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goosefoot family

Habit Herbs, annual. Herbs, shrubs (rarely small trees), annual or perennial, monoecious, dioecious, or polygamous, evergreen or deciduous.
Roots

fibrous, taprooted, sometimes fusiform or bulbous, fleshy and thickened in Beta.

Stems

erect, ascending, or procumbent, much branched, obtusely angled, 1–6 dm, stout, scurfy when young.

sometimes succulent and apparently jointed, or with slippery and aromatic bark, sometimes spiny, alternate or opposite;

pubescence silvery, sometimes stellate or glandular, often scurfy from inflated salt glands that senesce into white flakes.

Leaves

alternate or proximalmost opposite or subopposite;

petiole to 1.5 cm or sessile;

blade paler abaxially, oblong or oval, lanceolate, or elliptic to broadly obovate or narrowly oblong, 10–40(–60) × (2–)4–20 mm, base rounded to cuneate, margin entire or undulate, rarely with 1–2 teeth, apex rounded to acute, mucronate, thin, often densely white scurfy beneath, grayish green and glabrate above.

simple, usually alternate, occasionally opposite, lacking stipules, petiolate or sessile, sometimes reduced to small scales, or fleshy;

blade linear to broadly triangulate in outline, margins entire to serrate, serrate-dentate, or lobed.

Inflorescences

flowers solitary or clustered in axillary or terminal glomerules or in short, cylindric spikes;

bracts absent or 1–5, deciduous or persistent, of various shapes.

Flowers

bisexual or unisexual, uniseriate, radially or rarely bilaterally symmetric;

bracteoles absent or 1–5, connate basally, green;

perianth segments 5, sometimes 1 or absent, green, inconspicuous, fleshy in Salicornia and Sarcocornia, strongly imbricate in Nitrophila;

petals absent;

stamens absent or 1–5, usually as many as and opposite perianth lobes;

pistils absent or (1–)2(–3);

styles 1–3, sometimes with stylopodium;

ovary usually superior, half-inferior in Sarcobatus, inferior and connate with receptacle in fruit in Beta, 1-locular with single, basally attached ovule.

Pistillate flowers

fascicled in axils.

Ovules

usually 1, campylotropous, bitegmic, crassinucellate.

Seeds

reddish brown, 2 mm.

1 per flower, black, brown, reddish brown, or mixture, flattened vertically or rounded, margins winged or not winged, surfaces smooth and shiny or reticulate, regulate, verrucate, prickly, or indistinct, morphology variable and strongly influenced by plant photoperiod;

seed coat smooth, striate, or verrucate when pericarp is removed;

embryo large, curved to annular or spirally coiled;

radicle position median or basal, ascending or pointing outward;

endosperm usually digested by developing embryo and food storage taken over by perisperm.

Staminate

glomerules terminal or in dense or interrupted, terminal or axillary, naked spikes or shortly branched panicles.

Fruiting

bracteoles subsessile or with stipes to 1 mm, cuneate-orbiculate, compressed, 4.5–7 × 3.5–5.6 mm, typically longer than wide, united to middle, apex rounded, 3–5-toothed, teeth subequal, sides irregularly tuberculate or with 2 lateral dentate crests, rarely not appendaged.

structures: bracteoles or fruiting bracts brown, black, or reddish brown, monomorphic or sometimes dimorphic;

perianth segments deciduous or persistent in mature fruits and of various shapes and ornamentation, accrescent around fruits;

fruits achenes or utricles, vertical or horizontal within perianth parts, pericarp (ovary wall) adherent or nonadherent, chartaceous or papery, sometimes reticulate, mottled or smooth.

Polyploidy

common.

x

= 9.

2n

= 18.

