''Halothamnus'' is a genus in the 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, ...

of 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 goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making it ...

(s.l., now including Chenopodiaceae). The scientific name means saltbush, from the Greek ἅλς (''hals'') "salt" and θαμνος (''thamnos'') "bush". This refers either to salty habitats or to the accumulation of salt in the plants. The genus is distributed from Southwest and Central Asia to the Arabian peninsula and East Africa.

Description

Most species of ''Halothamnus'' are small

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

s or

sub-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 De ...

s, two species are

annual Annual may refer to: * Annual publication, periodical publications appearing regularly once per year **Yearbook ** Literary annual * Annual plant * Annual report * Annual giving * Annual, Morocco, a settlement in northeastern Morocco * Annuals (b ...

s. The alternate leaves are sitting without basal narrowing at the branches. They are simple, entire, half-terete or flat and slightly fleshy (

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 ...

). The bisexual inconspicuous flowers are sitting solitary in the axil of a green

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 of ...

and two green

bracteole 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 of ...

s. The five free

tepals 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 ...

are green with membranous margins above a transverse line and colourless below. Five

stamens 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 filame ...

arise from a cup-like structure (hypogynous disc) at the base of the flower. One

ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...

is formed by two

carpels Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

, with a thick

pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

, and two stigmas. The one-seeded, horizontally flattened fruit keeps being enclosed by the hardening

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 ...

. Wings develop from the transverse zone at the outer side of the tepals. Below the wings, the perianth forms a thick and lignified tube with a widened base. Its bottom side has five pits arranged in a ring, surrounded by a rim. These fruit characteristics are essential for the genus ''Halothamnus''. file:Halothamnus_lancifolius_fruit.JPG, Fruit (lateral view) of ''Halothamnus lancifolius'' file:Halothamnus_iranicus_fruit.JPG, Fruit (bottom) of ''Halothamnus iranicus'' Halothamnus Pollen 1

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 gametophyt ...

grains are nearly ball-shaped, 18-38 µm in diameter, with 12-29 pores scattered all over the surface (pantoporate), just like it is typically for the goosefoot-family. The species differ from each other in pollen diameter and number of pores, the southern species with the smallest and the northern species with the largest pollen grains.

Pollination and dissemination

In all species the stamens blossom first, and the stigmas unfold later (

protandry Sequential hermaphroditism (called dichogamy in botany) is a type of hermaphroditism that occurs in many fish, gastropods, and plants. Sequential hermaphroditism occurs when the individual changes its sex at some point in its life. In particular, ...

). ''Halothamnus subaphyllus'' was proved to be pollinated by insects (

entomophily Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, some ...

). There are no observations for the other species, but they are considered to be pollinated by insects, too. The fruits enclosed by the hardened perianth are dispersed by the wind (

anemochory In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...

), lifted up by their wings up to 20 mm in diameter.

Distribution

The distribution of the genus extends from the

Caucasus mountains The Caucasus Mountains, : pronounced * hy, Կովկասյան լեռներ, : pronounced * az, Qafqaz dağları, pronounced * rus, Кавка́зские го́ры, Kavkázskiye góry, kɐfˈkasːkʲɪje ˈɡorɨ * tr, Kafkas Dağla ...

and

Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...

through

Central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...

Southwest Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion of the larger geographical region of Asia, as defined by some academics, UN bodies and other institutions. It is almost entirely a part of the Middle East, and includes Anat ...

southward to

Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constituti ...

. The westernmost localities are at the

Sinai Peninsula The Sinai Peninsula, or simply Sinai (now usually ) (, , cop, Ⲥⲓⲛⲁ), is a peninsula in Egypt, and the only part of the country located in Asia. It is between the Mediterranean Sea to the north and the Red Sea to the south, and is a l ...

, the easternmost in

Dzungaria Dzungaria (; from the Mongolian language, Mongolian words , meaning 'left hand') is a geographical subregion in Northwest China that corresponds to the northern half of Xinjiang. It is thus also known as Beijiang, which means "Northern Xinjiang" ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...

. Most species occur in Iran,

Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan,; prs, امارت اسلامی افغانستان is a landlocked country located at the crossroads of Central Asia and South Asia. Referred to as the Heart of Asia, it is bordere ...

