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Pakistan | Family List | Pakistan V. 204 | Chenopodiaceae

23. Salsola Linnaeus, Sp. Pl. 222. 1753; Gen. pl. ed. 5: 104. 1754; Moq., Chenop. monogr. enum. 134. 1840; Moq. in DC., Prodr. 13,2: 179. 1849; Boiss., Fl. Or. 4: 951. 1878; Ulbr. in Engl. & Prantl, Nat. Pflanzenfam. ed.2, 16c: 563. 1934; Iljin in Fl. Gen. Flow. Pl. 2: 275.1993; Freitag in Rech.f., Fl. Iran. 172: 154. 1997; Rilke in Bibl. Bot. 149: 51. 1999. SSSR 6: 200. 1936; Aellen in Hegi, Illustr. Fl. Mittel-Eur. ed. 2, 3,2: 739. 1968; Botsch. in Bot. Zhurn. SSSR 54: 989. 1969 (and numerous further works); 1968; Kuhn in Kubitzki, Fam.

Caroxylon Thunb., Nov. gen.. 2: 37. 1782; Salsolaria Moq., Chenop. Monogr. enum. 144. 1840 p.p.; Hypocylyx Wol. in Stapf, Denkschr. Akad. Wiss. Wien, Math.-Nat. Kl. 51: 275. 1886; Darniella Maire & Weiller in Bull. Soc. Hist. Nat. Afrique N. 30: 301. 1939; Climacoptera Botsch. in Sborn. Posv. Akad. Sukachevu: 111. 1956; Nitrosalsola Tzvelev in Ukrayins´k. Bot. Zhurn. 50: 80. 1993; Tzvelev in Fl. Eur. Or. 9: 87. 1996; Xylosalsola Tzvelev l.c. Ukrayins´k. Bot. Zhurn. 50: 81. 1993; Tzvelev in Fl. Eur. Or. 9: 90. 1996.

Annuals, dwarf shrubs, shrubs rarely small trees. Glabrous or with very diverse types of indumentum. Young stems whitish, purplish or fresh green with pale longitudinal lines. Leaves alternate, sometimes subopposite or opposite, rarely distichous; usually semiterete or terete and ± succulent, more or less linear, eventually with a spiny apex, isolateral, mostly with 1 large central bundle and numerous smaller peripheral bundles near to the Kranz layer, usually short-lived and functionally replaced in flowering time by bracts and bracteoles; leaf base very variable in shape. Bracts more or less leaf-like and succesively shorter. Bracteoles 2, green at least in upper part, usually much shorter and more uniform in length, enclosing at least the lower 2/3 of the flowers. Flowers sessile, solitary or clustered, in simple or paniculate spikes; usually bisexual, protandrous, anemophilous or/and entomophilous. Tepals 5, free from base, 1-4 mm long, varying in outline from circular to narrow ovate and from very thin and delicate to moderately thickened or stiff and coriaceous, with or without a distinct green blotch on back, the outer wider than the inner, with 1 to several veines, in lower 1/2 usually with a distinct transverse line associated with the later formation of a wing-like outgrowth. Stamens 5, antitepalous, anthers with thecae more or less deeply divided from base, apically with an appendage of very different size and shape; filaments band-shaped, mostly inserted on the outside of a glandular, lobed interstaminal disc, more rarely disc absent. Ovary globular, ovoid or depressed, style shorter or longer, rarely absent; stigmas 2, rarely 3, spreading, recurved or revolute, flat, papillose on upper side only (in our species); ovule 1, basal. Fruit usually made up of an utricle with adnate fruiting perianth, both shed together as an anemophilous diaspore; utricle commonly enclosed, sometimes exposed on upper side. Tepals in fruit with 5 subequal wings or the 2 inner much smaller, wings horizontally spreading, flat or somewhat flexuose, often partly overlapping, very thin and translucent with delicate radial veins, or thicker, coriaceous, and then pale or brown, when young and fresh often purple, pinkish, orange or yellow; tepal back above the wings often variously enlarged and thickened, usually connivent, the lobe apices often forming very typical dome-, cone- or column-like structures; tepals below the wings slightly to moderately hardened, rarely indurated, forming a bowl- or cup-like structure, its base usually flattened, with 5 shallow grooves and a polygonal rim surrounding the central scar, more rarely the base rounded; sometimes 1 or 2-3 fruits on condensed lateral spikes forming wingless complex fruits by their greatly enlarged, fused and indurated bracts and bracteoles. Utricle drum-, disc-or pot-shaped, or semiglobular with the upper side flattened; pericarp membranous. Seed horizontal, more rarely oblique or vertical, with membranous testa; endosperm almost absent; embryo flat to conical spiral or helical, greenish.