Atriplex mucronata

Chenopodiaceae

Phenology Flowering summer–fall.
Habitat Sandy seashores, salt marshes
Elevation 0 m (0 ft)
Distribution
from FNA
AL; FL; LA; MA; MD; MS; NC; NH; NJ; TX; VA
[BONAP county map]
Worldwide; especially in desert and semidesert regions; often in alkaline or saline habitats
Discussion

Problems with prior applications of the name Atriplex mucronata lie in the mistaken determination of the place of publication as Amer. Monthly Mag. & Crit. Rev. 2: 176. 1818 (where the name is only mentioned), instead of 2: 119. 1817 (where accompanied by a description and notes). H. M. Hall and F. E. Clements (1923) evidently relied on P. C. Standley’s (1916) interpretation, which indicated the wrong citation (see also A. dioica Rafinesque as an identical example). Hall and Clements applied the name mucronata to their interpretation of A. patula subsp. hastata or to A. hastata (i.e., to A. prostrata according to this treatment).

The treatment of Obione by C. H. B. A. Moquin-Tandon (1849) included “A. mucronata Rafin.!” as a synonym of O. arenaria. Possibly a sheet in the Prodromus herbarium at Geneva was the basis for that decision. It has two mounted specimens, one labeled A. arenaria Nuttall, collected by Nuttall in “N. Jersey, 1826,” and a second labeled “Atriplex mucronata Rafinesque (A. arenaria Nuttall, N. Jersey) Maritime NY, Rafinesque 1819.” From that information (although the year is 1819, not 1817), and from the description of the taxon, it seems clear that A. arenaria Nuttall is a later synonym of A. mucronata Rafinesque by at least half a year.

H. A. Gleason and A. Cronquist (1991) indicated that this taxon, by whatever name, is “perhaps better treated as a variety of the more tropical Atriplex pentandra (Jacquin) Standley, but the proper nomenclatural combination not yet made.” The present writer agrees with that conclusion, but such subjugation might indicate further contraction into the species of additional closely related taxa, e.g., A. wrightii, which is clearly closely allied as well.

Plants from the coastal states from New England south to New Jersey are much alike and seldom, if ever, display prominent, terminal, naked spikes or panicles with beadlike glomerules of staminate flowers. Plants from Florida westward sometimes have such staminate spikes or panicles.

Specimens from Florida and some from Texas can be distinguished from Atriplex pentandra only with difficulty, especially those individuals with entire leaves. However, the fruiting bracteole length-width ratio and overall shape, with some allowance for overlap, can serve to distinguish most specimens; those of A. mucronata are proportionately longer than broad and, on average, larger.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera ca. 100, species ca. 1500 (27 genera, 168 species in the flora).

A number of species introduced from Europe and Asia are weedy in North America. The widespread distribution of the family in the deserts of Eurasia and Australia is indicative of the ancient status of the family. Fossil pollen from this family dates to the Maestrichtian, providing the oldest known fossils in the Caryophylliidae.

Plants in this family typically have Crassulacean Acid Metabolism, have either a C3 or C4 photosynthetic pathway (W. V. Brown 1975; G. W. Welkie and M. Caldwell 1970), accumulate organic acids, free nitrates, and oxalates, and often contain alkaloids. Along with other members of the Caryophyllales, members of the family contain pigments called betalains (named for the genus Beta) rather than anthocyanins.

Economically important members of this family include spinach and chard (Spinacia oleracea) and beets (Beta vulgaris). Seeds in this family generally provide a rich source of protein, and one species, Chenopodium quinoa, is gaining widespread acceptance as a cereal crop. Toxicity from high levels of nitrates or oxalates has been reported for a number of species (J. M. Kingsbury 1964), and the pollen is known to be allergenic (T. C. Fuller and E. McClintock 1986). The nutritional characteristics of many species that we share with northern Asia were described by M. M. Iljin (1936).