, and southern Central Asia. All species grow in dry habitats of

desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...

s and

semidesert A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi-ar ...

s from 0–2800 m above sea-level. They live on rocky, clayish, or sandy soils; many species tolerate saline conditions.

Photosynthesis pathway

All species of ''Halothamnus'' show leaf anatomy with Kranz arrangement of the ''Salsola''-type. Physiological investigations confirmed that they are all C4-plants.

Economic importance

Some species of ''Halothamnus'' are important grazing plants for camels, sheep and goats,G. K. Kinzikaeva: ''Aellenia''. - In: P. N. Občinnikov (Hrsg.): ''Flora Tadžikskoj SSR'' 3: p. 419-425. - Leningrad: Nauka, 1968 and are therefore used for the recultivation of pasture, for example ''Halothamnus subaphyllus'', ''Halothamnus auriculus'' and ''Halothamnus glaucus''. ''Halothamnus subaphyllus'' is planted for the stabilisation of mobile sands and for the production of the medically used

alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...

Salsolin. ''Halothamnus somalensis'' is used as medicinal plant against parasitic worms. In former times ''Halothamnus subaphyllus'' and ''Halothamnus glaucus'' were used to extract

potash Potash () includes various mined and manufactured salts that contain potassium in water-soluble form.

for making soap.P. Aellen: ''Chenopodiaceae''. - In: G. Hegi (Hrsg.: K. H. Rechinger): ''Illustrierte Flora von Mitteleuropa'', 2. Aufl. 3(2): p. 533-747. - Berlin/Hamburg: Parey, 1961.

Systematics

Halothamnus glaucus as Kali fruticosum spicatum by Buxbaum

The genus ''Halothamnus'' belongs to the subfamily

within the family

, (formerly

Chenopodiaceae 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 ...

). It is grouped to Tribus

Salsoleae 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, ...

, Subtribus Sodinae, and is closely related to genus ''

Salsola ''Salsola'' is a genus of the subfamily Salsoloideae in the family Amaranthaceae. The genus ''sensu stricto'' is distributed in central and southwestern Asia, North Africa, and the Mediterranean. A common name of various members of this genus and ...

''. The genus ''Halothamnus'' was first described in 1845 by

Hippolyte François Jaubert Count Hippolyte François Jaubert (28 October 1798 – 5 December 1874) was a French politician and botanist. Jaubert was born in Paris, the son of François Hippolyte Jaubert (a commissioner of the French Navy, killed at the Battle of the Nil ...

and

Édouard Spach Édouard Spach (23 November 1801 – 18 May 1879) was a French botanist. The son of a merchant in Strasbourg, in 1824 he went to Paris, where he studied botany with René Desfontaines (1750–1831) and Antoine-Laurent de Jussieu (1748–1836). ...

(in ''Illustrationes Plantarum Orientalium'', 2, 50, pl. 136). The type of the genus is ''Halothamnus bottae'', at that time the only species. In 1981,

Victor Petrovič Botschantzev Victor Petrovitch Botchantsev (russian: Виктор Петрович Бочанцев) (20 October 1910 - 30 August 1990) was a Russian botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of ...

included the genus ''Aellenia'' here and enlarged ''Halothamnus'' to 25 species. After a revision by

Gabriele Kothe-Heinrich Gabriele is both a given name and a surname. Notable people with the name include: Given name Surname *Al Gabriele, American comic book artist * Angel Gabriele (1956–2016), American comic book artist *Corrado Gabriele (born 1966), Italian polit ...

(1993), the genus is classified into 2 sections with 21 accepted species: *Sect. ''Pungentifolia'' Kothe-Heinr. **'' Halothamnus beckettii '' Botsch. *Sect. ''Halothamnus'' **''

Halothamnus somalensis ''Halothamnus somalensis'' is a species of the plant genus ''Halothamnus'', that is now included into the family Amaranthaceae, (formerly Chenopodiaceae). It is endemic to the Horn of Africa (Djibouti, Somalia, and Ethiopia) and is used for tradi ...

'' (N.E.Br.) Botsch. **''

Halothamnus bottae ''Halothamnus bottae'' is a species of the plant genus ''Halothamnus'', that belongs to the subfamily Salsoloideae of the family Amaranthaceae. It occurs on the Arabian peninsula. Morphology ''Halothamnus bottae'' is a shrub 30–50 cm h ...

