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WOODYCHENOPODSUSEFULFOR
RANGELAND RECLAMATION IN
WESTERN NORTH AMERICA
Stewart C. Sanderson
Howard C. Stutz
gardneri. From these parents it obtains a tendency to
produce four-winged fruits and also an ability to rootsprout. All of the genetic variability from the parents is
combined in this variable species. It occurs in the northwestern Great Plains, often in badlands, but where effective precipitation may be somewhat greater than in valleys of the Intermountain area.
Atriplex canescens (fourwing saltbush) (2x-20x; 2x-6x
in the Intermountain area) is usually a medium to large
shrub. It has numerous chromosome races, mostly found
in the warm deserts of the southwestern United States
and northern Mexico (Sanderson and Stutz, in review;
Stutz and Sanderson 1993). Because of the chromosomal
diversity, care should be taken to select seed sources of like
chromosome numbers to avoid the development of chromosomal sterility in later generations. Many of the chromosomal races of fourwing saltbush are described as species or varieties. Some of these are limited in nature to
sand dunes; others are found both on sand and heavy
soils. Leaf length of fourwing saltbush races tends to
decrease while leaf width may increase somewhat with
ploidy.
Fourwing saltbush is able to resprout when browsed or
burned (Tirmenstein 1986a). Rare, highly rootsprouting
individual plants have been found that are also able to
spread extensively by vegetative means (Barrow, in review). Such plants, if frost hardy, seem to have promise
in annual grass remediation. In the Intermountain area
diploid, tetraploid, hexaploid, and possibly a few octoploid
fourwing saltbush populations are found. The most abundant are tetraploids, which grow in all of the western
States and on a variety of soil types.
There are three diploid fourwings in the Intermountain
area. Atriplex canescens var. gigantea is a large bush
with unusually long leaves, growing only on the Little
Sahara Dunes in western Utah. Var. angustifolia, identified by its very narrow leaves, is another large bush growing on dunes. It grows chiefly in Mexico, but is also found
in southern Arizona and New Mexico. In New Mexico
there are some roadside plants as far north as Farmington.
The third diploid is A. garrettii, or A. canescens var.
garrettii, a small yellowish-green bush with round or oval
leaves, found only along the Colorado River in southeastem Utah.
Tetraploid fourwing saltbush occupies a variety of habitats. Leaves of the tetraploid are shorter than those of
vars. gigantea and angustifolia. 'Rincon,' from northern
New Mexico, and 'Marana,' from southern Arizona, are
released cultivars of tetraploid fourwing saltbush.
ABSTRACT
A number of woody and semiwoody chenopods are potententially useful in annual grassland reclamation. Ceratoides lanata, Grayia spinosa, Atriplex canescens, and
other species of Atriplex, especially those showing the potential of fire tolerance through an ability to rootsprout
or crownsprout, may be valuable in regulating cheatgrass.
INTRODUCTION
Members of the chenopod family tend to tolerate high
soil salinity and drought, permitting them to grow in
places where competition from other plants is limited.
Many woody chenopod species are equipped with C" photosynthesis, which is advantageous for growth in deserts
and summer heat.
Some chenopods defend themselves from mammalian
and insect herbivores with spines or thorns. Members of
the family also may protect themselves by accumulating
unpalatable or toxic substances such as oxalates, salts, or
triterpene saponins. However, these are harmful only under particular conditions and in particular species (Ihnat
1989; Oakenfull and Sidhu 1989; Young and James 1988).
They do not detract from the forage value of the family as
a whole.
Chenopods, due to their overall benign defensive syndrome and high protein content, tend to be very desirable
browse plants. This is fortunate because in the salt deserts
of western North America they constitute almost the only
vegetation on millions of acres of rangeland. This paper
outlines a number of the more valuable chenopod species.
Tables 1 and 2list species' characteristics.
ATRIPLEX SPECIES
Atriplex aptera (aptera fourwing) (4x) is more appropriately classified as a variety of A. canescens, where it becomes the type variety, var. canescens, as explained elsewhere (Sanderson and Stutz, in review). It is a highly
variable hybrid of tetraploid fourwing saltbush and A
Paper presented at the Symposium on Ecology, Management, and Restoration of Intermountain Annual Rangelands, Boise, ID, May 18-22, 1992.
Stewart C. Sanderson is Plant Geneticist, Shrub Sciences Laboratory,
Intermountain Research Station, Forest Service, U.S. Department of
Agriculture, Provo, UT 84606, and Research Associate, Department of
Botany and Range Science, Brigham Young University, Provo, UT 84602.
