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Chenopodiaceae Ventenat

藜科 li ke

Authors: Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants

Herbs annual, subshrubs, or shrubs, rarely perennial herbs or small trees. Stems and branches sometimes jointed (articulate); indumentum of vesicular hairs (furfuraceous or farinose), ramified (dendroid), stellate, rarely of glandular hairs, or plants glabrous. Leaves alternate or opposite, exstipulate, petiolate or sessile; leaf blade flattened, terete, semiterete, or in some species reduced to scales. Flowers monochlamydeous, bisexual or unisexual (plants monoecious or dioecious, rarely polygamous); bracteate or ebracteate. Bractlets (if present) 1 or 2, lanceolate, navicular, or scale-like. Perianth membranous, herbaceous, or succulent, (1-)3-5-parted; segments imbricate, rarely in 2 series, often enlarged and hardened in fruit, or with winged, acicular, or tuberculate appendages abaxially, seldom unmodified (in tribe Atripliceae female flowers without or with poorly developed perianth borne between 2 specialized bracts or at base of a bract). Stamens shorter than or equaling perianth segments and arranged opposite them; filaments subulate or linear, united at base and usually forming a hypogynous disk, sometimes with interstaminal lobes; anthers dorsifixed, incumbent in bud, 2-locular, extrorse, or dehiscent by lateral, longitudinal slits, obtuse or appendaged at apex. Ovary superior, ovoid or globose, of 2-5 carpels, unilocular; ovule 1, campylotropous; style terminal, usually short, with 2(-5) filiform or subulate stigmas, rarely capitate, papillose, or hairy on one side or throughout. Fruit a utricle, rarely a pyxidium (dehiscent capsule); pericarp membranous, leathery, or fleshy, adnate or appressed to seed. Seed horizontal, vertical, or oblique, compressed globose, lenticular, reniform, or obliquely ovoid; testa crustaceous, leathery, membranous, or succulent; embryo annular, semi-annular, or spiral, with narrow cotyledons; endosperm much reduced or absent; perisperm abundant or absent.

Probably about 100 genera and 1400 species (depending on taxonomic opinions): mainly in arid areas, deserts, and coastal and saline habitats of N and S Africa, Asia, Australia, Europe, and North and South America; 42 genera (two endemic, two introduced) and 190 species (21 endemic, six introduced) in China.

Many species of Chenopodiaceae are adapted to, and are major components of, arid or ruderal environments. They are often intimately involved with the daily life of people. For example, Beta vulgaris is one of the most important sources for sugar; Chenopodium quinoa is a new high-protein crop; Spinacia oleracea and Beta vulgaris are excellent vegetables; Dysphania ambrosioides and Salsola collina are used medicinally; seeds of Agriophyllum squarrosum are called “sand-rice” locally and are edible; seeds of Corispermum declinatum are used for making gin; the ash of Halogeton arachnoideus and some species of Salsola contains soda which is used in noodle-making; and Anabasis aphylla can be used as an insecticide. Many species are important as animal forage in desert, semidesert, and steppe regions, and some species make good windbreaks and soil binders. Haloxylon ammodendron has been used extensively in biological reconditioning of the desert.

Kung Hsien-wu, Chu Ge-lin, C. P. Tsien Cho-po, Ma Cheng-gung & Li An-jen. 1979. Chenopodiaceae. In: Kung Hsien-wu & C. P. Tsien Cho-po, eds., Fl. Reipubl. Popularis Sin. 25(2): 1–194.

