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Transcript
Chenopodiaceae Vent.
~ Amaranthaceae.
Including Atriplicaceae Juss., Corispermaceae Link, Farinaceae Dulac, Salicornieae
(Salicorniaceae) J.G. Agardh, Salsolaceae Moq.-Tandon; excluding Dysphaniaceae, Halophytaceae,
Sarcobataceae.
Habit and leaf form. Herbs (mostly halophytic), or shrubs (some), or trees (few, small), or lianas
(few). ‘Normal’ plants and switch-plants; sometimes more or less ‘cactoid’, with succulent,
photosynthetic stems (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 without conspicuous 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; leaf development not ‘graminaceous’.
General anatomy. Plants with ‘crystal sand’, or without ‘crystal sand’.
Leaf anatomy. The leaf lamina when not much reduced, dorsiventral (rather uncommonly), or
bifacial to centric. Hydathodes present (sometimes), or absent. Stomata universally on both surfaces (or in
all parts of the epidermes); anomocytic (usually), or paracytic (in a few species and genera). Hairs of
numerous kinds present (see illustration); eglandular and glandular; mostly multicellular. Multicellular
hairs uniseriate (eclusively?); branched and simple (commonly 2-armed, sometimes candelabra type).
Complex hairs present, or absent; sometimes stellate. Adaxial hypodermis present, or absent. The
mesophyll containing crystals. The crystals mainly druses (and sand). Minor leaf veins without phloem
transfer cells (6 genera).
Axial (stem, wood) anatomy. Cork cambium present, or absent; when present, initially
deep-seated, or initially superficial. Nodes unilacunar. Primary vascular tissues comprising a ring of
bundles; collateral (?). Internal phloem absent. Cortical bundles absent. Medullary bundles present, or
absent (ostensibly common, but supposedly representing leaf traces laid down before the onset of
secondary thickening), or absent. Secondary thickening developing from a conventional cambial ring
(rarely?), or anomalous (nearly always). The anomalous secondary thickening usually via concentric
cambia (these comprising rings or arcs, generating numerous vascular bundles in spirals, in rings or
distributed irregularly).
The wood semi-ring porous (occasionally), or diffuse porous. The vessels small (sometimes extremely
so); in radial multiples, or clustered, or in tangential arcs (typically in groups or clusters internal to the
phloem strands). The vessel end-walls simple. The vessels with spiral thickening (commonly), or without
spiral thickening. The axial xylem with tracheids, or without tracheids; without fibre tracheids; with
libriform fibres; without septate fibres. The fibres without spiral thickening. The parenchyma paratracheal
(and conjunctive). ‘Included’ phloem present (very commonly, typically of the concentric type but
occasionally foraminate), or absent. The wood not storied (VP, in all those with secondary growth).
Reproductive type, pollination. Plants 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 solitary or paired, axillary; when flowers aggregated, in
cymes (the cymes mostly in racemose arrangements). The ultimate inflorescence units cymose.
Inflorescences terminal, or axillary; mostly racemes of cymes. Flowers minute, or small; regular; cyclic;
when hermaphrodite, usually tricyclic (sometimes 2-cyclic). Free hypanthium present (the stamens then
inserted ‘on a disk on the calyx’), or absent. Hypogynous disk often present (but not always?).
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 (nearly always), or polysepalous (in Salsola?); commonly becoming
appendaged with spines, tubercles or wings; fleshy (commonly), or non-fleshy (then usually
herbaceous or membranous); persistent (in the fruit); accrescent (usually, enclosing the fruit and often
appendaged with wings, tubercles or spines), or non-accrescent; imbricate.
Androecium 3–5. Androecial members free of the perianth, or adnate (to the base of the perianth); all
equal (by contrast with most Amaranthaceae); free of one another (usually, by contrast with most
Amaranthaceae), or coherent; when coherent 1 adelphous (connate at the base); 1 whorled.
Androecium nearly always exclusively of fertile stamens (by contrast with many Amaranthaceae).
Stamens 3–5; usually isomerous with the perianth; oppositisepalous (when ascertainable). 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; spinulose; 3-celled (7 genera).
Gynoecium (2–)5 carpelled. The pistil 1 celled. Gynoecium 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 capsular-indehiscent (commonly a
utricle); enclosed in the fleshy hypanthium, or enclosed in the fleshy perianth (commonly), or without
fleshy investment external to the original ovary. Gynoecia of adjoining flowers combining to form a
multiple fruit, or not forming a multiple fruit. Seeds more or less non-endospermic. Perisperm present, or
absent (the variation of classificatory interest). 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, phytochemistry. C3, or C4. C3 physiology recorded directly 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
physiology recorded directly 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 (see illustration), Bassia (see illustration),
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, Salsola (see illustration), Sarcocornia, Sclerostegia, Suaeda, Tecticornia etc.).
Inulin recorded (? — Beta). Cyanogenic, or not cyanogenic. Alkaloids present, or absent.
Anthraquinones detected (Salsola); polyacetate derived. Arbutin absent. Iridoids not detected. Betalains
present. Saponins/sapogenins present, or absent. Proanthocyanidins absent. Flavonols present;
kaempferol, or kaempferol and quercetin. Ellagic acid absent (7 species, 7 genera). Aluminium
accumulation not found. Plants often accumulating free oxalates. Sieve-tube plastids P-type; type III (c).
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. APG 3 core angiosperms; core
eudicot; Superorder Caryophyllanae; Order Caryophyllales (under Amaranthaceae!).
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,
Sarcocornia, Scleroblitum, Sclerochlamys, Sclerolaena, Sclerostegia, Seidlitzia, Sevada, Spinacia,
Stelligera, Suaeda, Suckleya, Sympegma, Tecticornia, Tegicornia, Teloxys, Threlkeldia, Traganopsis,
Traganum, Zuckia.
General remarks. Comparison of this description with that of Amaranthaceae (q.v.) shows
markedly differing tendencies in some overlapping characters, especially in habitat preferences and in
details of the androecium (equal, free stamens and no staminodes in most chenopods); but there seem to
be no convincing absolute differences, and it seems legitimate to regard the chenopods as mainly
halophytic Amaranthaceae.
Economic uses, etc. Contributes sugarbeet and Swiss chard (Beta vulgaris), spinach (Spinacia
oleracea), and ‘saltbush’ pastures.