Paschke2003 PDF

land degradation & development
Land Degrad. Develop. 14: 459–480 (2003)
Published online 9 July 2003 in Wiley InterScience (www.interscience.wiley.com). DOI: 10.1002/ldr.568
BIOLOGY AND ESTABLISHMENT OF MOUNTAIN SHRUBS ON

MINING DISTURBANCES IN THE ROCKY MOUNTAINS, USA
M. W. PASCHKE,1* E. F. REDENTE1 AND S. L. BROWN2
1
Colorado State University, Department of Rangeland Ecosystem Science, Fort Collins, CO, USA
2
Colorado Department of Natural Resources, Division of Minerals and Geology, Denver, CO, USA
Received 28 January 2002; Revised 6 September 2002; Accepted 6 December 2002
ABSTRACT
The Rocky Mountains of the western United States contain many economically important natural resources. Increasing
development of these resources has lead to land degradation, which often requires restoration efforts. A common type of
disturbance in this region is mineral extraction and these activities often occur in zones of vegetation dominated by shrubs.
These mined lands have proven to be particularly challenging to restore to native shrub cover. Mountain shrubland species such
as big sagebrush (Artemisia tridentata), antelope bitterbrush (Purshia tridentata), Saskatoon serviceberry (Amelanchier
alnifolia), black chokecherry (Prunus virginiana), mountain mahogany (Cercocarpus montanus), and snowberry (Symphor-
icarpos sp.) are crucial species for wildlife in this region due to the seasonal dependence of many wildlife species on the
mountain shrubland zones. The development of successful restoration techniques for these key shrub species is therefore
important for the continued and sustainable utilization of natural resources in the Rocky Mountains.
The purpose of this literature review is to provide a reference to land managers working on woody plant establishment on
mining disturbances in mountain shrublands in the Rocky Mountains. This review covers the biology, ecology, and propagation
of six shrub species that are of primary importance for meeting regulatory compliance in this region. Based upon this survey of
the published literature, we conclude that successful establishment of these species has most often involved: (1) utilization of
local shrub ecotypes, varieties or subspecies in reclamation efforts, (2) protection from browsing during the establishment
phase, (3) strategies for avoiding herbaceous competition, and (4) providing a source of mutualistic soil organisms. Additional
specific recommendations for each of these species are discussed. Copyright # 2003 John Wiley & Sons, Ltd.
key words: mined land; seeding; shrubland restoration; topsoil; transplants; wildlife habitat
INTRODUCTION
There is a need for an increased knowledge of the biology and methods for establishment of woody plants on
disturbed lands in mountain shrub zones of the Rocky Mountains. This region contains many economically
important natural resources including minerals, wildlife, water, timber, forage, recreational opportunities, and
food. The development of these resources often leads to degradation of plant communities, which may require
restoration efforts. In the Rocky Mountains, where mineral extraction is an important part of the local economies,
the restoration of mountain shrub communities on mined lands is an important and difficult task.
Areas of coal extraction in the Rocky Mountains often occur in habitats of shrubby vegetation that are referred
to as ‘mountain brush’ or ‘mountain shrub’. This is an important habitat type in much of the area because big game
is concentrated in mountain shrublands during winter periods. The quality of these habitats is often the key
determinant of the ecosystem carrying capacity for big game population (Wallmo et al., 1976). Mountain
shrublands also serve as a habitat for a wider variety of wildlife including threatened Columbian sharp-tailed
Correspondence to: M. W. Paschke, Department of Forest, Rangeland and Watershed Stewardship, Colorado State University, Fort Collins, CO
80523-1478, USA.
E-mail: Mark.Paschke@colostate.edu
Copyright # 2003 John Wiley & Sons, Ltd.

460 M. W. PASCHKE ET AL.
grouse and sage grouse. Mountain shrubland species such as bitterbrush (Purshia tridentata), serviceberry
(Amelanchier sp.) and mountain mahogany (Cercocarpus montanus) are some of the most palatable North
American plant genera for big game, while sagebrush (Artemisia tridentata) is less palatable but among the most
important browse species due to its abundance and availability (Merrill, 1971).
The mountain shrub plant association is widespread in the Rocky Mountain region from western Colorado
through Utah and Nevada and from western Wyoming south to northern Arizona. Mountain shrublands typically
occur above valley and foothill sagebrush or pinyon-juniper communities and below montane conifer or aspen
(Populus tremuloides) forests. These mountain shrub communities are diverse and vary in species composition
with elevation, aspect, soils, and disturbance history. Despite the wide diversity of mountain shrub habitat types,
there are relatively few dominant shrub species found in this vegetation type. Notable among these common shrubs
are big sagebrush, antelope bitterbrush, mountain mahogany, Saskatoon serviceberry (Amelanchier alnifolia),
black chokecherry (Prunus virginiana), snowberry (Symphoricarpos sp.), maple (Acer sp.), and Gambel oak
(Quercus gambelii) (Terwilliger and Tiedeman, 1978; Tiedeman and Terwilliger, 1978; Hoffmann, 1979; Hoffman
and Alexander, 1980; Hess and Wasser, 1982; Hoffman and Alexander, 1983; Alexander, 1985; Hess and
Alexander, 1986; Alexander, 1987; Banner, 1992; Colorado Natural Areas Program, 1998).
On surface-mined lands in Rocky Mountain shrublands, the establishment of woody plants at densities required
for bond release, and within the liability period, has proven to be very difficult (Mathews and Savage, 1990). This
difficulty arises from numerous factors including grazing and browsing pressures, competition from other plant
species, short growing season, lack of seeds or planting stock, and a lack of technical information regarding the
cost-effective methods for establishing shrubs on disturbed lands.
The purpose of this literature review is to provide a reference to land managers working on woody plant
establishment on disturbed mountain shrublands in the Rocky Mountains. This review will cover the biology,
ecology, and propagation of six species (Table I) that are of primary importance for meeting regulatory compliance
in this region. The information presented here is intended to improve success in establishing these species on
mining and other disturbances in shrublands of the Rocky Mountains in the United States.
BIOLOGY OF IMPORTANT MOUNTAIN SHRUBLAND SPECIES

