''Halothamnus'' is a genus in the subfamily
Salsoloideae 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
annuals. 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
bracteoles. 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 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''

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). 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), 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 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 in
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

The genus ''Halothamnus'' belongs to the subfamily
Salsoloideae within 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 ...
, (formerly
Chenopodiaceae). It is grouped to Tribus
Salsoleae, Subtribus
Sodinae, and is closely related to genus ''
Salsola''.
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 included the genus ''Aellenia'' here and enlarged ''Halothamnus'' to 25 species. After a revision by
Gabriele Kothe-Heinrich (1993), the genus is classified into 2 sections with 21 accepted species:
*Sect. ''Pungentifolia'' Kothe-Heinr.
**''
Halothamnus beckettii '' Botsch.
*Sect. ''Halothamnus''
**''
Halothamnus somalensis '' (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 '' (Bornm.) Botsch.
**''
Halothamnus iliensis '' (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 '' (Gilli) Botsch.
**''
Halothamnus schurobi '' (Botsch.)
**''
Halothamnus turcomanicus '' Botsch.
**''
Halothamnus ferganensis '' Botsch.
**''
Halothamnus sistanicus '' (De Marco & Dinelli) Kothe-Heinr.
**''
Halothamnus oxianus '' Botsch.
**''
Halothamnus seravschanicus '' Botsch.
**''
Halothamnus iraqensis '' Botsch.
**''
Halothamnus subaphyllus '' (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 WebpageDigital herbarium specimens from Herbarium Berlin
*
''Halothamnus'' Jaub. & Spachat Tropicos
{{Taxonbar, from=Q149213
Halothamnus,
Amaranthaceae genera