Southwestern

Western

Great Plains

Northern

Intermountain

C a olo o r

B

ad

m

P t lateau/Gr

a

e asi l n/Co u bi Basin

United States

Department

of Agriculture

Forest Service

Rocky Mountain

Research Station

General Technical

Report RMRS-GTR-125

February 2004

Fourwing Saltbush (Atriplex canescens)

Seed Transfer Zones

Stewart C. Sanderson and E. Durant McArthur
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Sanderson, Stewart C., and McArthur, E. Durant. 2004. Fourwing saltbush (Atriplex canescens) seed 

transfer zones. Gen. Tech. Rep. RMRS-GTR-125. Fort Collins, CO: U.S. Department of Agriculture, Forest 

Service, Rocky Mountain Research Station. 10 p.

Abstract

Atriplex canescens (Pursh.) Nutt. is the most widespread species of perennial Atriplex in North America. 

Throughout its distributional range, A. canescens shows considerable between-population variation. Some of 

this variation may be due to phenotypic plasticity but most of it appears to be genetic. Mutations, polyploidy, introgressive hybridization, and segregation from interspecific hybrids all appear to have contributed to its extensive heritable variation. Polyploidy is unusually common with numerous chromosome races (2x, 4x, 6x, 8x, 10x, 

12x, 14x, 20x).

Fourwing saltbush is widely used for reclamation plantings. Proper identification is important to the utilization 

of fourwing saltbush in such plantings. While many of the races have been formally named as varieties, others 

have not. Even though differentiated by ploidy, chemical constituents, geographic distribution, and statistical distribution of morphological characters, races may lack sufficient diagnostic characters to allow facile identification, 

at least in the herbarium. Rather than combining unnamed races under those that do have a taxonomic name, 

it seems better at present not to use the formal infraspecific categories in treating the fourwing saltbushes, but 

to consider them all as races. Seed transfer should be within the geographical distribution limits of each race. 

The most common race, by far, is Occidentalis. We recommend four overlapping seed transfer zones for race 

Occidentalis in the United States: (1) Northern Intermountain, (2) Western Great Plains, (3) Colorado Plateau/

Great Basin/Columbia Basin, and (4) Southwestern. Source seed populations from near the planting sites generally do well; and populations generally perform better when moved south and/or to lower elevations than when 

moved north and/or up in elevation.

The Authors

Stewart C. Sanderson is a research geneticist and E. Durant McArthur is a research geneticist and project 

leader of the Shrubland Biology and Restoration Research Work Unit, Shrub Sciences Laboratory, Provo, Utah. 

Dr. Sanderson has been at the Shrub Sciences Laboratory since 1981; he received B.S. and M.S. degrees 

from Brigham Young University, a Ph.D. from the University of Texas at Austin, and postdoctoral training at the 

University of Durham, England. Dr. McArthur has been with Forest Service Research since 1972 (at the Shrub 

Sciences Laboratory since 1975 and project leader since 1983); his B.S., M.S., and Ph.D. degrees are from the 

University of Utah. He spent a postdoctoral sojourn at the University of Leeds, England.
Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Application of Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Planting Recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Selected Germplasms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Acknowledgments

This work was stimulated and partially funded by the USDI Bureau of Land Management’s Great Basin Native 

Plant Selection and Increase Project and by the USDA Forest Service’s National Fire Plan. It draws from work initiated many years ago by our colleagues A. Perry Plummer of the Intermountain Forest and Range Experiment 

Station (now Rocky Mountain Research Station) and Howard C. Stutz of Brigham Young University. We appreciate review comments from Mike Pellant, Dan Ogle, Loren St. John, Nancy Shaw, and Howard Stutz on earlier 

versions of the manuscript.

USDA Forest Service RMRS-GTR-125. 2004 1

Introduction

Fourwing saltbush, Atriplex canescens (Pursh.) 