Following is a brief discussion of the biological characteristics of the six woody species that are the focus of this
review. These species are diverse, but they share many important features (Table II). For instance, all of these
species support important wildlife, and most show large genetic variation, which translates into large geographic
ranges, local ecotypes, varieties, and numerous subspecies. The adaptability of these species to local conditions is
an important part of their biology. The following descriptions generalize many of their qualities, and the reader
should be aware that exceptions to these characteristics will occur. It is also important to note that the adaptations
of these species to specific sites should be considered when planning restoration projects.
Artemisia tridentata Nutt. (Big Sagebrush) vars. wyomingensis (Wyoming Big Sagebrush) and vaseyana
(Mountain Big Sagebrush)
Big sagebrush is the most abundant and widespread shrub in the western United States. It is often thought of as a
symbol of the Old West. It provides important food and cover for domestic animals and wildlife. Once thought of
as an undesirable component of grasslands, it is increasingly recognized for its importance as a wildlife habitat,
especially in light of recent declines in sage grouse populations. Extensive loss of big sagebrush communities in
the Great Basin (Anderson and Inouye, 2001), which is often associated with cheatgrass (Bromus tectorum)
invasion and dominance (Pyke, 1999; Young and Evans, 1978), have prompted much recent interest in restoration
of big sagebrush communities.
Big sagebrush is a long-lived woody shrub that reproduces from seed. Pollination and seed dispersal of big
sagebrush are primarily facilitated by wind, but seeds may also be transported by animals (Bay, 1975; Pendleton
et al., 1989). Like many woody plants of mountain shrublands, germination of big sagebrush seed is poor and is
usually 35 per cent or less (Vines, 1960). Big sagebrush seeds readily establish on bare ground in early spring in the
Copyright # 2003 John Wiley & Sons, Ltd. LAND DEGRADATION & DEVELOPMENT, 14: 459–480 (2003)
Table I. Woody plant species covered by this review. Nomenclature follows that of USDA (USDA, NRCS, 2001)
Scientific name Synonyms Family Common name
Artemisia tridentata Nutt. ssp. Artemisia tridentata Nutt. var. wyomingensis (Beetle & Young) Welsh Asteraceae Wyoming big sagebrush
wyomingensis Beetle & Young Seriphidium tridentatum (Nutt.) W.A. Weber ssp. wyomingense

and (Beetle & Young) W.A. Weber and
Artemisia tridentata Nutt. ssp. Artemisia tridentata Nutt. var. pauciflora Winward & Goodrich
vaseyana (Rydb.) Beetle Artemisia tridentata Nutt. var. vaseyana (Rydb.) Boivin Mountain big sagebrush
Artemisia vaseyana Rydb.
Seriphidium tridentatum (Nutt.) W.A. Weber ssp. vaseyanum (Rydb.) W.A. Weber
Seriphidium vaseyanum (Rydb.) W.A. Weber
Purshia tridentata (Pursh) DC. Rosaceae Antelope bitterbrush
Amelanchier alnifolia (Nutt.) Nutt. Rosaceae Saskatoon serviceberry
ex M. Roemer.
Prunus virginiana L. var. Padus melanocarpa (A. Nels.) Shafer
melanocarpa (A. Nels.) Sarg. Padus virginiana (L.) M. Roemer ssp. melanocarpa (A. Nels.) W.A. Weber Rosaceae Black chokecherry
Prunus melanocarpa (A. Nels.) Rydb.
Prunus virginiana L. ssp. melanocarpa (A. Nels.) Taylor & MacBryde
Cercocarpus montanus Raf. Rosaceae Mountain mahogany
Symphoricarpos spp. Caprifoliaceae Snowberry
S. occidentalis Hook. Western snowberry
S. oreophilus Gray Symphoricarpos rotundifolius Gray var. oreophilus (Gray) M.E. Jones Mountain snowberry
Symphoricarpos tetonensis A. Nels.
Symphoricarpos utahensis Rydb.
BIOLOGY AND ESTABLISHMENT OF MOUNTAIN SHRUBS
Symphoricarpos vaccinioides Rydb.

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462
Table II. Select ecological characteristics of the woody plants described in this literature review. Information from Pendleton et al. 1989; USDA and NRCS, 2001
Big Antelope Saskatoon Black Mountain Western Mountain
sagebrush bitterbrush serviceberry chokecherry mahogany snowberry snowberry
C:N ratio Low Medium/low Medium High Medium/low Medium Medium

Dinitrogen fixation None Low None None Low None None
Palatablity to browsers Low High High High High High Low

Palatablity to grazers Low Medium Low Low Low Low Medium
Protein potential Medium Medium High Medium Medium Low Medium
Fire tolerance High Low High High High High High
Precip. range (cm) 15–40 20–50 30–76 33–165 25–64 30–114 30–100
Min. frost free days 90 100 100 120 90 110 100
Min. temp. ( C) 42 38 39 42 39 38 36
Shade tolerance Intolerant Intermediate Intermediate Intolerant Intermediate Intermediate Intermediate
Pollination Wind Animal Animal Animal Wind Animal Animal
Seed dispersal Wind Mammal Bird Bird Wind Bird Bird
Soil texture adaptation:
Fine No No Yes Yes No No Yes
Medium Yes Yes Yes Yes Yes Yes Yes
Coarse No Yes Yes Yes Yes No No
M. W. PASCHKE ET AL.
Fertility requirement Low Medium Medium Medium Low Medium Low

