''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, maki ...

(s.l., now including Chenopodiaceae). The scientific name means "saltbush", from the

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

() and () . This refers either to salty habitats or to the accumulation of salt in the plants. The genus is distributed from

Southwest The points of the compass are a set of horizontal, radially arrayed compass directions (or azimuths) used in navigation and cartography. A '' compass rose'' is primarily composed of four cardinal directions—north, east, south, and west— ...

and

Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...

to the

Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...

and

East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...

Description

Most species of ''Halothamnus'' are small

shrub A shrub or 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 by their multiple ...

s or

sub-shrub A subshrub (Latin ''suffrutex'') or undershrub is either a small shrub (e.g. prostrate shrubs) or a perennial that is largely herbaceous but slightly woody at the base (e.g. garden pink and florist's chrysanthemum). The term is often intercha ...

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 (band), a ...

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, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...

and two green

bracteole In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...

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 ve ...

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

stamens The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filamen ...

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

ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...

is formed by two

carpels Gynoecium (; ; : gynoecia) 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) ...

, with a thick

pistil Gynoecium (; ; : gynoecia) 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 (botany), whorl of a flower; it consists ...

, 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. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...

. 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 most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

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 one of the two types of hermaphroditism, the other type being simultaneous hermaphroditism. It occurs when the organism's sex changes at some point in its life. A sequential hermaphrodit ...

). ''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 * * Azerbaijani: , * * * * * * * * * * * is a mountain range at the intersection of Asia and Europe. Stretching between the Black Sea and the Caspian Sea, they are surrounded by the Caucasus region ...

and

Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...

through

Southwest Asia West Asia (also called Western Asia or Southwest Asia) is the westernmost region of Asia. As defined by most academics, UN bodies and other institutions, the subregion consists of Anatolia, the Arabian Peninsula, Iran, Mesopotamia, the Armenia ...

southward to

Somalia Somalia, officially the Federal Republic of Somalia, is the easternmost country in continental Africa. The country is located in the Horn of Africa and is bordered by Ethiopia to the west, Djibouti to the northwest, Kenya to the southwest, th ...

. The westernmost localities are at the

Sinai Peninsula The Sinai Peninsula, or simply Sinai ( ; ; ; ), 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 land bridge between Asia and Afri ...

, the easternmost in

Dzungaria Dzungaria (; from the Mongolian words , meaning 'left hand'), also known as Northern Xinjiang or Beijiang, is a geographical subregion in Northwest China that corresponds to the northern half of Xinjiang. Bound by the Altai Mountains to the n ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

. Most species occur in Iran,

Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan, is a landlocked country located at the crossroads of Central Asia and South Asia. It is bordered by Pakistan to the Durand Line, east and south, Iran to the Afghanistan–Iran borde ...

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

desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

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 sem ...

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 broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...

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 (biology), type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 ...

). 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 Australia, central and southwestern Asia, North Africa, and the Mediterranean. Common names of various members of this ...

''. 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 Nile ...

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). H ...

(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 '' 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 '' (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 ''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 ''Halothamn ...

'' (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 {{Authority control Halothamnus, Amaranthaceae genera