Molecular and morphologic studies provide evidence supporting the inclusion of the Chenopodiaceae within Amaranthaceae (Angiosperm Phylogeny Group 1998; W. S. Judd and I. K. Ferguson 1999; J. E. Rodman 1990). However, until discordant elements within these lineages, such as Sarcobatus (H.-D. Behnke 1997), are interpreted within a larger evolutionary scheme, the disposition of family groups remains problematic.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Leaves scalelike; stems fleshy or succulent; plants of saline habitats
→ 2
1. Leaves well developed, not scalelike; stems not fleshy (except Arthrocnemum, and Sarcocornia, which is fleshy when young); plants of various habitats
→ 5
2. Leaves alternate
Allenrolfea
2. Leaves opposite
→ 3
3. Plants annual herbs; all stems terminated by an inflorescence
Salicornia
3. Plants perennial shrubs; many stems entirely vegetative
→ 4
4. Flowers distinct in each cyme, not adnate; outer seed coat hard, tuberculate
Arthrocnemum
4. Flowers of each cyme adnate to distal branch of inflorescence; outer seed coat thin, covered with hooked or straight hairs
Sarcocornia
5. Leaves opposite; perianth strongly imbricate
Nitrophila
5. Leaves alternate, rarely opposite; perianth slightly, if at all imbricate
→ 6
6. Inflorescence leaves or bracts tipped with spine or spinelike bristle
→ 7
6. Inflorescence leaves and bracts not tipped with spine or bristle
→ 9
7. Leaves triquetrous, triangular in cross section, terminating in soft or rarely rigid, flattened bristle
Polycnemum
7. Leaves terete or flat in cross section, terminating in stiff, mucronate spine tips or bristle
→ 8
8. Fruiting perianth abaxially winged; leaves linear to subulate, herbaceous or fleshy, spine-tipped or not; bracts of inflorescence ovate-lanceolate, spine tipped
Salsola
8. Fruiting perianth apically winged; leaves terete, fleshy-succulent, bristle- tipped; bracts of inflorescence similar to leaves
Halogeton
9. Leaves sub- or semicylindric to linear, usually fleshy
→ 10
9. Leaves with flattened blades, not especially fleshy
→ 12
10. Shrubs armed with thorny branches; flowers unisexual, staminate flowers in spikes, pistillate flowers solitary or paired and axillary
Sarcobatus
10. Shrubs or herbs, not armed; flowers bisexual or both bisexual and pistillate, solitary or 2-5 in axillary clusters
→ 11
11. Fruiting perianth developing horizontal, membranceous wings or tubercles; leaves herbaceous
Kochia
11. Fruiting perianth without wings or tubercules, unchanged from flowering peri- anth; leaves fleshy
Suaeda
12. Plants stellate-pubescent annual herbs
Axyris
12. Plants glabrous, farinose, or with dendritic hairs; if the latter, then plants woody perennials
→ 13
13. Plants densely tomentose with at least some stellate hairs becoming golden brown in age, subshrubs
Krascheninnikovia
13. Plants hairy or glabrous, not as above, shrubs or herbs
→ 14
14. Pistillate flowers usually lacking perianth, at least some flowers enclosed by 2 accrescent or connate bracts in fruit; flowers unisexual or, rarely, bisexual
→ 15
14. Perianth present, not enclosed by paired bracts in fruit; flowers bisexual or unisexual (Micromonolepis) or some also pistillate
→ 19
15. Stigmas 4 or 5; plants cultivated herbs
Spinacia
15. Stigmas 2; plants not or seldom cultivated
→ 16
16. Bracts (or bractlike perianth segments) dorsally compressed, usually triangular, smooth, tuberculate, vertically keeled, or winged; pubescence usually of inflated hairs or, sometimes, none; axillary rounded buds absent or inconspicuous
→ 17
16. Bracts laterally compressed, keeled, or winged, lacking appendages; pubescence of simple or branched hairs
→ 18
17. Bractlike perianth segments united below middle, vertically keeled; plants annual
