Salsoloideae

The Salsoloideae are a subfamily of the Amaranthaceae, formerly in family Chenopodiaceae.

Salsoloideae
Salsola oppositifolia, in fruit
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Amaranthaceae
Subfamily: Salsoloideae
Raf.
Genera

About 35 genera, see text

Description

These are herbs, subshrubs, shrubs and some trees. Stems and leaves are often succulent. The ovary contains a spiral embryo. In most genera, scarious wings develop at the outside of the fruiting perianth, allowing for dispersal by the wind (anemochory). In tribe Caroxyleae, the stamens have vesiculose anther appendages, discolor with anthers, that probably play a role for insect pollination. In tribe Salsoleae the anther appendages are absent or small and inconspicuous.

Distribution

The area with most species (center of diversity) are the deserts and semideserts of Central-Asia and the Middle East. Distribution of the subfamily extends to the Mediterranean, to Middle-Europe, north and south Africa, and Australia, some species have also been introduced to America. Many species grow in dry habitats (xerophytes) or tolerate salty soils (halophytes), some are ruderals.

Photosynthesis pathway

Salsoloideae are C4 plants (with a few exceptions in tribe Salsoleae). Tribe Caroxyleae is exclusively of the NAD-malic enzyme C4 subtype. Most Salsoleae also use the NADP-malic enzyme.[1]

Taxonomy
Climacoptera turcomanica, in fruit
Cornulaca monacantha
Halothamnus subaphyllus, in fruit
Saxaul, Haloxylon ammodendron
Hammada griffithii, in fruit
Kali tragus

The type genus of subfamily Salsoloideae is Salsola L. According to recent research, the former classification did not reflect the phylogenetic relationship, so that the subfamily had to be reclassified; it is now split in two tribes, Caroxyleae and Salsoleae (in the strict sense).[2] The tribe Camphorosmeae is now treated in its own subfamily, Camphorosmoideae.[3]

Caroxyleae

Caroxyleae Akhani & E. H. Roalson (as "Caroxyloneae")[2]

  • Caroxylon Thunb., with 43 species
  • Climacoptera Botsch., with 41 species
  • Halarchon Bunge, with one species
    • Halarchon vesiculosum (Moq.) Bunge
  • Halimocnemis C. A. Mey., with 27 species (Syn. Gamanthus Bunge, Halanthium C. Koch, Halotis Bunge)
  • Halocharis Moq., with 7 species
  • Kaviria Akhani & E. H. Roalson, with 10 species
  • Nanophyton Less., with ca. 10 species
  • Ofaiston Raf., with one species
    • Ofaiston monandrum (Pall.) Moq.
  • Petrosimonia Bunge, with 12 species
  • Piptoptera Bunge, with one species
    • Piptoptera turkestana Bunge
  • Physandra Botsch., with one species
    • Physandra halimocnemis (Botsch.) Botsch.
  • Pyankovia Akhani & E. H. Roalson, with one species
    • Pyankovia brachiata (Pall.) Akhani & E. H. Roalson

Salsoleae
  • Anabasis L. ( incl. Fredolia (Coss. & Durieu ex Bunge) Ulbr.), with 29 species
  • Arthrophytum Schrenk, with 9 species
  • Cornulaca Delile, with 5 species
  • Cyathobasis Aellen, with one species:
    • Cyathobasis fruticulosa (Bunge) Aellen
  • Girgensohnia Bunge ex Fenzl, with 4 species
  • Halogeton C. A. Mey, with 5 species. (Syn. Agathophora (Fenzl) Bunge, Micropeplis Bunge)
  • Halothamnus Jaub. & Spach, with 21 species
  • Haloxylon Bunge, with 2 species
  • Hammada Iljin, with 12 species
  • Horaninowia Fisch. & C. A. Mey, with 6 species
  • Kali Mill., with 13 species:
  • Lagenantha Chiov. (Syn.: Gyroptera Botsch.) Classification not sure. With 1-3 species.
  • Noaea Moq., with 3 species
  • Nucularia Batt., Classification not sure. With one species:
    • Nucularia perrini Batt.
  • Rhaphidophyton Iljin, with one species
    • Rhaphidophyton regelii (Bunge) Iljin
  • Salsola L., with 25 species. (Syn. Darneilla Maire & Weiller, Fadenia Aellen & Townsend, Neocaspia Tzvelev, Hypocylix Wol., Seidlitzia Bunge ex Boiss.)
  • Sympegma Bunge
  • Traganum Del., with 2 species
  • Traganopsis Maire et Wilczek, with one species
    • Traganopsis glomerata Maire & Wilczek
  • Turania Akhani & E. H. Roalson, with 4 species
  • Xylosalsola Tzvelev, with 4 species
  • Classification within Salsoleae unclear:
    • "Canarosalsola"-Clade:
      • Salsola divaricata Masson ex Link.
    • "Collinosalsola"-Clade:
      • Salsola arbusculiformis Drob.
      • Salsola laricifolia Turcz. ex Litw.
    • "Oreosalsola"-Clade:
      • Salsola abrotanoides Bunge
      • Salsola botschantzevii Kurbanov
      • Salsola flexuosa Botsch.
      • Salsola junatovii Botsch.
      • Salsola lipschitzii Botsch.
      • Salsola maracandica Iljin
      • Salsola masenderanica Botsch.
      • Salsola montana Litw.
      • Salsola oreophila Botsch.
      • Salsola tianschanica Botsch.
    • Others:
      • Salsola genistoides Juss. ex Poir.
      • Salsola pachyphylla Botsch.
      • Salsola webbii Moq.

Classification in subfamily not sure
  • Iljinia Korovin, with one species:
    • Iljinia regelii (Bunge) Korovin

References
  1. Pyankov, V.; Ziegler, H.; Kuz'min, A.; Edwards, G. (2001). "Origin and evolution of C4 photosynthesis in the tribe Salsoleae (Chenopodiaceae) based on anatomical and biochemical types in leaves and cotyledons". Plant Systematics and Evolution. 230 (1–2): 43–74. doi:10.1007/s006060170004. ISSN 0378-2697. S2CID 25370346.
  2. Akhani, H.; Edwards, G.; Roalson, E.H. (2007). "Diversification of the Old World Salsoleae s.l. (Chenopodiaceae): Molecular phylogenetic analysis of nuclear and chloroplast data sets and a revised classification". International Journal of Plant Sciences. 168 (6): 931–956. doi:10.1086/518263. ISSN 1058-5893.
  3. Kadereit, G.; Freitag, H. (2011). "Molecular phylogeny of Camphorosmeae (Camphorosmoideae, Chenopodiaceae): Implications for biogeography, evolution of C4-photosynthesis and taxonomy". Taxon. 60 (1): 51–78. doi:10.1002/tax.601006.
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