Howard C. Stutz is Professor Emeritus, Department of Botany and Range
Science, Brigham Young University, Provo, UT 84602.
374
Table 1-Characteristics of selected species of North American perennial Atriplex
Taxon
A. canescens
A. confertifolia
A.aptera
A.gardneri
A. falcata
A. tridentata
Hybrids with
A. canescens
Ploidy
Forage use
Soil salinity
Rootsprouting
Remediation potential
2x-20x
2x-10x
4x
2x,4x
2x
4x,6x
palatable
spiny
palatable
palatable
palatable
palatable
medium
medium to higher
medium to higher
medium to higher
low to higher
high
crown sprouts
no
yes
yes
yes
yes
good
usually poor
moderately good
moderately good
good
specialized sites only
4x, 6x
palatable
low medium
yes
good
Populations of tetraploids contain occasional plants that
are hexaploid; occasionally entire populations of hexaploids are found. As shown in Stutz and Sanderson (1979),
hexaploids are quite common in Arizona, New Mexico,
and Nevada. In addition there is a small hexaploid population growing at Grantsville, UT, which we called
"tooelensis." It lacks saponins, has furfuraceous foliage,
and flowers very late (mostly in September). The hexaploid race in western Nevada has a distinctive flavonoid
pattern compared to other tetraploids or hexaploids, and
because it is shorter in stature, is referred to as "nana."
It seems that polyploids have an advantage in being able
to grow in saltier soils (Sanderson and others 1987).
"Nana" grows in transitional areas between the warm
desert and cold desert.
Fourwing saltbush races and their ecotypes provide a
rich variety of useful genotypes that may prove helpful
in revegetation. In addition to those mentioned, there
are numerous others existing in the southwestern United
States and Mexico that would be useful in warmer climates.
Atriplex confertifolia (shadscale) (2x-10x) occurs from
Oregon and California to North Dakota and Texas, and
marginally into Mexico. It consists of numerous chemical
and chromosomal races (Sanderson and others 1990;
Stutz and Sanderson 1983). For this reason, as with A
canescens, care must be taken in selecting seed sources of
shadscale so that incompatible chromosome races are not
interplanted. Shadscale is usually a small- to mediumsized shrub. Although spring growth is palatable and nutritious and the plants are saponin free, young branches
rapidly develop into spines, reducing the accessibility of
forage. Shadscale palatability is lower at other times of
year (Dayton 1937). Shadscale is the Atriplex dominant
most often replaced by cheatgrass because, except for occasional restablishment from seed, the species does not
recover from fire. Unless cultivars with much more favorable characteristics can be developed, it seems unlikely
that shadscale will be of value in recovering cheatgrassinfested sites. It will remain important on sites where
annuals are not a problem.
Atriplex falcata (falcate saltbush) (2x) is a low-growing
subshrub that can grow on badland outcrops or on welldrained valley slopes. It is mainly found in the States of
Utah, Nevada, Idaho, and Oregon. The spring leaves are
linear, somewhat like those of fourwing saltbush, but
these are replaced by very small, persistent leaves in the
fall and winter. Falcate saltbush tends to exclude other
vegetation, forming monotypic stands. Even cheatgrass
may be excluded on more saline soils. Although predominantly caespitose, it sprouts readily from the crown and
roots when damaged, and most populations also occasionally vegetatively reproduce by roots prouts.
Because it forms small, isolated populations (suggesting
that larger, more continuous populations are unstable),
and has been observed to die out suddenly, disease problems may exist. Nevertheless, because of its resprouting
ability, A. falcata is a species that should be useful in fireaffected areas. Falcate saltbush can be distinguished
from A tridentata, which is vegetatively similar but occupies a different ecological setting, by the presence of bitter
saponins in the foliage and by the distinctive falcate
(curved) fruit beak (fig. 1).
Atriplex gardneri (A gardneri var. gardneri) (Gardner
saltbush) (2x, 4x) is a prostrate to upright, low shrub
found on saline and often barren clays, frequently growing
in large pure stands, mainly in Wyoming and Montana.
With age, plants spread into circular patches. It differs
from A falcata and A. tridentata in having roundish, oval
to cuneate or obcuneate leaves. Gardner saltbush resists
fire through low flammability and also through resprouting
(Tirmenstein 198Gb). Often found with grasses or sagebrush, it has at least a degree of competitive ability.