Suckleya
17. Bracts united above middle, not vertically keeled; plants pe- rennial or annual
Atriplex
18. Shrubs with divaricate, often spinescent branches; bracts with margins thickened, spongy within; pubescence of branched hairs
Grayia
18. Shrubs with erect thornless branches; bracts with margins not spongy-thickened, either obcompressed or dorsiventrally compressed and 6-ribbed; pubescence of scurfy or moniliform hairs
Zuckia
19. Perianth horizontally winged in fruit
→ 20
19. Perianth not horizontally winged in fruit
→ 21
20. Leaf blade margins sinuate-dentate; plants ± villous or tomentulose, becoming glabrous with maturity
Cycloloma
20. Leaf blade margins entire; plants usually glabrous, sometimes slightly pubescent
Kochia
21. Proximal leaves petiolate, middle ones merely sessile, distal ones cordate-clasping; style branches 3
Aphanisma
21. Leaves not as above; style branches mainly 2
→ 22
22. Perianth segments each armed with spiniform, hooked, or conic appendages
Bassia
22. Perianth segments rounded or keeled abaxially, lacking wings or spines
→ 23
23. Ovary partly inferior; plants cultivated, rarely naturalized
Beta
23. Ovary superior; plants native or naturalized, not cultivated
→ 24
24. Perianth lobes 5, largely enfolding and concealing to exposing fruits; stamens usually 5
→ 25
24. Perianth lobes 1-3 or absent; fruits largely exposed; stamens 1-3(-5)
→ 26
25. Plants (at least some parts) with glandular or glandular-vesicular hairs
Dysphania
25. Plants farinose or glabrous
Chenopodium
26. Plants dichotomously branched, ultimate branches filiform; leaves oblong, succulent
Micromonolepis
26. Plants not dichotomously branching, ultimate branches not filiform; leaves triangular-lanceolate to oblanceolate, spatulate, or linear
→ 27
27. Leaves triangular-lanceolate to oblanceolate or spatulate; perianth segments usually 1 (2-3 segments in central flowers); stamens usually 1; plants widespread, of many habitats
Monolepis
27. Leaves lanceolate, linear-lanceolate, or filiform; perianth segments 1-3; stamens 1-3(-5); plants of sandy, low elevation sites.
Corispermum
Source FNA vol. 4, p. 362. FNA vol. 4, p. 258. Authors: Stanley L. Welsh, Clifford W. Crompton, Steven E. Clemants.
Parent taxa Chenopodiaceae > Atriplex > subg. Obione > sect. Obione > subsect. Arenariae
Sibling taxa
A. acanthocarpa, A. amnicola, A. argentea, A. californica, A. canescens, A. confertifolia, A. cordulata, A. coronata, A. corrugata, A. coulteri, A. covillei, A. dioica, A. elegans, A. fruticulosa, A. gardneri, A. garrettii, A. glabriuscula, A. gmelinii, A. graciliflora, A. heterosperma, A. holocarpa, A. hortensis, A. hymenelytra, A. joaquiniana, A. klebergorum, A. laciniata, A. lentiformis, A. leucophylla, A. lindleyi, A. linearis, A. littoralis, A. matamorensis, A. nudicaulis, A. nummularia, A. oblongifolia, A. obovata, A. pacifica, A. parishii, A. parryi, A. patula, A. pentandra, A. phyllostegia, A. pleiantha, A. polycarpa, A. powellii, A. prostrata, A. pusilla, A. rosea, A. saccaria, A. semibaccata, A. serenana, A. spinifera, A. suberecta, A. suckleyi, A. tatarica, A. torreyi, A. truncata, A. tularensis, A. watsonii, A. wolfii, A. wrightii
Subordinate taxa
Allenrolfea, Aphanisma, Arthrocnemum, Atriplex, Axyris, Bassia, Beta, Chenopodium, Corispermum, Cycloloma, Dysphania, Grayia, Halogeton, Kochia, Krascheninnikovia, Micromonolepis, Monolepis, Nitrophila, Polycnemum, Salicornia, Salsola, Sarcobatus, Sarcocornia, Spinacia, Suaeda, Suckleya, Zuckia
Synonyms A. arenaria, A. cristata var. arenaria, A. pentandra subsp. arenaria
Name authority Rafinesque: Amer. Monthly Mag. & Crit. Rev. 2(2): 119. (1817) Ventenat
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