Camphorosmeae is a species-rich tribe of the

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 i ...

, formerly Chenopodiaceae, with 20 genera and about 179 species. It is classified as a single

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

of subfamily Camphorosmoideae.

Description

The Camphorosmeae are mostly

dwarf shrub A subshrub (Latin ''suffrutex'') or dwarf shrub is a short shrub, and is a woody plant. Prostrate shrub is a related term. "Subshrub" is often used interchangeably with "bush".Jackson, Benjamin, Daydon; A Glossary of Botanic Terms with their Der ...

s or annuals (rarely

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...

herbs) with spreading or ascending branches. The plants are more or less densely covered with appressed or spreading hairs. The alternate leaves are often

succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...

, only a few annual species have thin and flat leaves. The inconspicuous flowers sit solitary or in axillary clusters of 2–3 (5) in the axil of a subtending

bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or o ...

. They differ from the related subfamily

Salsoloideae The Salsoloideae are a subfamily of the Amaranthaceae, formerly in family Chenopodiaceae. Description These are herbs, subshrubs, shrubs and some trees. Stems and leaves are often succulent. The ovary contains a spiral embryo. In most genera ...

by the absence of bracteoles. The flowers are mostly bisexual. The

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla ( petals) or tepals when ...

consists of (3–) 5 membranous or scarious

tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...

s, which are often fused for about 1/5 to 4/5 of their length. 4–5

stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the fila ...

s are basally fused in a hypogynous disc. They have mostly exserted anthers without appendages. The

pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...

grains differ from Salsoloideae by greater diameter, and higher number of smaller pores with fewer spinulae per operculum. The horizontal or more rarely vertical ovary is uniovulate, with a distinct style and 2 filiform stigmas with papillae on the entire surface. The perianth persists end encloses the fruit. The tepals can enlarge or develop wings, spines or long hairs, or become fleshy or woody. The seed with thin testa contains an annular or folded embryo sometimes engirdling a rudimentary central perisperm.

Photosynthesis pathway

The species of ''Chenolea'' clade and the large ''Sclerolaena'' clade are plants. In the ''Bassia''/''Camphorosma'' clade, all species are plants except '' Sedobassia sedioides'' which is – intermediate.

Distribution and evolution

The Camphorosmeae are distributed in mainly in Australia (c. 147 species) and in the temperate and subtropical regions of the northern hemisphere: Eurasia including North Africa (c. 27 species), and North America (2 species), in South Africa (3 species). A few species are naturalized worldwide. They grow in different habitats as shores, salt marshes and deserts in mediterranean climate to forests, steppes and deserts in climates with summer rains, from the Sahara to the alpine zone in Central Asia. Very often they grow in dry, saline or disturbed ( ruderal) sites. The group evolved in the Early

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

, probably deriving from halophytic plants growing at seashores in a warm-temperate climate. The species of the ''Chenolea'' clade are regarded as remnants of an early line of evolution. The subfamily spread from Eurasia to Australia, North America and at least two times to South Africa. The Australian lineage diversified strongly, the other lineages remained species-poor.

Systematics

The taxon "Camphorosmeae" has been published in 1837 by Stephan Ladislaus Endlicher as a subtribe within the Chenopodieae.

Alfred Moquin-Tandon Christian Horace Benedict Alfred Moquin-Tandon (7 May 1804 – 15 April 1863) was a French naturalist and doctor. Moquin-Tandon was professor of zoology at Marseille from 1829 until 1833, when he was appointed professor of botany and director o ...

classified it as a tribe in 1840, and A J Scott raised it to subfamily level as "Camphorosmoideae" in 1978.

Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...

research by Kadereit & Freitag (2011) revealed that the traditional classification of the group did not reflect evolutionary relationships. Most of the genera, especially '' Kochia'' and '' Bassia'', were found to be highly

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...

, so some of their species had to be transferred to separate genera: '' Eokochia'', ''

Spirobassia ''Spirobassia'' is a monotypic plant genus with the species ''Spirobassia hirsuta''. It has cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the wor ...

'', '' Grubovia'' and '' Sedobassia''. The Australian species of Camphorosmeae form a relatively young group still in the process of speciation and with some hybridization between species. In phylogenetic research by Cabrera et al. (2009), the genera were not clearly separated. Probably ''

Neobassia ''Neobassia'' is a genus of small shrubs in the family Chenopodiaceae (''sensu stricto''), which are included in Amaranthaceae family, ('' sensu lato'') according to the APG classification. Species are endemic to Australia. Description Species ...

'', ''

Threlkeldia ''Threlkeldia '' is a genus of annuals or short-lived perennials in the family Amaranthaceae. There are two species, both of which are endemic to Australia. Together they occur in all Australian states except Queensland. *'' Threlkeldia incho ...

'' and ''

Osteocarpum ''Osteocarpum'' is a genus of flowering plants belonging to the family 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 g ...

'' should be included in ''

Sclerolaena ''Sclerolaena '' is a genus of annuals or short-lived perennials in the family Chenopodiaceae (''sensu stricto''), which are included in Amaranthaceae (''sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number ...

''. Likewise, '' Enchylaena'' should be included in '' Maireana''. The species-rich genera ''Sclerolaena'' and ''Maireana'' were found to be polyphyletic, so that further investigations are needed.

''Bassia''/''Camphorosma'' clade

plants (except ''Sedobassia sedoides'' which is intermediate). Widely distributed in Eurasia and southern Africa * '' Bassia'' All., (Syn. ''Kochia, Londesia, Panderia, Kirilowia, Chenoleioides''), with about 20 annual and subshrubby species, native from western Mediterranean to East Asia, introduced in America and northern Europa. * '' Camphorosma'' L., with 4 species, from western Mediterranean to Central Asia ** ''

Camphorosma annua ''Camphorosma'' is a genus of flowering plants in the family Amaranthaceae, found in northern Africa, southern and eastern Europe, Crimea, Russia, Anatolia, the Caucasus, Iran, Afghanistan, Pakistan, Central Asia, the Altai, western Siberia, Xinj ...