Nutt., is an important shrub for revegetation plantings 

(Plummer and others 1966; Blauer and others 1976; 

Plummer 1984). Its uses include mine land rehabilitation (Aldon 1981), shrub-herb interplantings for animal 

forage (Pendery and Provenza 1987; Rumbaugh and 

others 1982), fire rehabilitation plantings (Ott and others 2003), rangeland improvement (Monsen and Shaw 

1995), and stock for nutrient-poor soils (Glenn and others 1996, 1998). It is widespread in North America, 

being found from southern San Luis Potosi, Mexico, to 

southern Alberta and Saskatchewan, Canada, and from 

the Pacific Coast of California and Baja California to 

Texas, Oklahoma, Nebraska, Kansas, and the Dakotas 

(figures 1-5). It is a quality forage plant. Not only does it 

have fine nutritive quality but it is palatable to wild and 

domestic animals and tolerates browsing (Cibils and others 1998; Maywald and others 1998; Welch and Monsen 

1981, 1984). It shows abundant variation in response to 

seasons and to rainfall, as well as genetic differentiation. 

This has resulted in increasing confusion, even by reputable authors (see for example Taylor and Wilkens 1993 

who do not recognize the extent of subspecific variation). Fourwing saltbush is widely used for reclamation 

and restoration plantings. Over 100,000 pounds of seed 

is sold annually (McArthur and Young 1999). Last year, 

one seed company alone sold 70,000 pounds (McArthur 

and Sanderson, personal communication). With this 

much seed in commerce, land management agencies 

and seed vendors need guidance in planting. This paper 

is intended to meet that need.

The species has been planted successfully not only 

in restoration habitats, i.e., sites on which it naturally 

occurs, but also on sites in which it does not naturally 

Figure 1. Distribution of tetraploid races of fourwing saltbush. Occidentalis and Aptera overlap extensively in Wyoming.
2 USDA Forest Service RMRS-GTR-125. 2004 

Figure 2. Distribution 

of hexaploid races 

of fourwing saltbush.

Figure 3. Distribution of 

diploid races of fourwing saltbush.
USDA Forest Service RMRS-GTR-125. 2004 3

Figure 4. Distribution 

of races that are 

octoploid and higher in Nevada and 

in the Mojave and 

Sonoran deserts.

Figure 5. Distribution of races 

that are hexaploid and higher 

in the Mexican 

portion of the 

Chihuahuan 

Desert.
4 USDA Forest Service RMRS-GTR-125. 2004 

Table 1. Distinguishing characteristics of races of fourwing saltbush.

Race Bush height Leaves Four-winged fruits Habitata

Angustifolia (2x) 3 to 6 ft Narrow to very Unusually large Sand (U.S.), other soils 

 narrow, often long (Mexico)

Aptera (4x) 1 to 4 ft Moderately wide, Wings often small Badlands and grasslands

 with curving margins and irregular 

Brevis (2x) 1 to 2 ft, a few 3 ft Narrow to very Size normal to Grasslands on old lava flows

 narrow, shorter somewhat small

Diegensis (6x) 3 ft to as much Moderately wide, Normal size and Sea coast, ravines, canyons

 as 6 ft often green colored shape

Garrettii (2x) Usually 1 ft or less Leaves roundish, Normal Shale slopes, often southwest

 often yellow-green facing

Gigantea (2x) 4 to 8 ft, often Not very narrow, Unusually large Active dunes

 partly buried but often long

Grandidentatum (20x) 3 to 6 ft Bluish colored, small Strongly toothed, Sandy coastal areas 

 to very large, ends can be large near 

 often rounded the beach

Laciniata (8x) 6 to 8 ft Small to long Wings strongly Bottomlands

 and wide toothed

Laciniata (12x) 4 to 6 ft Extremely small Wings with many Sandy desert soils

 to large teeth and cuts

Laciniata (14x) 4 to 6 ft Short, or large Deeply toothed Coastal dunes

 and long. Bluish

Linearis (2x) Usually 1 to 2 ft Usually very narrow, Wings very small Bottomlands, may be 

