tropical pitcher plant native to northern Queensland, Australia. It is the third Nepenthes species recorded from the continent and its second endemic species. Nepenthes tenax is closely related to the two other Australian Nepenthes species: N. mirabilis and N. rowaniae.

Nepenthes tenax grows to a height of around 100 cm with pitchers rarely exceeding 15 cm. The stem is usually self-supporting. In its natural habitat, it is sympatric with N. mirabilis and N. rowaniae. Simple and complex natural hybrids involving both of these species have been found.

Nyctaginaceae

Nyctaginaceae, the four o'clock family, is a family of around 33 genera and 290 species of flowering plants, widely distributed in tropical and subtropical regions, with a few representatives in temperate regions. The family has a unique fruit type, called an "anthocarp", and many genera have extremely large (>100 µm) pollen grains.

The family has been almost universally recognized by plant taxonomists. The APG II system (2003; unchanged from the APG system of 1998), assigns it to the order Caryophyllales in the clade core eudicots.

A phylogenetic study by Levin has justified the combination of Selinocarpus and Ammocodon into the genus Acleisanthes. The genus Izabalea is now considered a synonym of Agonandra, a genus in Opiliaceae. A more recent study by Douglas and Manos clarified the relationships among almost all of the genera in the family and demonstrated that a substantial diversification of herbaceous genera has occurred in arid North America. Many genera of Nyctaginaceae possess unusual characters. Notable examples include sticky bands on the stems between the nodes, cleistogamous flowers (which self-pollinate without opening), or gypsophily, the ability to grow on soils with a high concentration of gypsum.

Phytolaccaceae

Phytolaccaceae is a family of flowering plants. Such a family has been almost universally recognized by taxonomists, although its circumscription has varied. It is also known as the Pokeweed family.

The APG II system, of 2003 (unchanged from the APG system, of 1998), also recognizes this family and assigns it to the order Caryophyllales in the clade core eudicots. The family comprises five genera, totalling 33 known species. It is divided into the subfamilies Agdestioideae and Phytolaccoideae, with the former Rivinioideae in the Takhtajan system. now placed in its own family Petiveriaceae

Plumbaginaceae

Plumbaginaceae is a family of flowering plants, with a cosmopolitan distribution. The family is sometimes referred to as the leadwort family or the plumbago family.

Most species in this family are perennial herbaceous plants, but a few grow as lianas or shrubs. The plants have perfect flowers and are pollinated by insects. They are found in many different climatic regions, from arctic to tropical conditions, but are particularly associated with salt-rich steppes, marshes, and sea coasts.

The family has been recognized by most taxonomists. The APG II system (2003; unchanged from the APG system of 1998), recognizes this family and assigns it to the order Caryophyllales in the clade core eudicots. It includes ca 30 genera and about 725 species.The 1981 Cronquist system placed the family in a separate order Plumbaginales, which included no other families. The Dahlgren system had segregated some of these plants as family Limoniaceae.

Polygonaceae

The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus Polygonum, and was first used by Antoine Laurent de Jussieu in 1789 in his book, Genera Plantarum. The name refers to the many swollen nodes the stems of some species have. It is derived from Greek; poly means many and goni means knee or joint.

The Polygonaceae comprise about 1200 species distributed into about 48 genera. The largest genera are Eriogonum (240 species), Rumex (200 species), Coccoloba (120 species), Persicaria (100 species) and Calligonum (80 species). The family is present worldwide, but is most diverse in the North Temperate Zone.

Several species are cultivated as ornamentals. A few species of Triplaris provide lumber. The fruit of the sea grape (Coccoloba uvifera) is eaten, and in Florida, jelly is made from it and sold commercially. The seeds of two species of Fagopyrum, known as buckwheat (sarrasin in French), provide grain (its dark flour is known as blé noir (black wheat) in France). The petioles of rhubarb (Rheum rhabarbarum and hybrids) are a food item. The leaves of the common sorrel (Rumex acetosa) are eaten in salads or as a leaf vegetable.Polygonaceae contain some of the worst weeds, including species of Persicaria, Rumex and Polygonum, such as Japanese knotweed.

Portulacaceae

The Portulacaceae are a family of flowering plants, comprising 115 species in a single genus Portulaca. Formerly some 20 genera with about 500 species, were placed there, but it is now restricted to encompass only one genus, the other genera being placed elsewhere. The family has been recognised by most taxonomists, and is also known as the purslane family; it has a cosmopolitan distribution, with the highest diversity in semiarid regions of the Southern Hemisphere in Africa, Australia, and South America, but with a few species also extending north into Arctic regions. The family is very similar to the Caryophyllaceae, differing in the calyx, which has only two sepals.

The APG II system (2003; unchanged from the APG system of 1998) assigns it to the order Caryophyllales in the clade core eudicots. In the APG III system, several genera were moved to the Montiaceae, Didiereaceae, Anacampserotaceae and Talinaceae, thus making the family monotypic and only containing the genus Portulaca.

Superasterids

The superasterids are members of a large clade (monophyletic group) of flowering plants, containing more than 122,000 species.

The clade is divided into 20 orders as defined in APG IV system. These orders, in turn, together comprise about 146 families.The name is based upon the name "Asteridae", which had usually been understood to be a subclass.

Tamaricaceae

The Tamaricaceae are a flowering plant family (the tamarisk family) containing four genera and a total of 78 known species. In the 1980s, the family was classified in the Violales under the Cronquist system; more modern classifications (Angiosperm Phylogeny Group) place them in the Caryophyllales.

The family is native to drier areas of Europe, Asia, and Africa. Many grow on saline soils, tolerating up to 15,000 ppm soluble salt and can also tolerate alkaline conditions. The leaves are generally scale-like, measure 1–5 mm long, overlap each other along the stem, and in some species are encrusted with salt secretions.

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