pH range 6–82 56–84 56–84 52–84 6–8 66–8 52–75
Min. root depth (cm) 50 50 60 50 50 46 30
Moisture use Medium Low Medium Medium Low Low Low
Anaerobic tolerance None Low None Medium None None None
CaCO3 tolerance High High High High High High Low
Drought tolerance High High Medium Medium High High High
Salinity tolerance Low None None None None Low Low
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BIOLOGY AND ESTABLISHMENT OF MOUNTAIN SHRUBS 463
absence of cheatgrass competition. Mountain big sagebrush may reach reproductive maturity in three to five years
and can thus establish quickly (Bunting et al., 1987). Perryman et al. (2000) studied 27 stands of big sagebrush in
Wyoming and found that natural recruitment in these stands occurs in pulses with mean recruitment intervals of
23 years for Wyoming big sagebrush and 22 years for mountain big sagebrush.
Root sprouts are occasionally formed by big sagebrush (Vines, 1960), but it is primarily dependent on
reproduction from seed. Big sagebrush can produce numerous seedlings after a fire, but frequent fires, such as
occur with cheatgrass invasion, can eliminate big sagebrush from a site (Blaisdell, 1953; Wright, 1971; Wright et al.,
1979), especially Wyoming big sagebrush (Bunting et al., 1987). In a review of the literature, Bunting et al. (1987)
found that mountain big sagebrush seeds usually establish readily after a fire whereas, Wyoming big sagebrush
reestablishes slowly after fire.
Big sagebrush occurs on moderately shallow to deep, well-drained, sandy to silt loam soils. It is typically found
on deep soil pockets (Vines, 1960). It can tolerate neutral to slightly alkaline soils, but is generally an indicator of
soils largely free of alkalinity (Vines, 1960). The various subspecies of big sagebrush are distributed on different
substrate types and climatic zones. A. tridentata wyomingensis occurs primarily at warmer lower elevations with
less precipitation and poorly developed soils (West et al., 1978 Hironaka et al., 1983). The subspecies A. tridentata
vaseyana is more common at cooler mid- to high elevations with greater precipitation and more developed soils
(Hironaka et al., 1983; West et al., 1978).
Big sagebrush communities provide critical habitats for about 250 species of vertebrate wildlife (Sands et al.,
1999). These wildlife values are described in detail by Wallmo (1973), McArthur et al. (1978), Blaisdell et al.
(1982), McArthur (1983), Nydegger and Smith (1986), Roberson (1986) and Sands et al. (1999). McArthur (1983)
lists 87 wildlife species that use sagebrush for a habitat, as well as their degree of association, in eight
Intermountain states.
Purshia tridentata (Pursh) DC. (Antelope Bitterbrush)

Antelope bitterbrush is a widespread shrub in the western United States, occurring from the east slope of the Rockies
to the Pacific Ocean, and from British Columbia to Arizona. It is actinorhizal, which means that it is capable of
forming a dinitrogen-fixing symbiosis with Frankia bacteria in root nodules. Bitterbrush and mountain mahogany in
the western United States form some of the most extensive stands of actinorhizal plants in the world, with bitterbrush
alone occupying some 136 million ha in 11 states (Hormay, 1943; Paschke, 1997). Antelope bitterbrush is a key
browse species throughout its range. Recent declines in bitterbrush have reduced mule deer populations (Updike
et al., 1990) and generated interest in the restoration of this species. Following World War II, there was much interest
in increasing the availability of browse on depleted winter ranges of the western United States. Much of the ensuing
research in the 1950s and 1960s focused on antelope bitterbrush for this purpose (Ferguson, 1983).
Flowers of antelope bitterbrush are pollinated by a wide assortment of insects, including honeybees (Bay, 1975;
Pendleton et al., 1989). Natural reproduction of antelope bitterbrush is closely tied to the collection and seed
caching activities of rodents (Evans et al., 1983; Scholten, 1983; Vanderwall, 1994), which act as primary dispersal
vectors (Pendleton et al., 1989). Good seed crops occur at widely varying intervals from two to six years (Young
and Young, 1992).
Antelope bitterbrush can be severely damaged by fire (Pechanec et al., 1948; Blaisdell, 1950; Nord, 1965;
Weaver, 1967; Wright, 1971; Murray, 1983). If soils are wet at the time of a burn, or shortly after a burn, then
bitterbrush will usually resprout (Blaisdell, 1953; Wright, 1971; Driver, 1983; Martin and Driver, 1983). Where
fires are not followed by rain, bitterbrush may not resprout and could be eliminated from the site (Weaver, 1967;
Wright, 1971). However, under good conditions, antelope bitterbrush seedlings may rapidly become established
following fires (Martin, 1982; Driver, 1983; Martin and Driver, 1983). In a literature review of bitterbrush–fire
relationships, Rice (1983) cautions that bitterbrush response to fire appears to be site specific, and it is therefore
difficult to predict what the effects of fire will be in a bitterbrush stand.
The highly palatable nature of bitterbrush often detracts from its use on disturbed lands, and unless grazing and
browsing animals are controlled, its establishment can be quite difficult (Wasser, 1982). Small mammals may also
girdle and kill seedlings in plantings, necessitating the use of chemical deterrents (Wasser, 1982).
Antelope bitterbrush occurs in a variety of habitats including plains, foothills, mountain slopes, mesas and
open woodlands (Stubbendieck et al., 1992), and on a variety of soils (Vines, 1960), but is most abundant on
young, well-drained, deep, coarse-textured soils (Nord, 1965; Stubbendieck et al., 1992). In a survey of 29 natural
bitterbrush stands in California, Nord (1965) found antelope bitterbrush rarely occurred on soils of clay or
clay-loam texture, or where there was a claypan or hardpan within 61 cm of the surface. Nord (1965) also reported
that antelope bitterbrush was rarely found on saline soils that were mildly alkaline or calcareous within 09 to
12 m of the surface. Bitterbrush’s ability to form a dinitrogen-fixing symbiosis with Frankia allows it to grow
on nutrient-poor substrates (Paschke, 1997), and it often grows as a pioneer species on volcanic deposits (Nord,
1965).
Antelope bitterbrush is highly palatable to big game and livestock and it is one of the most widely planted
browse shrubs in North America (Bay, 1975). It is typically regarded as an important winter browse species.
However, Austin and Urness (1983) found the summer diet of mule deer in Utah consisted of 8 per cent bitterbrush
in May and increased steadily to 92 per cent by September. Woodis (1989) found that over a two-year period,
Nevada mule deers’ diets consisted of 51 per cent bitterbrush. Seeds are a valuable food source for chipmunks
(Grant, 1997). Bitterbrush has a low foliar C : N ratio, high protein content, and high N content of dormant tissues.
Bitterbrush is also able to withstand heavy browsing and it seems that browsing may actually stimulate growth of
antelope bitterbrush (Jones, 1983; Urness and Jensen, 1983; Bilbrough and Richards, 1993).
Amelanchier alnifolia (Nutt.) Nutt. ex M. Roemer. (Saskatoon Serviceberry)