 bluish, the sides (1/8 in), the body seasonally flooded

 straight may be swollen

Navajoensis (4x) One to 2 ft Shortened and Normal Red or blue shale 

 exceptionally wide

Nevadensis (6x) 1 to 4 ft, Wider than 4x at the Usually more Desert sands and slopes

 rarely more same season toothed than 4x

Obtusifolia (8x) 1 to 2 ft Egg-shaped with a Wings small, Alkaline bottomlands

 blunt end, greenish toothed

 gray

Occidentalis (4x) 2 to 6 ft Small to large, May be toothed Valleys and hills, sometimes 

 narrower than 6x on sand

Piochensis (2x) 1 to 6 ft Narrow to very Normal in size to Hills, commonly with juniper

 narrow rather large

Prosopidium (8x) 1 to 4 ft Short and wide, Wings small, Valleys

 bluish toothed

Prosopidium (10x) 1 to 4 ft Short and wide, Wings small, Valleys

 bluish toothed

Prostrata (6x) 1 to several Moderately long Normal Alkaline valley bottoms

 inches, flat 

 topped

Rachelensis (6x) 1 to 5 ft high Wider than 4x Smallish, toothed Valley

Rachelensis (8x) 1 to 2 ft high Very gray Smallish, toothed Alkaline valley bottoms

Vallis (6x) Usually 1 to Wider than 4x Normal to smallish Bottomlands

 3 ft high, larger 

 in West

a These specific type sites need to be combined with the general race distribution as shown in figures 1-5.
USDA Forest Service RMRS-GTR-125. 2004 5

occur, e.g., Wyoming big sagebrush habitats and pinyon-juniper habitats (Monsen and Shaw 1995; Ott and 

others 2003). It has also been used with success in the 

Mediterranean Basin and western and southern Asia 

(Belal and others 1993; Hyder 1981; Khalii and others 1986; Le Houérou 1992, 2000; Sankary and Goodin 

1986; Thompson and others 1998).

Fourwing saltbush includes populations with different 

chromosome numbers that are distributed in distinct geographical patterns. Sanderson and Stutz (2001) described 

and characterized this variation as chromosome number 

or chromosomal races based on previously described 

formal taxonomic designations and geographical, morphological, chemical, and cytogenetic information that 

they presented (table 1). The racial names in fourwing 

saltbush are derived, for the most part, from previously 

applied formal specific or varietal names but some racial 

names come from geographical or morphological traits 

of races that had not been formally characterized prior 

to the description of chromosomal races by Sanderson 

and Stutz (2001). This chromosomal race knowledge is 

useful in clarifying the limits of the species and its subspecific and population variation. Chromosome number 

races range from 2x to 20x—2x, 4x, 6x, 8x, 10x, 12x, 

14x, 20x (Dunford 1984, 1985; Sanderson and Stutz 

1994, 2001; Stutz and others 1975; Stutz and Sanderson 

1979)—and apparently account for much of the existing genetic differentiation. By far the most widespread 

of the races is the tetraploid Occidentalis (figures 1-5). 

Fourwing saltbush’s dioecious or trioecious outbreeding 

fosters genetic variation and contributes to the species’ 

successful occupation of diverse habitats (Freeman and 

others 1993; McArthur and others 1992). The reason 

the widespread race is called Occidentalis rather than 

“Canescens” is that the variety name canescens has to 

be given to plants from the type locality (site from which 

the species was first collected and described). The type 

locality for fourwing saltbush is a badland area along the 

banks of the Missouri River Big Bend in South Dakota, 

where the plant was collected by the explorers Lewis 

and Clark and described by Frederick Pursh. The type 

specimen, and plants growing there now, have relatively 

wide leaves and belong to race Aptera (4x), or perhaps 

to their own separate race because they show abundant 

sprouting from the roots (in figure 1 they are shown in 

the distribution map with Aptera).