A genus of about 100 species centred in the Old World desert belt from the Canary Islands through S Europe and N Africa to SW and C Asia, southwards to tropical E Africa and to SW Africa; a few species along seashores, on ruderal and naturally disturbed sites penetrating into adjacent semi-humid or even humid areas; 3 species of sect. Kali adventive in semi-arid parts of N and S America, E and S Africa, and in Australia.

18 species are recorded from Pakistan. 4 more species are included which do occur in adjacent parts of SE Afghanistan and very likely will be found sooner or later in the geologically, altitudinally and climatologically similar parts on this side of the border.

*H.Freitag, Morphologie u. Systematik der Pflanzen, Fachbereich Biologie/ Chemie, Universität Kassel, Heinrich Plett-Str. 40, D-34132 Kassel, Germany.

Species erroneously recorded from the area:

S. dendroides Pall. in R. R. Stewart, Ann. Cat. Vasc. Pl. Pak. Kashm. 226. 1972 – a perennial herb of sect. Caroxylon, extending eastwards to C and NE Afghanistan only; probably confused with S. imbricata Forssk.

S. lanata Pall. in Burkill, Work. List Fl. Pl. Baluch. 65. 1909 and R. R. Stewart, Ann. Cat. Vasc. Pl. Pak. Kashm. 226. 1972 – annual species of sect. Physurus from the Irano-Turanian; related to and mistaken for S. crassa.

S. vermiculata L. in Parker, Forest Fl. Punjab ed. 2: 418. 1924 – a Mediterranean species of sect. Caroxylon, extending eastwards to W. Iran only; confused with S. imbricata Forssk.

S. verrucosa M. Bieb. in Burkill, Work. List Fl. Pl. Baluch. 66. 1909 – a dwarf shrub of sect. Caroxylon restricted to an area from NE Anatolia, Armenia, Azerbaijab and NW Iran, probably confused with S. imbricata Forssk.