1 Embryo spiral; perisperm separated into two parts by embryo, or perisperm absent.   (2)
+ Embryo annular or semi-annular; perisperm copious, surrounded by embryo.   (17)
2 (1) Bractlets rudimentary, membranous, scale-like, hidden by perianth; stigmas papillose or hairy throughout; embryo planospiral.   (3)
+ Bractlets developed, herbaceous or succulent, navicular or similar to leaves, surrounding perianth (in Sympegma bractlets absent, but flowers fascicled at apex of branchlets); stigmas papillose only adaxially; embryo conic-spiral rarely planospiral (in tribe Salsoleae).   (4)
3 (2) Flowers unisexual; perianth of female flowers pellucid-membranous, lobed at apex, enlarged, incrassate, and berrylike in fruit; perianth segments of male flower spreading, caducous; anther 0.6-0.7 mm.   27 Borszczowia
+ Flowers bisexual; perianth segments not spreading; anther to 0.5 mm.   28 Suaeda
4 (2) Perianth segments connate into a tube, with 5 membranous teeth at apex; perianth adaxially with an acicular appendage adnate to it in fruit; axil of bractlets with a fascicle of villous hairs.   29 Cornulaca
+ Perianth segments not connate into a tube, without an acicular appendage; axil of bractlets glabrous or ± hairy, but hairs not fascicular.   (5)
5 (4) Perianth segments without abaxial appendage; branchlets not jointed.   (6)
+ Perianth segments with a well-developed or rudimentary wing or tuberculate appendage.   (8)
6 (5) Subshrubs cushionlike; perianth conspicuously enlarged and exposed in fruit; leaves subulate.   40 Nanophyton
+ Herbs annual; perianth not exposed in fruit; leaves linear, semiterete.   (7)
7 (6) Perianth segments connate, hardening and forming an urceolate body in fruit; anther appendage inflated and bladderlike.   41 Halimocnemis
+ Perianth segments free, proximally becoming leathery and adaxially concave in fruit; anther appendage thick, not inflated and bladderlike, with 3 teeth.   42 Petrosimonia
8 (5) Branchlets jointed; leaves opposite (except in Horaninovia).   (9)
+ Branchlets not jointed; leaves alternate (except in Salsola brachiata).   (13)
9 (8) Seed vertical.   (10)
+ Seed horizontal.   (11)
10 (9) Subshrubs, or woody stems specialized into tuberous caudex, or absent; anthers without appendage; leaves obtuse or acute at apex, sometimes with blunt spines.   33 Anabasis
+ Herbs annual; anthers with a slender, mucronate appendage at apex; leaves spinose at apex.   34 Girgensohnia
11 (9) Herbs annual; leaves and bracts with a long spine at apex.   30 Horaninovia
+ Subshrubs, shrubs, or small trees; leaves and bracts without a long spine at apex.   (12)
12 (11) Shrubs or small trees; flowers emerging from lateral, dwarf branches on second year's growth; perianth membranous, with a winged appendage in fruit; utricle slightly concave; disk inconspicuous.   31 Haloxylon
+ Subshrubs; flowers emerging from annual branches; perianth slightly succulent, with a rudimentary winged appendage; disk conspicuous.   32 Arthrophytum
13 (8) Flowers usually 3, borne at apex of a dwarf branchlet.   35 Sympegma
+ Flowers solitary or glomerulate in leaf axils.   (14)
14 (13) Winged appendage of perianth segment attached subapically.   (15)
+ Winged appendage of perianth segment attached at middle.   (16)
15 (14) Herbs annual; flowers glomerulate; perianth conic; leaves expanded at base.   36 Halogeton
+ Subshrubs; flowers solitary; perianth subglobose; leaves not expanded at base.   37 Iljinia
16 (14) Portion of perianth segment below wing enlarged, hardened, and woody in fruit.   38 Halothamnus
+ Portion of perianth segment below wing not enlarged, hardened, or woody in fruit.   39 Salsola
17 (1) Fruit a pyxidium, dehiscent by a lid.   1 Acroglochin
+ Fruit a utricle, indehiscent or irregularly dehiscent.   (18)
18 (17) Perianth basally adnate to ovary, enlarged, incrassate, and hardened in fruit.   2 Beta
+ Perianth free from ovary, not enlarged, incrassate, or hardened in fruit (in tribe Atripliceae ovary is attached to 2 specialized bractlets).   (19)
19 (18) Flowers borne in axil of succulent bracts, appearing sunken into rachis; leaves reduced to scales or succulent-tuberculate, decurrent if terete.   (20)