Atriplex gardneri is a species that merits experimentation
on cheatgrass sites.
a
b
Figure 1-Fruits of Atriplex tridentata (a, b)
and A. falcata (c, d). The seed is covered
by two clamshell-like bracts that have teeth
along their margins or a long beak. Both
species can have bumps or soft spines on
the sides of the fruiting bracts, but they are
more common in A. falcata.
375
c
d
Atriplex tridentata (trident saltbush) (mostly 6x) is a
strongly rootsprouting subshrub similar in appearance to
falcate saltbush. However, in contrast to that species, it
is free of bitter saponins and is found in highly saline bottomlands, often with Sarcobatus vermiculatus. Because
of abundant rootsprouting, it may spread into large clonal
patches. The plants have narrow, linear leaves like those
of falcate saltbush, but the fruits have three to seven marginal teeth rather than a long beak (fig. 1). As discussed
later, trident saltbush has a tendency to form hybrids and
hybrid derivative populations by crossing with fourwing
saltbush.
backcrossing to one of the parental species, these gametes
might have given rise to either tetraploid or hexaploid hybrid derivatives. However, those in nature that clearly
belong to this parental combination are, so far, all 2n = 54
(hexaploid). The numerous distinctive populations of
these hybrid derivatives provide a rich source of genetic
variation for use in reclamation.
SHRUBBY NON-ATRIPLEX
CHENOPODS
Allenrolfea occidentalis (iodine bush) is a succulent
shrub or subshrub with jointed green stems that grows
at the margins of salt marshes and on salt flats, from
Oregon and California to Texas and Mexico (Welsh and
others 1987). Great Basin plants are small shrubs to subshrubs, but plants from other places may be larger. Those
in California's Central Valley are strong, woody shrubs
several feet high, and perhaps deserve to be classified as
a separate species or variety. The consumption of its foliage by herbivores is limited by a high salt content. Because it is very halophilic and dependent on high water
tables, it does not grow to any extent in cheatgrass habitats, and is therefore not likely to be useful in remediation
of such areas.
Ceratoides lanata (winterfat) is a densely pubescent
shrub or subshrub occurring from southern Canada to
California and Texas. Winterfat often forms huge, pure
stands excluding other shrubs. It also grows withAtriplex
or Artemisia. Although its hairiness would be a deterrent
to insect attack (Smith 1989), the plant is eaten readily by
mammals and is an important source of browse. There
are two widespread forms of winterfat, a short-statured
one more northern in distribution, and a large one (for example, Hatch winterfat) mostly found in warm deserts.
Winterfat resprouts after fire (Holifield 1987a); however,
a solid stand of winterfat often partly excludes cheatgrass,
reducing the frequency of fire. It is readily established
from seed (McArthur and Monsen, in review).
Grayia brandegii (spineless hopsage) is palatable but
uncommon (Holifield 1987b), found only on steep, welldrained outcrops of highly saline and seleniferous shales
on the Colorado Plateau, mostly in eastern Utah but including portions of neighboring Wyoming, Colorado, New
Mexico, and Arizona. These shales are often brightly colored red, or white and purple, such as the Morrison Shale
formation. Usually, little cheatgrass is present, and it is
expected that the competitive ability of spineless hopsage
is low except on unusually saline soils.
Grayia spinosa (spiny hopsage), which occurs from
Washington to Montana and California to New Mexico,
is common in the upper elevations of the Mojave Desert.
It grows in valleys and hills of the Great Basin well into
the pinyon-juniper community, on soils of low to moderate
salinity. It is very palatable when bearing leaves but its
usefulness as a range plant is limited; it sheds leaves and
fruits in early summer (McArthur and Monsen, in review). Spiny hopsage resprouts readily after light burning (Holifield 1987c), but can sometimes be killed by tire.
Kochia americana (gray molly) occurs from Oregon to
Montana and from California to New Mexico. It is a small
subshrub that forms stands of rather widely spaced plants
ATRIPLEX HYBRIDS
Atriplex canescens x A. gardneri: Individual first generation plants having this evident parentage are common
where fourwing saltbush plants have established themselves along roadsides near Gardner saltbush populations. Hybrids are also found occasionally at considerable
distances from plants of fourwing saltbush, apparently
due to wind transport of the pollen. The hybrids are easy
to find because they tower above Gardner saltbush plants.