1 Stems of annual shoots fresh green except for narrow longitudinal lines. Leaves, bracts and bracteoles spine-tipped, often pungent (fig.25,A) sect. Kali.   (10)
+ Stems of annual shoots nor fresh green, eventually a short green area just below the base of leaves and bracts present. Leaves, bracts and bracteoles obtuse, acute or apiculate, never spiny nor pungent.   (2)
2 (1) Stems of annual shoots with a short green area just below upper leaves and bracts (fig.25, G). Anther appendages large, vesicular, stalked (fig. 26,1).sect. Physurus.   22a subsp. turcomanica
+ Stems uniformly whitish, creamy or reddish. Anther appendages small or large, never stalked.   (3)
3 (2) Annuals. Grey with a dense indumentum of short, thiln-walled, vesicular or saussage-shaped hairs (surface mealy). sect. Cardiandra.   (18)
+ Perennials or annuals. Variously hairy but never with vesicular or saussage-shaped hairs.   (4)
4 (3) Plants glabroius, papillose, hispidulous or hispid by stiff, thick-walled, 1-celled hairs.   (5)
+ Plants throughout or at least the young stems with long or short few-celled hairs of various shape.   (6)
5 (4) Leaf blade arising from a compact, indurated and persistent pulvinus (fig. 25, C,D). Flowers in clusters. Perianth lobes not thickened.(fig.29,A). sect. Cocco-salsola.   (7)
+ Leaf blade near base petiole-like constricted, arising from a dilated, hump-shaped, persistent base (fig.25,B). Flowers solitary. Lobes of the fruiting perianth above the wings ± thickened (fig.28, A-C). Utricle enclosed by the fruiting perianth. sect. Arbuscula.   (8)
6 (4) Upper leaves and bracts beneath near base distinctly gibbous (fig. 25, E). Tepal back with green blotch. Antgher appendages very short. sect. Caroxylon.   (14)
+ Leaves and bracts at base never gibbous, the upper ± sheathing (fig. 25, F). Tepal back without green blotch. Anther appendages large, at least 1/3 of thecae lengh (fig. 26, C,D). sect. Belanthera.   (20)
7 (5) Small shrub up to 1 m with divaricate branching. Upper leaves and bracts from distinctly clavatge to dumb-bell-shaped (fig. 25, D; fig. 29,A). sect. Coccosalsola.   1 S. drummondii
+ Subshrub up to 0.4 m. All leaves and bracts cylindrical or semi-cylindrical (fig. 25, C). C. Makran only.   2 S. makranica
8 (5) Loosely branched elegant shrub up to 2 m. Lower leaves 3-8 cm long, often pendant. Fruiting perianth 14-18 mm diam., with a stiff cone formed by the apical tepal lobes (fig. 28, C). Baluchistan   5 S. richteri
+ Densely branched smaller plants. Leaves shorter, never pendant. Fruiting perianth smaller, with different stgructgures formed by the tepal lobes.   (9)
9 (8) Small shrub up to 1 m, the branches divaricate, rigid and often becoming spiny. Fruiting perianth 1013 mm diam., with the lobes above the wings forming a thick, indurated, circular bulge crowned by a flower-like cup (fig. 28, B). To expect in Waziristan and Upper Baluchistan. sect. Arbuscula.   4 S. arbuscula
+ Subshrub up to 30 cm, with numerous erect annual shoots, the woody branches not becoming spiny. Fruiting perianth 7-10 mm diam., the lobes forming a narrow bulge and continuing into a flat dome-like structure (fig. 28, A). Waziristan, Upper Baluchistan.   3 S. montana
10 (1) Small shrub or subshrub. Axillary hairs in dense, wool-like tufts. Anthers 1.8-2.8 mm long. Baluchistan.   10 S. griffithii
+ Annuals. Axillary hairs less prominent, fugacious. Anthers shorter.   (11)
11 (10) Fruits without wings, up to 2 mm wide. Spikes elongated, dense, narrow, with imbricate bracts and bracteoles. Outer tgepals 1-veined. Seed vertical or oblique (fig. 27, B1). N.W.F.P., Kashmir.   7 S. collina
+ Normal fruits winged, at least 5 mm wide. Spikes with spreading bracts and bracteoles. Outer tepals 3-veined. sect. Kali.   (12)
12 (11) Fruiting perianth above the wilngs with 5 parallel delicate spine-like indurated tepal lobes (fig. 27, D). Baluchistan.   9 S. paulsenii
+ Fruiting perianth above the wings with 0-2(3) indurated tepal lobes.   (13)
13 (12) Plant robust, usually glabrous. Tepals 2-2.5 mm long, glabrous. Fruits 6-8 mm diam. (fig. 27, A1).   6 S. tragus
+ Plant delicate, always hispid-puberulent. Tepals 1.5-2 mm long, densely papillose on back and long ciliate at the margins. Fruits 4-7 mm diam. (fig. 27, C).N.W.F.P., Kashmir.   8 S. jacquemontii
14 (6) Annuals.   (15)
+ Subshrubs or shrubs.   (16)
15 (14) Tepal back with stiff, ascending hairs, in fruit thickened and the lobes hook-like incurved (fig. 28, H). Upper bracts and bracteoles with persistent hairs. sect. Caroxylon. Expected for Waziristan and Upper Baluchistan.   13 S. incanescens
+ Tepals back glabrous, in fruit not significantly thickened, the lobes not hook-like incurved (fig. 28, G). Upper bracts and bractgeoles glabrous or glabrescent.   12 S. nitraria
16 (14) Subshrub with divaricate, spinescent branches. Leaves, bracts, bracteoles and tepals white with adpressed, smooth hairs (fig. 28, D). Disc very dilstinct. Makran, Lower Baluchistan.   15 S. cyclophylla
+ Subshrubs or shrubs without divaricate, spinescent branches. Indument not adpressed, hairs warty or denticulate.   (17)
17 (16) Leaves and lower bracts all or predominantly linear. Fruiting perianth 6-9 mm diam. Hairs branched. Expected for Waziristan and Upper Baluchistan.   14 S. orientalis
+ All bracts scale-like and leaves predominantly so. Fruiting perianth up to 6 mm diam. Hiars spinulose or almost smooth. sect. Caroxylon.   11 S. imbricata
18 (3) Plant procumbent, with divaricate, rigid branches. Long hairs stiff, rough from rounded warts. Fruiting perianth 4-6 mm diam. Baluchistan.   16 S. chorassanica
+ Plants ±erect, with slender, ascending branches. Long hairs delicate, smooth or denticulate, interwoven, forming an arachnoid indumentum (young plants!).   (19)
19 (18) All leaves needle-like, the lower 0.5-0.75 mm wide. Tepals 1.5-2.5 mm long. Anthers 0.9-1.4 mm long. Stigmas 0.7-1 mm long. Wings of fruiting perianth with a prominent keel on upper side (fig. 29, C). Baluchistan, Waziristan.   17 S. turkestanica
+ Lower leaves flat, 0.72-1.2 mm wide. Tepals 3-3.5 mm long. Anthers 1.3-1.8 mm long. Stigmas 1-1.5 mm long. Wings of fruiting perianth not or only the inner indistinctly keeled (fig. 29, D). Psammophyte. Baluchistan.   18 S. sclerantha
20 (6) Leaves flat, not or only slightly succulent, covered by a persistent silky indument of smooth hairs. Spikes few, very long, thin, strictly erect. Tepals 3-7-veined. Expected for Waziristan.   19 S. canescens
+ Leaves semi-terete, strongly succulent, with scattered spinulose hairs, glabrescent. Spikes numerous by profuse branching, short, spreading. Tepals 2-3-veined.   (21)
21 (20) All leaves and bracts helically arranged, the upper not clasping at base. Chagai; expected for Upper Baluchistan and Waziristan.   20 S. tomentosa
+ Lateral vegetative shoots and spikes conspicuously flattened by distichous position of half-clasping leaves and bracts. C. Makran.   21 S. rubescens

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