Saskatoon serviceberry is a common shrub or small tree of mountain shrublands and prairies of the western United
States and Canada. The fruits of Saskatoon serviceberries, ‘Saskatoons’, were an important source of food for
Native Americans and early settlers (Harris, 1966). Saskatoons are also an important source of food for migratory
songbirds and other wildlife.
Saskatoon serviceberry is a clump-forming, long-lived tree or shrub, which reproduces from animal dispersed
fruits, or from suckers and root sprouts (Harris, 1966; Wasser, 1982). Flowers of Saskatoon serviceberry are
pollinated by animals (insects) and seeds are primarily dispersed by birds (Pendleton et al., 1989).
Wright (1971) describes Saskatoon serviceberry as being severely damaged by fire although, Wasser (1982)
describes stands as being fire tolerant and recovering from sprouts in about 3 years. Lyon and Stickney (1976)
found that saskatoon serviceberry regenerated well from root crowns on three catastrophic fire sites. Similarly,
Noste et al. (1989) working in western Montana tested various intensities and timing of fire on Saskatoon
serviceberry and found that shrubs resprouted well under all treatments, although larger individuals seem to
resprout more vigorously than smaller ones.
Saskatoon serviceberry is a variable species, which occurs in a variety of habitats including brushy hillsides,
open woods, canyons, and riparian areas (Vines, 1960; Stubbendieck et al., 1992). It is a common component in
most mountain shrub communities throughout the Rocky Mountains and Intermountain west. It is adapted to a
wide range of soil types, from fine to coarse textures (USDA and NRCS, 2001). It is most abundant on deep, well-
drained, medium-textured soils, but occasionally grows around bogs (Stubbendieck et al., 1992; Wasser, 1982).
Major and Rejmanek (1992) describe soils where Saskatoon serviceberry is a dominant species as typically young
and rocky with andesite, limestone and/or quartzite parent materials. It will not tolerate saline soils and high
watertables (Wasser, 1982).
Saskatoon serviceberry is excellent cover and browse for deer, elk, and moose, and the young growth is fair to
good for livestock (Vines, 1960; Stubbendieck et al., 1992; Grant, 1997). Austin et al. (1984) reported that the June
diet of mule deer in Utah consisted of 65 per cent Saskatoon serviceberry. The fruit of Saskatoon serviceberry is
an important food for chipmunks, squirrels, black bears, blue grouse, Columbian grouse, sooty grouse,
Richardson’s grouse, mountain quail, and numerous songbirds, and the bark is often eaten by beavers and
marmots (Vines, 1960; Stubbendieck et al., 1992; Grant, 1997). On Yellowstone National Park winter ranges, Kay
(1995) found that browsing by big game may drastically reduce berry production by Saskatoon serviceberry,
suggesting that browse and berry production (for birds or other wildlife) may be incompatible management
objectives for Saskatoon serviceberry.
Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg. (Black Chokecherry)

Black chokecherry is a loose thicket-forming low shrub to small tree with a wide geographic range throughout the
temperate portion of western North America. The fruits of black chokecherries are relished by songbirds and other
wildlife. Black chokecherries are often collected by humans to make jelly and Native Americans regarded it as an
important food and medicinal plant.
Black chokecherry reproduces from bird dispersed seeds (Pendleton et al., 1989), rhizomes or basal sprouts.
Flowers are pollinated by insects (Pendleton et al., 1989). Extensive thickets are often formed from root sprouts
(Vines, 1960). It sprouts readily after fire or mechanical damage from stumps and roots (Wasser, 1982). McKell
(1950) found that black chokecherry increased in abundance two years after fire in Utah, and, after 18 years, it had
twice the cover as an adjacent unburned reference area.
Black chokecherry is common on moist sites, such as drainages, ditches, and road shoulders, and in cool and
moist montane and canyon habitats (Wasser, 1982). Black chokecherry is adapted to a wide range of soil textures,
but will not grow on dense clay and is intolerant of poor drainage and high watertables (Wasser, 1982). It is most
common on moderately deep, silty or sandy soils with good fertility (Wasser, 1982). Black chokecherry will
tolerate moderately acidic to moderately basic soil and weak salinity (Wasser, 1982).
Black chokecherry is regarded as among the most important shrubs for wildlife cover and food in Wyoming and
Colorado (Johnson and Anderson, 1980). Twigs of black chokecherry are good winter browse for wildlife. Fruits
and foliage are an important source of summer and fall food for bears, cottontails, mule deer, and over 40 species
of birds (Vines, 1960; Grant, 1997). Kay (1995) found that browsing by big game may drastically reduce berry
production in black chokecherry, suggesting that browse and berry production may be incompatible management
objectives for black chokecherry.
Cercocarpus montanus Raf. (Mountain Mahogany)