'' Jaub. & Spach - with 2 subspecies **'' Halothamnus iranicus '' Botsch. **''

Halothamnus hierochunticus ''Halothamnus hierochunticus'' is a species of the plant genus '' Halothamnus'', that belongs to the subfamily Salsoloideae within the family Amaranthaceae, (formerly Chenopodiaceae). It occurs in Southwest Asia and is partly considered as a weed ...

'' (Bornm.) Botsch. **''

Halothamnus iliensis ''Halothamnus iliensis'' is a species of the plant genus ''Halothamnus'', that belongs to the subfamily Salsoloideae of the family Amaranthaceae, (formerly Chenopodiaceae). It occurs in Central Asia. Morphology ''Halothamnus iliensis'' is an an ...

'' (Lipsky) Botsch. **'' Halothamnus auriculus '' (Moq.) Botsch. - with 2 subspecies **'' Halothamnus kermanensis '' Kothe-Heinr. **'' Halothamnus afghanicus '' Kothe-Heinr. **''

Halothamnus lancifolius ''Halothamnus lancifolius'' is a species of the plant genus ''Halothamnus'', that belongs to the subfamily Salsoloideae within the family Amaranthaceae, (formerly Chenopodiaceae). It occurs in Southwest Asia. Morphology ''Halothamnus lancifoli ...

'' (Boiss.) Kothe-Heinr. **'' Halothamnus cinerascens '' (Moq.) Kothe-Heinr. - with 2 subspecies **'' Halothamnus glaucus '' (M.Bieb.) Botsch. - with 3 subspecies **''

Halothamnus bamianicus ''Halothamnus'' is a genus in the subfamily Salsoloideae of the family Amaranthaceae (s.l., now including Chenopodiaceae). The scientific name means saltbush, from the Greek ἅλς (''hals'') "salt" and θαμνος (''thamnos'') "bush". This re ...

'' (Gilli) Botsch. **'' Halothamnus schurobi '' (Botsch.) **'' Halothamnus turcomanicus '' Botsch. **'' Halothamnus ferganensis '' Botsch. **''

Halothamnus sistanicus ''Halothamnus'' is a genus in the subfamily Salsoloideae of the family Amaranthaceae (s.l., now including Chenopodiaceae). The scientific name means saltbush, from the Greek ἅλς (''hals'') "salt" and θαμνος (''thamnos'') "bush". This re ...

'' (De Marco & Dinelli) Kothe-Heinr. **''

Halothamnus oxianus ''Halothamnus'' is a genus in the subfamily Salsoloideae of the family Amaranthaceae (s.l., now including Chenopodiaceae). The scientific name means saltbush, from the Greek ἅλς (''hals'') "salt" and θαμνος (''thamnos'') "bush". This re ...

'' Botsch. **''

Halothamnus seravschanicus ''Halothamnus'' is a genus in the subfamily Salsoloideae of the family Amaranthaceae (s.l., now including Chenopodiaceae). The scientific name means saltbush, from the Greek ἅλς (''hals'') "salt" and θαμνος (''thamnos'') "bush". This re ...

'' Botsch. **'' Halothamnus iraqensis '' Botsch. **''

Halothamnus subaphyllus ''Halothamnus subaphyllus'' is a species of the plant genus '' Halothamnus'', that belongs to the subfamily Salsoloideae within the family Amaranthaceae, (formerly Chenopodiaceae). It occurs in Southwest and Central Asia. Morphology ''Halotha ...

'' (C.Meyer) Botsch. - with 3 subspecies

References

* Gabriele Kothe-Heinrich: ''Revision der Gattung ''Halothamnus'' (Chenopodiaceae)''. Bibliotheca Botanica Bd. 143, Schweizerbart, Stuttgart 1993, (German) * Gabriele Kothe-Heinrich: ''Halothamnus''. In: Karl Heinz Rechinger et al. (Edit.): ''Flora Iranica'' 172, ''Chenopodiaceae'': 255-289 - Graz: Akad. Druck 1997,

External links

Schweizerbart Bibliotheca Botanica Bd. 143 Webpage

Digital herbarium specimens from Herbarium Berlin *
''Halothamnus'' Jaub. & Spach
at Tropicos {{Taxonbar, from=Q149213 Halothamnus, Amaranthaceae genera