 Fourwing saltbush plants from different areas or 

belonging to different races have been found to vary 

in tolerance to environmental factors (Glenn and others 1994, 1996; McArthur and others 1983; Mikhiel 

and others 1992; Plummer and others 1968; Van Epps 

1975). Especially important from the point of view of 

reclamation in cold desert settings is variation for tolerance to low temperatures (Van Epps 1975; Aberdeen 

Plant Materials Center staff 2002; USDA-NRCS and 

University of Idaho staff 2001).

Materials and Methods

Chromosome counts were accumulated over a period of 30 years in the laboratory of Howard C. Stutz, 

now professor emeritus, Brigham Young University in 

cooperation with the U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station, 

Shrub Sciences Laboratory. Recently, a ploidy analyzer instrument (Partec GmbH, Münster, Germany) has 

been obtained by the Shrub Sciences Laboratory, Provo, 

Utah, that facilitates the chromosome count work.

Meiotic chromosome counts were made from male 

flower buds preserved in the cold (-20 ̊ F) in vinegar or 

5% acetic acid. Pollen mother cells were squashed on 

a microscope slide and stained with saturated acetocarmine dye (Sanderson and Stutz 1994). More recently, 

ploidy levels were also determined on the ploidy analyzer using a DAPI stain for DNA and plants of known 

ploidy for comparison (Koontz 2001).

 Flavonoid chemistry was found to be useful in differentiating some races, due to presence or absence of 

6-methoxy flavonols, 3-O methylated flavonols, and 

the flavone tricin. Flavonoid aglycones were obtained 

from glycosides by acid hydrolysis with 1 N HCl, separated by paper chromatography, and examined over a 

black light (longwave ultraviolet) both before and after 

treatment with 5% aqueous aluminum chloride solution 

(Sanderson and others 1999). We have also examined 

isozymes to characterize the genetic variation of fourwing saltbush populations (McArthur and others 1986).

We call on our experience and reports in the literature in planting, transplanting, and survival of fourwing 

saltbush seed source populations taken together with the 

genetic and chemical variation data cited in the previous 

paragraphs to make recommendations for seed transfer 

for revegetation plantings for this important reclamation 

species.

Application of Results

Knowledge of chromosome numbers and flavonoid 

chemistry proved invaluable in understanding the variation observed in fourwing saltbush. Previously described 

varieties were validated and new races were discovered.
6 USDA Forest Service RMRS-GTR-125. 2004 

Fourwing saltbush is adaptable to seasonal drought 

and the leaves vary in size from large to small depending on growth status and moisture availability. This 

is particularly true of the high polyploids. Plants of 

Grandidentatum (20x) normally have small leaves of 

a bluish color, but when we grew plants from seed in 

a barrel of sand, the juvenile plants had leaves about 3 

inches long and 1 inch wide that looked somewhat like 

peach leaves. In spite of strong foliage variability, differences in leaf characters must often be relied upon in 

distinguishing the different races (table 1). There is a 

tendency for leaf width to increase with ploidy, at least 

at the lower ploidy levels. This is the most useful characteristic in distinguishing diploids from tetraploids. With 

the notable exception of race Garrettii (A. garrettii), 

diploids have very narrow leaves that are 10 times or 

more longer than broad, although they vary seasonally to some degree. With care, tetraploids can also 

be distinguished from hexaploids by leaf width alone, 

but plants have to be compared in the field and at the 

same time of year. Leaf color is also useful in some 

cases. The diploids, except for race Linearis with rather 

bluish gray-green leaves (sometimes blackish in herbarium specimens), all tend to have more yellowish foliage 

during dry seasons and winter due to chlorophyll loss, 

which has been found to prevent damage from excess 

light energy during times when the plant is not growing (Munné-Bosch and Alegre 2000). Yellowing may be 

seen slightly in tetraploid plants also, but not usually in 

higher polyploids. Like Linearis, several Mexican races 

of higher ploidy also have a bluish leaf color.