Mountain mahogany is a common shrub or small tree of mountain shrublands and chaparral foothills in the Rocky
Mountains. Related species include C. ledifolius (curlleaf mountain mahogany) and C. betuloides (birchleaf
mountain mahogany). Mountain mahogany and its related species are extremely valuable winter browse for elk,
deer, and bighorn sheep throughout the western United States (reviewed by Paschke, 1997). Like bitterbrush,
mountain mahogany is actinorhizal and thus capable of forming a dinitrogen-fixing symbiosis with Frankia
bacteria in perennial root nodules.
Mountain mahogany typically flowers from June through July, but it may flower from May to November
depending on location (Elmore, 1976; Stubbendieck et al., 1992). The flowers are wind-pollinated and seeds are
dispersed by gravity or wind (Pendleton et al., 1989). The typical life span for mountain mahogany is probably
over 150 years (Ferguson, 1983). Mountain mahogany has good tolerance to fires and usually resprouts vigorously
from root crowns with greater herbage production (Wasser, 1982).
Mountain mahogany is abundant throughout the Rocky Mountains where it occurs in a variety of habitats such
as rocky bluffs, mountain slopes, canyons, rimrock, breaks, and open woodlands (Vines, 1960; Stubbendieck et al.,
1992). Habitats are often rocky, gravelly, and thin-soiled (Wasser, 1982). Mountain mahogany grows on a variety
of soil types, but is typically found on well-drained moderately coarse to moderately fine-textured substrates
(Wasser, 1982). Mountain mahogany is very good browse for cattle, sheep, and goats, although it may contain a
cyanogenic glycoside, which may cause hydrocyanic acid poisoning (Stubbendieck et al., 1992). It is a critical
winter browse species for mule deer and bighorn sheep throughout the Rocky Mountains. Rominger et al. (1988)
found 74–92 per cent of the summer diet of bighorn sheep in the Front Range of Colorado consisted of mountain
mahogany leaves. Seeds are valuable for small game such as grouse (Grant, 1997).
Symphoricarpos occidentalis Hook. (Western Snowberry) and Symphoricarpos oreophilus Gray

(Mountain Snowberry)
Western snowberry is a common species of the Great Plains and Rocky Mountains. It provides important browse
for deer and livestock, and its seeds are eaten by a large variety of birds and mammals. Mountain snowberry is a
common species of mountain shrub zones in the Rocky Mountains. It is very similar to western snowberry in terms
of its biology and utilization.
Flowers of western and mountain snowberry are insect pollinated, and the seeds are dispersed by birds
(Pendleton et al., 1989). Snowberry may be harmed by burning, but populations often recover within 15 years
(Pechanec et al., 1948; Blaisdell, 1953; Wright, 1971) and may form thicker stands after fires (Wasser, 1982).
Although, Anderson and Bailey (1980) noted that annual repeated fires may dramatically reduce western
snowberry.
Snowberry is found in a wide variety of prairie, forest, and shrubland habitats. Western snowberry is most
common on disturbed or overgrazed sites, and in riparian areas, draws, and depressions (Wasser, 1982). Mountain
snowberry is a very common understory component of aspen-fir and spruce-fir forests of the Intermountain west
(Wasser, 1982).
Western snowberry is adapted to most soil textures but is least common on sandy sites (Wasser, 1982). Tueller
and Eckert (1987) studied 37 stands of mountain snowberry/mountain big sagebrush in Nevada and found that the
association occurred primarily on deep, productive, dark-colored soils. Snowberry is tolerant of poorly drained
soils and short-term flooding, but not high watertables. Western snowberry will tolerate weakly acidic to weakly
basic soils. It grows in areas that receive between 300 and 500 mm of mean annual precipitation (Wasser, 1982).
Mountain snowberry is common on sites where snow accumulates and is late to melt (Tueller and Eckert, 1987).
Snowberry is drought tolerant, but growth and seed production are greatly reduced by moisture stress.
Snowberry species are less preferred by game and livestock in winter than many of the other shrubs discussed in
this review. They are, however, sought in spring because they leaf out very early relative to associated browse
species (Plummer et al., 1968). On summer range, snowberry shrubs can be important browse. For instance, Austin
and Urness (1983) found that mountain snowberry made up 65 per cent of the July diet of Utah mule deer. The
fleshy fruits of snowberries are eaten by a variety of bird species (McAtee, 1936; Van Dersal, 1938).
CONSIDERATIONS IN THE RESTORATION OF THESE SPECIES
Weed Control
The control of herbaceous competition, and in particular annual weedy species, might be necessary during the first
few years of shrub establishment on reclamation sites (Cook, 1974). Competition from weeds can greatly reduce
the success of shrub restoration efforts. Herbaceous cover may also provide habitat for small mammals that girdle
and kill shrubs, and herbaceous growth can carry fires that kill shrubs (Frischknecht, 1977). Grasses in particular
can out-compete young shrubs (Kiger et al., 1987). Antelope bitterbrush has vigorous seedlings that may be able to
establish in the presence of grass competition (Monsen and Christensen, 1975). For the other species discussed in
this review, control of herbaceous competition may be necessary prior to planting shrubs (Giunta et al., 1975), and
for one or two complete seasons after planting (Monsen and Christensen, 1975). The use of herbicides prior to
planting of shrubs has been used with limited success (Ianson, 1996).
Since active weed control measures are often cost-prohibitive in reclamation efforts, strategies for avoiding
herbaceous competitors may be a better alternative. This might be accomplished by segregating shrub plantings in
areas where herbaceous competition is likely to be less, or managing weed populations soon after colonization to
avoid their spread and dominance.
Wildlife
Shrubs in mountain shrubland habitats are important winter browse for big game. Planting young succulent shrubs
on reclamation sites in the mountain shrub zone often turns out to be an expensive way to feed wildlife. Shrub
seedlings often require three or four years to establish before they can sustain browsing pressure (Cook, 1974).
Small plantings should be protected by fencing. For larger areas some measures of wildlife control may be
essential to establish shrubs. Meikle et al. (1996) reported that chemical browsing deterrents were ineffective,
while mechanical deterrents such as Tree Pro11 plastic tubes were successful on a mine site in Colorado. Cook
et al. (1974) recommend that only unpalatable shrub species be used in mountain shrub zones. However, where a
wildlife habitat is the desired post-mining land use, this strategy is not an acceptable alternative.
Shrub-seeding failures can often be attributed to seed consumption by rodents (reviewed by Medin and
Ferguson, 1971). Population sizes of rodents fluctuate widely, but these cycles are often predictable and seeding
programs that consider local rodent population levels before broadcast seeding may have greater success.
Measures to control rodents such as providing perches for raptors might help offset losses with minimal costs.
Fruits of Saskatoon serviceberry, snowberry, and black chokecherry are all dispersed by birds. Thus, the
provision of perches on disturbed sites may facilitate colonization by these and other native shrub species
(McClanahan and Wolfe, 1993). In addition, birds and small mammals have been linked to the dispersal of
symbiotic mycorrhizal fungi (Warner and French, 1970; McIlveen and H. Cole, 1976; Ponder, 1980; Warner et al.,
1987; Maser et al., 1988; Allen et al., 1992) and Frankia (Paschke and Dawson, 1993). The attraction of songbirds
to reclamation sites may thus speed the recovery of ecosystem processes by facilitating the input of important soil
biota via birds.
Inoculating with Symbiotic Organisms