Although Gardner saltbush has both diploid and tetraploid populations, most of the hybridization occurs at the
tetraploid level. Because of genetic segregation among
the progeny of hybrids, hybrids can form a large source
of variation for natural or artificial selection. There are
also vast areas where populations of both parents have
come together in the past and formed populations of
aptera saltbush, a new and still developing species, as
already mentioned.
Atriplex cansecens xA falcata: These hybrids have been
formed synthetically in the garden, and a number of distinct native hybrid populations in the Great Basin are suggested to have this parentage (Hanson 1962). Evidence
has been presented (Stutz and others 1979), however, that
some of the populations in question are derived from the
hybridization of A canescens and A tridentata, as discussed later. Even if there are no hybrids in nature having
A falcata as a parent, interesting selections could be produced using synthetic hybrids as a source of variation.
Atriplex canescens x A tridentata: An odd-ploid seedling
with a chromosome count of 2n = 45 (pentaploid) (Pope
1976) was grown from seed collected from a lone female
A. canescens plant near Grantsville, UT, pollinated by distant male bushes. Other seedlings from the same plant
had counts of 2n = 36 (tetraploid), which was inferred to
be the chromosome number of the A. canescens plant. A
search of the site revealed numerous plants of A tridentata close at hand; these proved to have a higher chromosome number of 2n =54, which could have produced the
pentaploid seed by hybridization. Several adult hybrids
were found. Many other such areas of hybridization were
eventually encountered-huge populations that appeared
to have resulted from hybridization of these parents
(Stutz and others 1979).
Although odd-ploidy is unstable in sexually reproducing
organisms because most of the gametes are unbalanced, it
was found in this case to rapidly resolve itself, apparently
because euploid (balanced) gametes, which are occasionally produced, have a viability advantage. Through
376
Table 2-Characteristics of shrubby North American chenopod species
Taxon
Allenrolfea occidentalis
Ceratiodes /anata
Grayia brandegii
Grayia spinosa
Kochia americana
Sarcobatus baileyi
Sarcobatus vermiculatus
Suaedato"eyana
Zuckia arizonica
Ploidy
2x
2x
2x,4x
4x
2x
12x
4x, ax
2x
2x
Forage use
low
palatable
palatable
early decid.
palatable
spiny
low
low
palatable
Habitats
Halophyllcity
salt flats, marshes
valleys, hills
saline shale outcrops
valleys, hills
saline barrens & hills
arid slopes
saline bottomlands
saline bottoms & flats
saline clay, shale
in barren, highly saline areas such as valley bottoms. Because it spreads vegetatively by occasional root- sprouting, it should withstand fire well. However, its habitat
suggests that most ecotypes of gray molly would have
relatively little competitive ability against communities
of introduced annuals.
Sarcobatus baileyi (Bailey greasewood) is a densely
spiny, deciduous shrub, up to about one-half meter in
height, that forms solid stands on arid hillsides in westem Nevada and adjacent California. It often grows with
Atriplex confertifolia. Bailey greasewood's leaves are
fleshy and narrow, like those of Sarcobatus vermiculatus.
It can presumably resprout after fire like that species. It
occurs slightly below and to the south of major cheatgrass
zones, but because it is not dependent on water tables like
its congener, Bailey greasewood may be worth a trial with
cheatgrass.
Sarcobatus vermiculatus (black greasewood) is a larger
deciduous shrub with much variability, found from
Sonora to Alberta and California to Texas. It has tetraploid and octaploid races, but plants in the Intermountain
region are all octaploid. It grows mainly in saline bottomlands having high water tables. There are some populations occurring in soils where cheatgrass is able to grow
abundantly, and in such cases greasewood resprouts vigorously from its roots after the grass has been burned
(Tirmenstein 1987).
Suaeda torreyana (inkbush) occurs from California and
Wyoming to Mexico. It is a plant of more saline soils, often
growing in the presence of high water tables. The leaves
are narrow and fleshy, similar in form to those of Sarcobatus. They tum inky black upon dying in the fall. Inkbush seldom grows where annual grasses are abundant.
Zuckia arizonica (zuckia) is an uncommon plant of barren clay or shale in the Colorado Plateau, in eastern Utah
and northern Arizona. It hybridizes with Grayia brandegii
and is therefore related to it, though the fruits are dissimilar. Like that species, it is not found in dense stands
of grass.
very strong
weak to moderate
strong
weak
strong
moderate
moderate to strong
strong
strong
Apparent cheatgrass
remediation potential
none
very useful
low
probably good
low
moderate? should be tested
good for specialized sites
low
low
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