'' Pall., annual, from Hungary to eastern Ukraine ** '' Camphorosma lessingii'' Litv., subshrub, from Transcaucasia to South Siberia ** ''

Camphorosma monspeliaca ''Camphorosma'' is a genus of flowering plants in the family Amaranthaceae, found in northern Africa, southern and eastern Europe, Crimea, Russia, Anatolia, the Caucasus, Iran, Afghanistan, Pakistan, Central Asia, the Altai, western Siberia, Xinj ...

'' L., subshrub from West Europe to South Siberia ** ''

Camphorosma songorica ''Camphorosma'' is a genus of flowering plants in the family Amaranthaceae, found in northern Africa, southern and eastern Europe, Crimea, Russia, Anatolia, the Caucasus, Iran, Afghanistan, Pakistan, Central Asia, the Altai, western Siberia, Xinj ...

'' Bunge, annual, from lower Wolga to Central Asia * '' Sedobassia'' Freitag & G. Kadereit, with one species ** ''Sedobassia sedoides'' (Pall.) Freitag & G. Kadereit (Syn. ''Bassia sedoides'' (Pall.) Asch.), annual, from Hungary to Siberia

''Chenolea'' clade

plants. With 4 genera and 5 disjunct species. * '' Chenolea'' Thunb., with one species ** ''Chenolea diffusa'' Thunb., a subshrub in southern Africa * '' Eokochia'' Freitag & G. Kadereit, with one species ** ''Eokochia saxicola'' (Guss.) Freitag & G. Kadereit (Syn. ''Kochia saxicola'' Guss.), a subshrub, endemic on the Mediterranean islands Ischia, Capri and Stromboli * '' Neokochia'' (Ulbr.) G.L.Chu & S. C. Sand., with 2 species in southwestern North America **'' Neokochia americana'' (S. Watson) G.L.Chu & S .C. Sand. (Syn. ''Kochia americana'' S. Watson), a subshrub in southwestern North America **'' Neokochia californica'' (S. Watson) G.L.Chu & S. C. Sand., a subshrub in southwestern North America * ''

'' Freitag & G. Kadereit, with one species ** ''Spirobassia hirsuta'' (L.) Freitag & G. Kadereit (Syn. ''Bassia hirsuta'' (L.) Asch.): annual, from North Mediterranean to South Siberia.

''Sclerolaena'' clade

plants. *''Grubovia'' subclade, with 3 species in Central Asia: ** '' Grubovia'' Freitag & G. Kadereit, with 3 species in Central Asia *** '' Grubovia dasyphylla'' (Fisch. & C. A. Mey.) Freitag & G. Kadereit (Syn. ''Bassia dasyphylla'' (Fisch. & C. A. Mey.) Kuntze): annual, from eastern Kazakhstan to Mongolia. *** '' Grubovia krylovii'' (Litv.) Freitag & G. Kadereit (Syn. ''Kochia krylovii'' Litv.): annual, from the Altai mountains to Mongolia. *** '' Grubovia melanoptera'' (Bunge) Freitag & G. Kadereit (Syn. ''Kochia melanoptera'' Bunge): annual, form Tian Shan mountains to Mongolia. *''Sclerolaena'' subclade, with about 147 species in Australia: ** '' Didymanthus'' Endl., with only one species: *** ''Didymanthus roei'' Endl., in Australia ** ''

Dissocarpus ''Dissocarpus'' is a genus of flowering plants belonging to the family 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 goo ...

'' F. Muell.,with 4 species in Australia ** '' Enchylaena'' R.Br., with 2 species in Australia. This genus should be included in '' Maireana'' ** ''

Eremophea ''Eremophea'' is a genus of flowering plants belonging to the family 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 goose ...

'' Paul G.Wilson, with 2 species in Australia ** '' Eriochiton'' (R. H. Anderson) A. J. Scott, with only one species: *** ''Eriochiton sclerolaenoides'' (F. Muell.) F. Muell. ex A. J. Scott, in Australia ** '' Maireana'' Moq., with about 57 species in Australia. This genus is polyphyletic ** ''

Malacocera ''Malacocera'' is a genus of flowering plants belonging to the family 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 goos ...

'' R. H. Anderson, with 4 species in Australia ** ''

'' A. J. Scott, with 2 species in Australia. This genus should be included in ''

'' ** ''

'' F. Muell., with 5 species in Australia. This genus should be included in ''

'' ** '' Roycea'' C. A. Gardner, with 3 species in Australia ** ''

'' R. Br. (incl. ''Sclerochlamys'' F. Muell., ''Stelligera'' A. J. Scott), with 64 species in Australia. This genus is polyphyletic. ** ''

'' R. Br., with 2 species in Australia. This genus should be included in ''

Economic importance

Some species of Camphorosmeae are of limited economic interest. '' Bassia scoparia'' var. ''trichophylla'' is cultivated as an

ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that ...

("summer-cypress"). '' Bassia prostrata'' is increasingly important for the improvement of rangeland and phytoremediation. ''

Bassia indica ''Bassia'' is a genus of flowering plants in the family Amaranthaceae. They are distributed in the western Mediterranean to eastern Asia. Some occur outside their native ranges as introduced species.Bassia scoparia'' are used as

forage Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also use ...

plants. ''

'' is a traditional medicinal herb.

References

{{Taxonbar, from1=Q15301790, from2=Q147213 Amaranthaceae Caryophyllales tribes