A problem with the identification of the diploids 

Angustifolia, Brevis, and Piochensis is the occurrence 

of hybrid intermediates between these and local plants 

of the tetraploid race Occidentalis. Hybrid intermediates are usually also tetraploid, and therefore interbreed 

with Occidentalis but not with their parental diploid. As 

a result the diploids remain unchanged and pure, but 

tetraploids can be somewhat mongrelized by the hybridization and begin to look more like diploids. Tetraploid 

plants in an area of past or present diploid occurrence 

are often sufficiently intermediate in form that it is difficult to tell from appearance alone where the diploid 

leaves off and the tetraploid begins. However, there 

are seasons when it is easier to distinguish them, and 

the tetraploids seldom become exactly like diploids in 

any characteristic. For instance, flavonoids, a kind of 

plant chemical, are different between diploids and the 

tetraploids of race Occidentalis (the latter produce 6-

methoxy flavonols and the former do not). In many cases 

that have been examined, hybrid derivatives always produce these compounds too.

Planting Recommendations

Many of the races are relatively localized (figures 1 to 

5). Many are also found in Mexico and have little ability to survive the winters of the temperate zone in the 

U.S. Therefore, they are not likely candidates for use in 

reclamation in this country. In localities where any of 

the distinctive less common races of fourwing saltbush 

are encountered, they alone should be used in reseeding. 

Ordinarily, however, it is the tetraploid Occidentalis, or 

sometimes Aptera, that are ordinarily used in reclamation or restoration plantings. When an appropriate seed 

source is chosen, the primary concerns are to seed each 

race in its own area of natural distribution and to be 

mindful of ecotypic variability of Occidentalis (see further discussion, below).

As might be expected, resistance to winter injury increases in a northward direction. Of the various Atriplex

species and races from warm desert areas or Mexico (and 

some from Australia) that were planted in gardens at 

Provo, Utah, over the years, none survived for long and 

most died during their first winter (Stutz and Sanderson, 

personal observations). Van Epps (1975) reported on 

winter survival of fourwing saltbush plants from a 

number of sites in the Intermountain area. There was 

variability, but generally plants from southern Utah and 

areas farther south suffered high mortality when grown 

in central Utah, and the best survival for this planting 

area was in plants obtained from Bridger in southern 

Montana. The Aberdeen, Idaho, Plant Materials Center 

has recently announced the release of a cold-adapted 

strain of fourwing saltbush hardier than strains presently in use (Aberdeen Plant Materials Center staff 2002; 

USDA-NRCS and University of Idaho staff 2001). It 

should be especially useful in colder areas, such as 

northern Nevada, northern Utah, and southern Idaho.

The race Aptera, mainly found in Montana (figure 1), 

is also very cold hardy. Its native habitat is saline soils 

such as are found in badlands, in areas of generally 

greater annual precipitation (>15 inches), especially 

summer rainfall, than is found over the general fourwing 

saltbush distributional range. Plants of Aptera from various locations were grown successfully for many years in 

gardens at Provo, Utah and at the Gordon Creek Wildlife 

Management Area near Price, Utah, where the annual 

rainfall is about 15 inches and 11.3 inches, respectively 

(Ashcroft and others 1992; McArthur and Welch 1982).

Plants from more northern areas generally perform 

well in areas to the south of their origin. On the other 

hand, plants from the south typically grow in northern 

areas with much vigor during summer but are eliminated during the cold season. Cold resistance probably 
USDA Forest Service RMRS-GTR-125. 2004 7

comes at a certain cost, and that is a reduced growth rate. 