+ Flowers free from rachis; leaves usually well developed.   (24)
20 (19) Herbs annual.   (21)
+ Shrubs or subshrubs.   (22)
21 (20) Branches and leaves opposite.   3 Salicornia
+ Branches and leaves alternate.   4 Halopeplis
22 (20) Branchlets not jointed; leaves alternate.   5 Kalidium
+ Branchlets jointed; leaves opposite.   (23)
23 (22) Subshrubs; spikes sessile.   6 Halocnemum
+ Shrubs; spikes pedunculate.   7 Halostachys
24 (19) Flowers unisexual (plants monoecious or dioecious).   (25)
+ Flowers bisexual or plants sometimes polygamous.   (31)
25 (24) Plant body covered with stellate or ramified indumentum.   (26)
+ Plant body glabrous or furfuraceous.   (28)
26 (25) Female flowers with perianth.   9 Axyris
+ Female flowers without perianth.   (27)
27 (26) Shrubs or subshrubs; bracts of female flowers united below middle, forming a tube, with 4 fascicles of villous hairs (with only ramified hairs in Krascheninnikovia compacta).   10 Krascheninnikovia
+ Herbs annual; bracts of female flowers connate to apex, both sides with an acicular appendage near apex.   14 Ceratocarpus
28 (25) Female flowers several, borne at base of a leaflike bract.   (29)
+ Female flowers borne in axil of a leaflike bract in a cup formed by 2 connate bracts.   (30)
29 (28) Female flowers with evident perianth.   11 Archiatriplex
+ Female flowers without evident perianth; bracts of female flowers 3-lobed, lateral lobes inflexed.   8 Microgynoecium
30 (28) Stigmas 4 or 5; plant body glabrous.   13 Spinacia
+ Stigmas 2; plant body ± covered with furfuraceous indumentum.   12 Atriplex
31 (24) Perianth segments 1-3, white, membranous; utricles exposed, compressed, with 2-fid beak at apex; plant body ± covered with ramified hairs.   (32)
+ Perianth (3 to)5-parted, succulent or herbaceous; utricles flattened, rarely only compressed, without beak; plant body without ramified hairs.   (33)
32 (31) Utricles convex on both sides; rostra nearly equal to length of kernel; seed free from pericarp; leaves and bracts acicular to spinulose at apex.   15 Agriophyllum
+ Utricles convex abaxially, plane or slightly concave adaxially; rostra 1/8-1/5 length of kernel; seed adherent to pericarp; leaves and bracts acute but not acicular at apex.   16 Corispermum
33 (31) Plant body usually furfuraceous, sometimes glabrous, or glandular and strongly aromatic.   (34)
+ Plant body pubescent; leaves terete, semiterete, rarely flattened.   (37)
34 (33) Flowers with 2 scale-like, membranous bractlets; seed vertical.   (35)
+ Flowers without bractlets; seed horizontal or oblique or, if vertical, perianth 3- or 4-parted.   (36)
35 (34) Leaves petiolate, leaf blade flattened.   17 Baolia
+ Leaves sessile, filiform, semiterete.   18 Polycnemum
36 (34) Plants covered with glandular hairs (subglabrous in Dysphania aristata, but then terminal inflorescence branches without flowers, ending with acute setae).   19 Dysphania
+ Plants covered with vesicular hairs (furfuraceous), occasionally glabrous (but then terminal inflorescence branches bearing flowers).   20 Chenopodium
37 (33) Perianth segments with a winged, acicular, or tuberculate appendage abaxially in fruit.   (38)
+ Perianth segments without an appendage in fruit.   (40)
38 (37) Appendage borne on distal portion of perianth segment; seed vertical or oblique.   23 Panderia
+ Appendage borne on middle of perianth segment; seed horizontal.   (39)
39 (38) Perianth segment appendage winglike, veined.   21 Kochia
+ Perianth segment appendage acicular, veinless.   22 Bassia
40 (37) Subshrubs; perianth teeth 4; stamens 4; leaves semiterete.   24 Camphorosma
+ Herbs annual; perianth teeth 5; stamens 5; leaves flattened.   (41)
41 (40) Perianth adnate to utricle, densely villous, resembling a woolly ball; seed horizontal (vertical in bisexual flowers); stigmas smooth; anthers yellow, ca. 0.7 mm.   25 Londesia
+ Perianth free from utricle, villous but not resembling a woolly ball; seed vertical; stigmas papillose; anthers usually purplish red, ca. 1.5 mm.   26 Kirilowia

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