Atriplex dioica |
Chenopodiaceae |
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
rillscale, saline orache, saline saltbush, spike saltbush, thick-leaf orache, thickleaf orach, thinleaf orache |
goosefoot family |
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Habit | Herbs, monoecious, typically erect and often branching, 3–15 dm; stems green or striped, angular. | 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 | opposite or subopposite at least proximally; petiole 1–3 cm; blade strongly 3-veined from near base, lanceolate to linear-lanceolate or often triangular-ovate or lance-ovate, 30–125 × 25–60(–80) mm, thickened, basal or subbasal lobes spreading to mainly antrorse, obtuse, blade otherwise entire to sparingly dentate, typically scurfy. |
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 | in terminal or terminal on lateral branches, in spiciform spikes, 2–9 cm, naked except at base. |
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. |
||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Ovules | usually 1, campylotropous, bitegmic, crassinucellate. |
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Seeds | ellipsoid, wider than long, dimorphic: brown, 1.5–3 mm wide, flattened at base, or black, 1–2 mm wide, shiny; radicle inferior. |
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 | bracteoles green, blackening in age, sessile, veined or veins obscure, broadly triangular to ovate, 3–10 mm, thick, usually longer than wide, base truncate to obtuse, margin united at base, apex acute lateral angles entire or with short, sharp teeth, faces smooth or with 2 tubercles, with an inflated inner spongy layer. |
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 | = 36, 54. |
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Atriplex dioica |
Chenopodiaceae |
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Phenology | Flowering summer–fall. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Habitat | Sea beaches, other saline habitats | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Elevation | 0-1500 m (0-4900 ft) | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Distribution |
AZ; CA; CO; IA; ID; KS; MA; ME; MN; MT; ND; NE; NH; NJ; NM; NV; OK; OR; RI; SD; UT; WA; WY; AB; BC; MB; NF; NS; NU; ON; QC; SK; YT
|
Worldwide; especially in desert and semidesert regions; often in alkaline or saline habitats |
||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Discussion | The distribution of the species is evidently bipartite, with an eastern coastal series extending northward mainly from New Jersey to Newfoundland and along the St. Lawrence seaway, and perhaps extending to James Bay of Hudson Bay. The western grouping lies mainly west of the 95th meridian of longitude, where it has been collected since early historical times to the present in saline marshes or other saline sites from the Yukon Territory and Northwest Territories, southward to southern California, northern Arizona, northern New Mexico, and Oklahoma. Rafinesque gave the following information: “Stem upright angular branched, leaves petiolate, deltoid, acute, thick, scaly, the proximal opposite toothed, the distal alternate, hastated, entire: flowers dioical glomerate, male spiked naked, female unequal, sepals deltoid, warty-crested.” The name Atriplex dioica Rafinesque antedates Chenopodium subspicatum Nuttall by half a year, being published in December 1817. Hence, it is the correct name for the widely ranging species, which has passed most recently under the name A. subspicata. Nuttall’s description of the habitat of Chenopodium subspicatum is: “In saline soils around Mandan Village, Missouri,” a designation of habitat that applies to this day. The species forms a mirror-image set of specimens with the remarkably similar Atriplex prostrata, from which it may be distinguished in most cases by the thickened, merely ovate to lanceolate leaf blades, and less commonly but in some localities exclusively triangular-hastate to lanceolate, mostly scurfy and prominently 3-veined leaf blades. In some specimens, including the types of both Chenopodium subspicatum and Atriplex carnosa, the blades bear a hastate lobe at or above the base and sometimes match triangular-hastate profile of A. prostrata. The leaves of A. prostrata are typically thin-textured, green, not scurfy, and the veins of the blade are obscure. I. J. Bassett et al. (1983) disallowed within A. dioica (as A. subspicata) any but those with lanceolate blades, including those with the proximalmost leaves with a pair of subbasal hastate lobes. However, there are numerous specimens in which the blades are thickened and transitional in that regard to the triangular-hastate profile as in A. prostrata. Certainly those specimens with triangular or triangular-hastate leaves taken prior to the introduction of A. prostrata sometime late in the nineteenth or early twentieth century, clearly belong to the indigenous A. dioica. Whether there are intermediates between diploid (2n = 18) A. prostrata and tetraploid or hexaploid (2n = 36, 54) A. dioica is not known. There does not seem to be any consistent feature or combination of features by which all specimens can be assigned to one or the other of the two taxa. It seems probable, however, that A. prostrata is a late introduction from Europe, and that it, along with the related A. heterosperma, is now invading habitats previously occupied exclusively by the indigenous A. dioica. (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 |
|
|||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Source | FNA vol. 4. | FNA vol. 4, p. 258. | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Parent taxa | Chenopodiaceae > Atriplex > subg. Atriplex > sect. Teutliopsis | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Sibling taxa | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Subordinate taxa | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Synonyms | A. patula var. subspicata, A. subspicata | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Name authority | Rafinesque: Amer. Monthly Mag. & Crit. Rev. 2(2): 119. (1817) | Ventenat | ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Web links |
|