Usually this is not an adaptive disadvantage. Long-term 

survival of seeded populations depends on successful recruitment by seeded plants. This may or may not 

happen depending of the site adaptation of the source 

population—it usually does not happen in off-site plantings, e.g., Wyoming big sagebrush sites (McArthur and 

Sanderson, personal observation). Seeded plants do 

produce viable seed in most locations (Noller and others 1984; Monsen and McArthur 1985; Sanderson and 

McArthur personal observations).

Our general recommendations for planting race 

Occidentalis are summarized in figure 6. Common garden plantings and revegetation plantings (Fitzsimmons 

and others 1998; Geist and Edgerton 1984; McArthur 

and others1983; Pendleton and others 1992; Peterson and 

others 1987; Plummer and others 1966; Van Epps 1975) 

point to a consensus that source populations should not 

be moved too far north or up in elevation. In general, 

source populations from near planting sites do well. So, 

we recommend that source populations not be obtained 

out of the general zones (figure 6) and not moved up 

in elevation more than 2,000 feet within their respective zones. The four zones are Northern Intermountain; 

Western Great Plains; Colorado Plateau/Great Basin/

Columbia Basin ; and Southwestern (figure 6).

Stutz and colleagues (Stutz 1982; Stutz and Estrada 

1995) recommend that revegetation be by natural evolutionary processes by seeding mixes of compatible seed 

sources (sources of the same ploidy level), thus stimulating an evolutionary trajectory for the specific seeded 

site. Should this be the management choice for revegetation, we recommend that the seed mix chosen include 

source populations from the zone to be rehabilitated.

Selected Germplasms

Five germplasms of fourwing saltbush have been 

released for reclamation plantings (Davis and others 

2002). Germplasm releases are plant materials that 

have been identified and depending on class (source 

identified, selected, tested, source identified, cultivar—

see Young 1995 and Young and others 1995 for 

criteria), evaluated for performance and made available for seed increase. Whereas there is a substantive 

continuing demand for fourwing saltbush seed for 

rehabilitation plantings (McArthur and Young 1999), 

most of this need is met by wildland collected seed. 

McArthur and others (1978) made an economic analysis on the potential profitability for growing fourwing 

saltbush in seed orchards. However, the ready availability 

Figure 6. Recommended seed transfer 

zones for race Occidentalis: (1) Northern 

Intermountain, (2) Western Great Plains, 

(3) Colorado Plateau/Great Basin/Columbia 

Basin, and (4) Southwestern.
8 USDA Forest Service RMRS-GTR-125. 2004 

of wildland-collected seed has dampened the demand 

for more expensive field-grown seed. Some commercial production, however, is available. The best way to 

determine if seed is available is to contact state Crop 

Improvement Associations or the plant materials specialists of the U. S. Department of Agriculture, Natural 

Resources Conservation Service.

The released germplasms are ‘Marana,’ ‘Rincon,’ 

‘Santa Rita,’ “Snake River Plains,” and ‘Wytana’ (Davis 

and others 2002). All but “Snake River Plains” were 

released as cultivars, implying that the indigenous population sources underwent selection for specific trait(s) 

and were tested in comparisons with other fourwing saltbush source populations. “Snake River Plains” is a tested 

plant material comprising a synthetic composite of four 

populations. The released plant materials are all of race 

Occidentalis except ‘Wytana’ which is of race Aptera. 

Both ‘Marana’ (originally collected from El Cajon, San 

Diego County, California at an elevation of 600 feet) 

and ‘Santa Rita’ (originally from south of Tucson, Pima 

County, Arizona at an elevation of 2400 feet) are from 

southern locations at relatively low altitudes and are best 

adapted to the Mojave and Sonoran Deserts, respectively (Davis and others 2002; Cable 1972; United States 