Soil is a habitat for numerous and diverse organisms including bacteria, fungi, algae, lichens, protozoa, nematodes,
microarthropods, macroarthropods, annelids, and molluscs. The soil community is an intricate part of the
functionality of most terrestrial ecosystems. When soil habitats are severely degraded, their physical and
biological attributes are compromised, resulting in far-reaching effects on the soil community.
The reestablishment of complete soil functionality following disturbance is dependent on the entire assemblage
of soil organisms and vegetation with their respective influences on the soil environment (Whitford, 1986). In
some instances the introduction of a single genera of organisms to a site can have profound effects. An example is
the practice of introducing symbiotic organisms such as mycorrhizae, or the diazotrophs Rhizobia and Frankia.
Introduction of these microbes is often used successfully to enhance survival and growth of woody plants used in
reclamation efforts. Unfortunately, commercial sources of these and other beneficial organisms are limited (Torrey,
1992). The use of direct-placed topsoil (as opposed to stockpiled) will often serve as an adequate source of these
and other beneficial soil organisms.
Topsoil
The use of topsoil on mine waste usually results in better establishment of vegetation on reclamation areas.
Generally, deeper topsoil treatments result in greater forage production (Nicholas, 1979; Sydnor and Redente,
2000) and topsoil may facilitate the establishment of some shrub species (Schuman et al., 1998, 2000). Shrubs,
however, may not be good competitors with grasses and forbs on spoils covered with relatively deep topsoil
(Draves and Berg, 1980; Redente et al., 1997; Meikle et al., 2000). Shrubs that are adapted to shallow rocky soils
may do better on spoils that are treated with less (or no) topsoil (Semmer and Mills, 1990; Redente et al., 1997),
whereas shrubs that are adapted to deeper- and finer-textured soils often survive better on topsoiled spoils (Monsen
and Richardson, 1984; Meikle et al., 2000). For example, Meikle et al. (2000), working on coal-mine spoils in
Colorado, found that 21 cm of topsoil significantly reduced survival of two out of three species of containerized
shrubs (47 versus 76 per cent survival). They hypothesized that the lower bulk density of the topsoil caused deer
and elk to pull up entire plants rather than simply nipping the tips of the shoots. The authors observed intact
extracted shrubs on the site. In contrast, the heavier spoil substrate seemed to anchor seedlings more securely and
prevented removal of the roots. Similar results have been noted at another mine in Colorado, where topsoil
application reduced the establishment of serviceberry, chokecherry and Gambel oak shrubs (Bitterroot
Consultants, 1998).
1
Mention of trade names does not constitute endorsement by Colorado State University or Colorado Department of Natural Resources Division
of Minerals and Geology.
Live-handled topsoil has advantages over stockpiled topsoil in that viable rhizomes or seeds of shrubs and other
perennial species are often present in fresh topsoil, but not in stockpiled soil (Howard and Samuel, 1979;
Pfannenstiel and Wendt, 1985; Harding and Agnew, 1990; Semmer and Mills, 1990; Schuman et al., 1998).
However, annual weedy species in fresh topsoil may be the resulting community on the reclamation site because of
an abundance of seed in the soil bank (Beauchamp et al., 1975).
Stockpiled topsoil may also have degraded soil communities relative to fresh-hauled topsoil. Since woody plants
compete better when symbiotic soil organisms are present, the use of biologically active topsoil may be very
important when establishing woody species (Stark and Redente, 1987; Schuman et al., 2000). Many of the reported
benefits to shrubs of topsoiling are primarily related to biological activity of the topsoil. The use of some biologically
active topsoil in locations where shrubs are to be established should therefore be considered in reclamation activities.
When handling biologically active topsoil, care should be taken to maintain the biological integrity of the material.
This can be accomplished by using live-hauled, rather than stockpiled topsoil, whenever possible.
Spatial Patterns
For drastically disturbed lands such as mine spoils, the planting of shrubs in clumps may result in more rapid
overall revegetation (West, 1989). Shrub clusters are more efficient at catching snow and soil particles. However,
Carpenter and West (1989) found that dispersed planting of mountain big sagebrush may have improved growth
relative to clumped plantings due to reduced competition for moisture. Shrub clumps or islands may also provide
perches for birds and cover for animals, which facilitate the introduction of beneficial soil organisms (McClanahan
and Wolfe, 1993; Reay and Norton, 1999), which may facilitate revegetation. The concentration of litter, moisture,
and soil organisms under shrub islands may result in acceleration of soil development (West, 1989). There may
also be economic advantages to planting shrubs in clumps, since more intensive management is possible when
shrub plantings are grouped on a limited area of the reclamation site.
SPECIFIC RESTORATION TECHNIQUES FOR THESE SPECIES
Big Sagebrush
Big sagebrush is a valuable reclamation shrub because of its rapid growth rate and exceptional ability to spread
naturally from seed. Wildings and seedlings are transplanted easily and these individuals can reproduce and spread
readily from seed within three to seven years (Plummer et al., 1968; McArthur et al., 1974). Big sagebrush is
difficult to establish by artificial seeding on reclamation sites (reviewed by Schuman et al., 2000). Success with
seeding of big sagebrush seems to be more common with mountain big sagebrush. Methods for seeding big
sagebrush on reclamation sites are not yet established (Kleinman and Richmond, 2000), as there are many
