Including Atriplicaceae Juss., Corispermaceae Link, Farinaceae Dulac, Salicorni(ac)eae J.G. Agardh, Salsolaceae Moq.-Tand
Excluding Dysphaniaceae, Halophytaceae
Habit and leaf form. Herbs (mostly), or shrubs (some), or trees (few, small), or lianas (few). `Normal' plants and switch-plants; sometimes more or less `cactoid' (e.g. Salicornia). Leaves well developed, or much reduced, or absent. Plants succulent, or non-succulent. Annual, or biennial, or perennial (often glaucous); with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting (usually), or climbing (few). Helophytic to xerophytic (nearly all halophytic). Leaves minute to large; alternate, or opposite; when alternate, spiral, or distichous (rarely); `herbaceous', or fleshy, or membranous; petiolate to sessile; when opposite, connate (not uncommonly, e.g. represented by lobed cupules at the internodes of succulent stems), or not connate; sheathing, or non-sheathing; simple; epulvinate. Lamina dissected, or entire; one-veined (?), or pinnately veined; hastate, or sagittate, or attenuate at the base, or cuneate at the base. Leaves exstipulate; without a persistent basal meristem.
General anatomy. Plants with `crystal sand', or without `crystal sand'.
Leaf anatomy. Hydathodes present (sometimes), or absent.
Minor leaf veins without phloem transfer cells (6 genera).
Stem anatomy. Cork cambium present; initially deep-seated, or superficial. Nodes unilacunar. Cortical bundles absent. Medullary bundles present (commonly), or absent. Internal phloem absent. Secondary thickening developing from a conventional cambial ring, or anomalous; when anomalous, via concentric cambia (very commonly), or from a single cambial ring (?). `Included' phloem present (very commonly), or absent. Xylem with libriform fibres; with vessels. Vessel end-walls simple. Wood not storied (VP, in all those with secondary growth); parenchyma paratracheal (and conjunctive). Sieve-tube plastids P-type; type III (c).
Reproductive type, pollination. Hermaphrodite (usually), or monoecious (sometimes), or andromonoecious (rarely), or gynomonoecious, or dioecious (rarely), or polygamomonoecious.
Inflorescence, floral, fruit and seed morphology. Flowers solitary (then axillary), or aggregated in `inflorescences' (or sometimes paired); when flowers aggregated, in cymes (the cymes mostly in racemose arrangements). The terminal inflorescence unit cymose. Inflorescences terminal, or axillary; mostly racemes of cymes. Flowers minute, or small; regular; cyclic; usually pentacyclic (sometimes 2-cyclic). Free hypanthium present (the stamens then inserted `on a disk on the calyx'), or absent. Hypogynous disk often present.
Perianth sepaline, or vestigial to absent; (1-)5(-6); joined (imbricate); 1 whorled; fleshy, or non-fleshy; persistent; accrescent, or non-accrescent. Calyx (interpreting the perianth as such) (1-)3-5(-6); gamosepalous; fleshy, or non-fleshy; persistent (in the fruit); accrescent, or non-accrescent; imbricate.
Androecium 3-5. Androecial members free of the perianth, or adnate (to the base of the perianth); all equal; free of one another, or coherent; when coherent 1 - adelphous (connate at the base); 1 - whorled. Androecium exclusively of fertile stamens. Stamens 3-5; usually isomerous with the perianth; oppositisepalous. Anthers bent inwards in bud; dehiscing via longitudinal slits; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer; of the `monocot' type. Tapetum glandular (usually), or amoeboid. Pollen grains aperturate; 8-90 - aperturate; (oligo-to poly-) foraminate; 3-celled.
Gynoecium (2-)5; syncarpous; synovarious to synstylovarious; superior (usually), or partly inferior (rarely). Ovary 1 locular. Styles (1-)2-3(-4); usually partially joined. Stigmas dry type; papillate; Group II type. Placentation basal. Ovules in the single cavity 1; pendulous, or ascending; non-arillate; campylotropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei `usually' fusing prior to fertilization. Antipodal cells formed; 3; proliferating (a little, e.g. Beta vulgaris), or not proliferating (usually); usually ephemeral. Synergids pear-shaped, or hooked. Endosperm formation nuclear. Endosperm haustoria present, or absent; when developed, chalazal. Embryogeny chenopodiad.
Fruit non-fleshy; indehiscent, or dehiscent; a nut (usually), or a capsule (infrequently); enclosed in the fleshy receptacle, or enclosed in the fleshy hypanthium. Capsules circumscissile. Fruits from adjoining flowers aggregated into compound fruits, or not aggregated into compound fruits. Seeds more or less non-endospermic. Perisperm present, or absent. Seeds with starch. Cotyledons 2. Embryo chlorophyllous (5/11), or achlorophyllous (3/5 (Atriplex, Chenopodium, Beta)); coiled (usually), or curved, or bent.