Department of Agriculture Soil Conservation Service 

and others 1980). The source population for ‘Rincon’ 

is at Rincon Blanco, Rio Arriba County, New Mexico, 

at 7,800 feet. This release was a result of selection and 

testing over a wide geographical area (McArthur and 

others 1984). The source high elevation site (ponderosa pine, pinyon-juniper, and big sagebrush community) 

preadapted this release to more northern habitats (as far 

north as central Utah and Nevada) than might be expected. The “Snake River Plains” tested germplasm was 

selected for its tolerance to the colder, windier northern conditions of Snake River Plain and Northern Great 

Basin areas (USDA-NRCS and University of Idaho 

staff, 2001). ‘Wytana’ is adapted not only to the Aptera 

geographical area (Fig. 1) but also to somewhat more 

southern areas (Sanderson and McArthur, personal observations).

We believe that each of the released plant materials should be considered for use in revegetation areas 

to which they are adapted but that wildland collected 

materials should be used prudently within their zones 

of adaptation (our recommendations are in Fig. 6). As 

a general rule, close habitat and geographical proximity 

are preferred. And, when greater movement from source 

sites occurs, down slope and south is better than upslope 

and north.

References

Aldon, Earl F. 1981. Long-term plant survival and density 

data from reclaimed Southwestern coal mine spoils. Great 

Basin Naturalist. 41: 271-273.

Aberdeen Plant Materials Center Staff. 2002. Fact sheet: 

Snake River Plains fourwing saltbush, selected class 

germplasm. Aberdeen, ID: USDA-NRCS, Aberdeen PMC, 

P.O. Box 296, Aberdeen, ID 83210. 1 p.

Ashcroft, Gaylen L.; Jensen, Donald T.; Brown, Jeffrey L. 

1992. Utah climate. Logan, UT: Utah Climate Center, Utah 

State University. 127 p.

Belal, A. H.; Rammah, A. M.; Hopkins, M. S.; Hansen, L. 

D.; McArthur, E. D. 1993. Studies of salt-tolerance and 

chemical analysis of fodder shrubs in Egypt and Utah 

(USA). In: Leith, H.; Al Masoom, A., eds. Towards 

the rational use of high salinity tolerant plants, Vol 2. 

Dordrecht, Netherlands: Kluwer Academic Publishers: 

213-220.

Blauer, A. Clyde; Plummer, A. Perry; McArthur, E. Durant; 

Stevens, Richard; Giunta, Bruce C. 1976. Characteristics 

and hybridization of important Intermountain shrubs. II. 

Chenopod family. Res. Pap. INT-177. Ogden, UT: U.S. 

Department Agriculture, Forest Service, Intermountain 

Forest and Range Experiment Station. 42 p.

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in southern Arizona. Journal of Range Management. 25: 

150-153.

Cibils, Andrés F.; Swift, David M.; McArthur, E. Durant. 

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of knowledge. Gen. Tech. Rep. RMRS-GTR-14. Ogden, 

UT: U.S. Department of Agriculture, Forest Service, 

Rocky Mountain Research Station. 31 p.

Davis, Kathleen, M.; Englert, John M.; Kujawski, Jennifer 

L. 2002. Improved conservation plant materials released 

by NRCS and cooperators through September 2002. 

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Resources Conservation Service. 57 p.

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adjacent Texas. Southwestern Naturalist. 29: 223-228.

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(Chenopodiaceae). Southwestern Naturalist. 30: 377-381.

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Stewart C.; Tiedemann, Arthur R. 1993. The influence 

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establishment in the Keating Uniform Garden—first 

year results. Res. Note PNW-416. Portland, OR: U.S. 

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Jody. 1998. Comparison of growth and morphology of 
USDA Forest Service RMRS-GTR-125. 2004 9

Atriplex canescens varieties occidentalis and angustifolia. 

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1994. How much sodium accumulation is necessary for 

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and salt tolerance of Atriplex canescens (Chenopodiaceae) 

genotypes. American Journal of Botany. 83: 997-1005.

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performance of some exotic species of Atriplex in Saudi 

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