conflicting results reported in the literature regarding cultural practices. Seeding success seems to be random and/
or site specific to date, as cultural practice effects are often masked by precipitation patterns.
Booth and Bai (2000) stress the importance of quality seed lots in determining revegetation success. These
authors recommend that seed buyers look for seed lots that contain less than 3500 seeds g 1, since larger seeds will
produce better establishment. Meyer and Monsen (1990) have shown that the probability of successful establish-
ment at a particular site was increased by using seedlots collected at similar sites.
In a series of studies, McDonough and Harniss found that mountain big sagebrush seeds from Idaho had after-
ripening requirements, which necessitated prechilling (McDonough and Harniss, 1974a,b, 1975). Young and
Young (1992) reported that mountain big sagebrush seeds collected from the western Great Basin did not have any
after-ripening or other forms of dormancy and seeds germinated at 80 per cent when incubated at alternating
temperatures of 15/20 C or 15/25 C.
Artificial seeding of big sagebrush is difficult because the small seeds are adapted for germination at or near the
soil surface (Jacobson and Welch, 1987) making proper placement of seed in seedbeds problematic (Young and
Young, 1992). Jacobson and Welch (1987) determined from careful studies that optimum seeding depth for
mountain big sagebrush was 5 mm or less. For drill seeding, Young and Young (1992) recommend that big
sagebrush seed be mixed with an extender such as rice hulls or vermiculite to control seed metering. A common
reason for failures of big sagebrush seedings, is from drilling seed too deep. Broadcasting sagebrush seed into
seedbeds prepared by mechanical roughing can increase success over drilling (McArthur et al., 1995). Meyer and
Monson (1990) reported 10 per cent establishment of broadcast big sagebrush seed on a mechanically prepared
seedbed. These authors attributed this success to the use of a locally adapted seedlot with high viability. The
importance of using locally-adapted seedlots when restoring big sagebrush was also stressed by McArthur et al.
(1995). Large reclamation areas can be broadcast seeded with aircraft in autumn through midwinter (Monsen,
1999). Autumn planted big sagebrush seed normally germinates in late winter or early spring, and often under
snow (Monsen, 1999). However, these seedlings may be eliminated by periods of soil drying and frost. Availability
of moisture during the early germination phase was found to be strongly related to establishment by Boltz (1994) at
two Idaho sites under a wide variety of cultural techniques.
Monsen (1999) points out that, when conditions are favorable, planting in well-prepared seedbeds can result
in good germination of uniform stands of big sagebrush. However, under unfavorable conditions total failure
may result. As a remedy for this, Monsen (1999) suggests that seeding irregular surfaces in the presence of
surface litter will result in less synchronous establishment and less intraplant competition, resulting in enhanced
survival.
The natural regeneration of big sagebrush is episodic (Perryman et al., 2000) and appears to be related to climate
(reviewed by Kleinman and Richmond, 2000). The practice of seeding big sagebrush as a one-time operation is
therefore contradictory to its biology (Meikle, 2000). Young and Evans (1986) reported widely varying success in
mountain big sagebrush seedings among seeding years, seed collection sites, and planting sites.
At a mine site in Idaho, Richardson et al. (1986) found that mountain big sagebrush established well from
seeding, but only when protected from grazing and seeded with low densities of competing grass and forb species.
In a review of sagebrush restoration on degraded wildlands, Meyer (1994) also concluded that it is important to
reduce early competition from both annual grass and seeded species as well as the manipulation of seedbed
conditions to improve favorable microsites. The importance of favorable microsites was demonstrated by Quinney
et al. (1996), who were able to establish Wyoming big sagebrush in Idaho by concentrating hand broadcasting
efforts in small favorable sites.
Wildings and seedlings of big sagebrush are easy to transplant onto reclamation sites (McArthur et al., 1974).
Big sagebrush transplants in spaced plantings grow rapidly and may produce significant amounts of seed in the
second year (Young and Young, 1992). Young and Evans (1986) reported good success in establishing contain-
erized stock of mountain big sagebrush in Nevada. Larger seedlings had higher initial survival (53 per cent)
compared to smaller containerized stock (15 per cent) on a Wyoming coal mine reclamation site (Meikle et al.,
1995).
The use of stem cuttings for propagation of big sagebrush has not been widely reported. Although, some limited
success with softwood cuttings was noted by Alvarez-Cordero and McKell (1979).
Wasser (1982) suggests that topsoil replacement and/or fertilization to correct for tested nutrient deficiencies
may be necessary to establish big sagebrush on disturbed sites. In addition, irrigation may ensure good
establishment. It should also be noted that fertilization with nitrogen may have the undesirable effect of increasing
competition from weedy annual species (McLendon and Redente, 1991; McLendon and Redente, 1992;
McLendon and Redente, 1994). Big sagebrush should be seeded one or two years prior to grass or forb species
to ensure seedling establishment (Kleinman and Richmond, 2000).
Direct-placed topsoil, mulching, and the presence of mycorrhizae have improved the establishment of big
sagebrush seedlings on reclaimed mined lands (Cockrell et al., 1995; Schuman et al., 1998, 2000). On abandoned
roads in Grand Teton National Park, Wyoming, the use of 15 cm of direct-placed topsoil resulted in 61 per cent
