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siltbush, spineless hopsage

goosefoot family

Habit Shrubs, branching from persistent woody base 0.5–2 dm; stems 1–5 dm. 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

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

13–80 × 15–42 mm.

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.

Staminate flowers

with perianth cleft to middle or below, 1.5–1.8 mm.

Pistillate flowers

few, intermixed in otherwise staminate spikes;

bracteoles laterally or vertically compressed, with vertical or horizontal achenes respectively, when mature either laterally flattened, 2(–4)-winged, 3.4–9 mm diam., or dorsiventrally compressed and not or 1–4(–5)-ribbed and 2-winged, (1.8–)2–2.5 mm diam.

Achenes

included within bracts, 1.2–2.2 × 1.2–2.2 mm.

Ovules

usually 1, campylotropous, bitegmic, crassinucellate.

Seeds

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.

Fruiting

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.

Zuckia brandegeei

Chenopodiaceae

Discussion

Varieties 3 (3 in the flora).

When he described Grayia brandegeei A. Gray (1876b) noted, “While pleased with an accession to this genus, and with the opportunity of associating it with the name of an excellent correspondent who discovered it, I must add that it does not much strengthen the genus.” Grayia, within which the species has long resided, shorn of G. brandegeei, consists of G. spinosa (Hooker) Moquin-Tandon. Grayia brandegeei was clearly regarded, even by its author, as not closely allied to G. spinosa, the type species of Grayia, even though they shared features of bracteole compression, seed position, and rounded axillary buds. The two species otherwise differ markedly in stature, vestiture, and in the nature of the bracts. In G. spinosa the bracteoles are thickened marginally and filled internally with a spongy cellular matrix. Those of G. brandegeei are thin margined and lack a spongy cellular matrix. Leaf shape, plant stature, and ecological associates also differ. Grayia spinosa is a plant mainly of low-salinity substrates, and is rather widely distributed over the West from Washington to Montana and south to California, Arizona, and New Mexico (S. L. Welsh et al. 1993).

Besides the similarity of stature, staminate inflorescences, and flowers, the plants herein considered within Zuckia occupy similar, relatively high-saline, gypsiferous, and seleniferous substrates. The major difference between the lone taxon, Z. arizonica, on which the genus initially rested, and G. brandegeei sensu lato, is to be found in the fruiting bracteoles, which in Z. arizonica are mainly four-ridged and narrowly two-winged, and accommodate the mostly horizontal arrangement of the achenes.

Inclusion of Grayia brandegeei, with its laterally flattened (rarely three- or four-winged) fruiting bracteoles and vertical achenes, within Zuckia, with its architecturally differing bracteoles, requires explanation. At first glance, the six-ribbed bracteoles around a horizontal achene appear to be both distinctive and diagnostic. The bracteoles, however, merely accommodate the shape of the achene, and what is apparently very distinctive is only a structural modification. Examination of the fruiting bracteoles of all taxa previously included within Grayia brandegeei demonstrates existence of a rather wide array of bracteole morphology. Bracteoles are typically laterally compressed and samaralike. However, even on the same individual plant, there occur three-winged bracteoles, and in some bracteoles there are evident veins on one or both surfaces. The veins appear in the same position as the lateral ribs on the transversely flattened bracteoles of the arizonica phase of the species. There is also considerable variation in the morphology of the transversely flattened bracteoles, ranging from merely oval with a hint of the wings, the lateral ribs lacking altogether, to very definitely winged and with two or more lateral ribs. The overall similarity of the plants, their growth habit, and their preference for fine-textured, saline, often seleniferous substrates rather narrowly confined within the Colorado Plateau indicates a rather close relationship best reflected in their alliance.

In spite of the similarity of substrate inhabited by the three taxa, their geographic ranges are mainly discrete. This is due at least partially to the autecological differences in the habitats they occupy, and to actual geographical separation, even though apparently sympatric when mapped.

(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.)

Distribution
from FNA
AZ; CO; NM; UT; WY
[WildflowerSearch map]
Worldwide; especially in desert and semidesert regions; often in alkaline or saline habitats
Parent taxa Chenopodiaceae > Zuckia
Subordinate taxa
Z. brandegeei var. arizonica, Z. brandegeei var. brandegeei, Z. brandegeei var. plummeri
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
Key
1.Fruiting bracteoles without or with (1-)2(-4) prominent and without or with 4-5 smaller crests or ridges; achenes usually horizontalZ. brandegeei var. arizonica
1.Fruiting bracteoles 2(-4)-winged, samaralike; achenes vertical→ 2
2.Leaves usually wider than 6 mmZ. brandegeei var. plummeri
2.Leaves usually narrower than 6 mmZ. brandegeei var. brandegeei
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 alternateAllenrolfea
2.Leaves opposite→ 3
3.Plants annual herbs; all stems terminated by an inflorescenceSalicornia
3.Plants perennial shrubs; many stems entirely vegetative→ 4
4.Flowers distinct in each cyme, not adnate; outer seed coat hard, tuberculateArthrocnemum
4.Flowers of each cyme adnate to distal branch of inflorescence; outer seed coat thin, covered with hooked or straight hairsSarcocornia
5.Leaves opposite; perianth strongly imbricateNitrophila
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 bristlePolycnemum
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 tippedSalsola
8.Fruiting perianth apically winged; leaves terete, fleshy-succulent, bristle- tipped; bracts of inflorescence similar to leavesHalogeton
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 axillarySarcobatus
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 herbaceousKochia
11.Fruiting perianth without wings or tubercules, unchanged from flowering peri- anth; leaves fleshySuaeda
12.Plants stellate-pubescent annual herbsAxyris
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, subshrubsKrascheninnikovia
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 herbsSpinacia
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 annualSuckleya
17.Bracts united above middle, not vertically keeled; plants pe- rennial or annualAtriplex
18.Shrubs with divaricate, often spinescent branches; bracts with margins thickened, spongy within; pubescence of branched hairsGrayia
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 hairsZuckia
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 maturityCycloloma
20.Leaf blade margins entire; plants usually glabrous, sometimes slightly pubescentKochia
21.Proximal leaves petiolate, middle ones merely sessile, distal ones cordate-clasping; style branches 3Aphanisma
21.Leaves not as above; style branches mainly 2→ 22
22.Perianth segments each armed with spiniform, hooked, or conic appendagesBassia
22.Perianth segments rounded or keeled abaxially, lacking wings or spines→ 23
23.Ovary partly inferior; plants cultivated, rarely naturalizedBeta
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 hairsDysphania
25.Plants farinose or glabrousChenopodium
26.Plants dichotomously branched, ultimate branches filiform; leaves oblong, succulentMicromonolepis
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 habitatsMonolepis
27.Leaves lanceolate, linear-lanceolate, or filiform; perianth segments 1-3; stamens 1-3(-5); plants of sandy, low elevation sites.Corispermum
Synonyms Grayia brandegeei, Atriplex brandegeei
Name authority (A. Gray) S. L. Welsh & Stutz: Great Basin Naturalist 44: 208. (1984) Ventenat
Source Flora of North America vol. 4, p. 304. Flora of North America vol. 4, p. 258.
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