Seedling. Germination phanerocotylar.
Physiology, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present, or absent. Iridoids absent. Proanthocyanidins absent. Flavonols present; kaempferol, or kaempferol and quercetin. Ellagic acid absent (7 species, 7 genera). Arbutin absent. Betalains present. Saponins/sapogenins present, or absent. Aluminium accumulation not found. Plants often accumulating free oxalates. Inulin recorded (? - Beta). C3, or C4. C3 recorded in Agriophyllum, Allenrolfia, Anthochlamys, Arthrocnemum, Atriplex, Bassia, Beta, Blitum, Ceratocarpus, Chenopodium, Eurotia, Halimione, Halisarcia, Halocnemum, Halosarcia, Haplopeplis, Halostachya, Kalidium, Micronemum, Pachycornia, Salicornia, Salsola, Sclerostegia, Spinacia, Suaeda, Tecticornia. C4 recorded in Aellinia, Agathophora, Anabasis, Arthrophytum, Atriplex, Bassia, Camphorosma, Climacoptera, Cornulaca, Gamanthus, Girgensohnia, Halanthium, Halarchon, Halimocnemis, Halocharis, Halogeton, Halosarcia, Halotis, Haloxylon, Hammada, Horaninovia, Hypocyclix, Kochia, Noaea, Panderia, Petrosimonia, Salsola, Suaeda, Theleophyton, Traganum. Anatomy C4 type (Aellenia, Anabasis, Arthrophytum, Atriplex, Bassia, Camphorosma, Chenolea, Girgensohnia, Halanthium, Halosarcia, Kochia, Panderia, Petrosimonia, Salsola, Suaeda, etc.), or non-C4 type (Allenrolfea, Arthrocnemum, Atriplex, Bassia, Ceratocarpus, Chenolea, Chenopodium, Halimione, Halocnemum, Halopeplis, Halosarcia, Kochia, Maireana, Pachycornia, Salicornia, Sarcocornia, Sclerostegia, Suaeda, Tecticornia etc.).
Geography, cytology. Temperate to sub-tropical. Widespread, but missing from Central America, Asia, Malaysia and most of Africa. X = (6-)9.
Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren's Superorder Caryophylliflorae; Caryophyllales. Cronquist's Subclass Caryophyllidae; Caryophyllales. Takhtajan's Subclass Caryophyllidae; Caryophyllanae; Caryophyllales. Species 1400. Genera about 105; Acroglochin, Aellenia (= Halothamnus), Agathophora, Agriophyllum, Alexandra, Allenrolfea, Anabasis, Anthochlamys, Aphanisma, Archiatriplex, Arthocnemum, Arthrophytum, Atriplex, Axyris, Baolia, Bassia, Beta, Bienertia, Blitum, Borsczowia, Camphorosma, Ceratocarpus, Chenopodium, Climacoptera, Corispermum, Cornulaca, Cremnophyton, Cyathobasis, Cycloloma, Didymanthus, Dissocarpus, Einadia, Enchylaena, Eremophea, Eriochiton, Exomis, Fadenia, Fredolia, Gamanthus, Girgensohnia, Grayia, Hablitzea, Halanthium, Halarchon, Halimione, Halimocnemis, Halocharis, Halocnemum, Halogeton, Halopeplis, Halosarcia, Halostachys, Halothamnus, Haloxylon, Hammada, Hemichroa (~ Amaranthaceae), Heterostachys, Holmbergia, Horaninovia, Iljinia, Kalidium, Kirilowia, Kochia, Krascheninnikovia, Lagenantha, Maireana, Malacocera, Manochlamys, Microcnemum, Microgynoecium, Monolepis, Nanophyton, Neobassia, Nitrophila, Noaea, Nucularia, Ofaiston, Oreobliton, Osteocarpum, Pachycornia, Panderia, Petrosimonia, Physandra, Piptoptera, Polycnemum, Rhagodia, Raphidophyton, Roycea, Salicornia, Salsola, Sarcobatus, Sarcocornia, Scleroblitum, Sclerochlamys, Sclerolaena, Sclerostegia, Seidlitzia, Sevada, Spinacia, Stelligera, Suaeda, Suckleya, Sympegma, Tecticornia, Tegicornia, Teloxys, Threlkeldia, Traganopsis, Traganum, Zuckia.
Economic uses, etc. Contributes sugarbeet and Swiss chard (Beta vulgaris), spinach (Spinacia oleracea), and `saltbush' pastures.
Illustrations. cheno638.gif cheno512.gif cheno179.gif
Additional, to be intercalated. Flowers when solitary or paired, axillary.