cover of mountain big sagebrush on unseeded plots compared to 00 per cent on control plots (Cotts et al., 1991).
On some reclamation sites, mulching can be important for preventing winter desiccation of seedlings (Kleinman
and Richmond, 2000).
Antelope Bitterbrush
Ease of establishment and rapid growth make antelope bitterbrush a valuable shrub for restoring wildlife habitat on
disturbed lands. It shows excellent success in establishing from direct seeding and bare-root transplants; but using
containerized stock can be more difficult (Plummer et al., 1968). Adaptability of site-specific strains of bitterbrush
warrants caution when purchasing seed or stock.
Bitterbrush seeds are usually dormant and require a prechilling at 0–5 C from three weeks to three months. This
prechilling requirement can be replaced by soaking in hydrogen peroxide (Everett and Meeuwig, 1975). The
natural regeneration of bitterbrush is facilitated by the seed caching activities of rodents (Evans et al., 1983;
Scholten, 1983; Vanderwall, 1994, 1995), and artificial seeding of bitterbrush often results in seed caching by
rodents (Young and Young, 1992).
Seeding antelope bitterbrush is often successful. Wasser (1982) states that: ‘few shrubs excel more than
bitterbrush in range and big game range seedings in its zones of adaptation in the Intermountain West’. For drill
seeding, the seedbed should be clean and firm, and the seed should typically be drilled at about 25 cm depth but
adjusted depending on moisture, texture, and seedbed firmness (Wasser, 1982). Broadcast seed should be covered
to a similar depth (Wasser, 1982). Most seedings of antelope bitterbrush are done in mixtures, so optimum rates for
pure seedings are not well-established. Seeding rates of 06–34 kg ha 1 have been recommended (Plummer et al.,
1968). Clements and Young (2002) seeded at a rate of 34 kg ha 1 and reported seedling densities that were too
high. Seeding is most often done in fall or early winter with untreated seed, but pretreated seed can be sown in
spring.
Dietz et al. (1980), working on a burn site in South Dakota, found 24 per cent germination of fall-seeded
antelope bitterbrush after six years, and a 22 per cent survival of these shrubs after 10 years. The seeds in this study
were sown 5 cm apart and 25 cm deep in hand-cultivated rows. Monson and Shaw (1982) reported 99 per cent
survival of drill-seeded bitterbrush after 39 years on a burn site in Idaho that was protected from grazing. An
adjacent moderately grazed area had no surviving bitterbrush. Clements and Young (2002) describe very good
success in seeding antelope bitterbrush with a no-till or rangeland drill. The use of a no-till drill can limit soil
disturbance and subsequent cheatgrass proliferation. On sites that were not accessible by a drill seeder, Clements
and Young (2002) report good success using a cased-hole punch seeder, which places seeds inside a protective
plastic cylinder that deters predation by rodents. At a mine site in Idaho, Richardson et al. (1986) found that
bitterbrush established well from seeding, but only when protected from grazing and seeded without competing
grass and forb species. Medin and Ferguson (1980) obtained 94 per cent survival of bitterbrush seedlings six years
after seeding a local seed source. Redente (1974) and Taylor (1977) both reported excellent to good establishment
from seeded antelope bitterbrush on surface disturbed sites in Colorado. Similarly, Sindelar et al. (1973) had
success with fall broadcast seeding of bitterbrush on untreated coal mine spoils in Montana (144 seedlings m 2
the following July). However, DePuit and Coenenberg (1979) had no success with a fall broadcast seeding of
bitterbrush on topsoiled spoils in Montana using a seeding rate of 11 kg ha 1.
In general, antelope bitterbrush transplants well (Young and Young, 1992). The use of containerized and bare
root stock of antelope bitterbrush on disturbed sites has been partially successful (Dietz et al., 1980; Young and
Young, 1992), but the operation should be conducted in early spring (Bay, 1975). Success of using nursery stock
depends on the use of dormant, fresh transplants (Bay, 1975; USDI and USDA, 1988). Dietz et al. (1980) found 20
per cent survival of spring-planted containerized stock after 10 years in the Black Hills. Carpenter (1983) reported
good success in transplanting 600 000 antelope bitterbrush seedlings (bare root and containerized) in Idaho.
Clements and Young (2000) reported varying (0–27 per cent two-year survival) success with bitterbrush bare root
transplants in California. They noted that seedlings protected from grazing and weeds, and provided with a source
of Frankia, had better survival.
Antelope bitterbrush can be readily propagated from rooted stem cuttings (Bay, 1975; Kituku et al., 1995).
Kituku et al. (1995) working in Wyoming found that propagation from cuttings and planting-out prior to winter
was a feasible method of preparing containerized bitterbrush for planting in the field.
Saskatoon Serviceberry
Saskatoon serviceberry can be established from seed, transplants, root cuttings or softwood cuttings (Alberta
Agriculture, 1980). Seeded stands are often slow to produce an abundance of forage. Whereas, planted stock often
grows well and can quickly become established on a site.
Embryos of Saskatoon serviceberry require prechilling to overcome dormancy (Dirr and Heuser, 1987; Young
and Young, 1992). The seedcoat may also retard germination, necessitating the use of sulfuric acid for scarification
(Young and Young, 1992). Dirr and Heuser (1987) recommend a 15–30 min treatment with concentrated H2SO4
followed by three months cold stratification. In nature, this scarification is provided by the digestive tract of birds
that consume serviceberry fruits and disperse the seeds (Robinson, 1986). These naturally scarified seeds then
overwinter and germinate in the spring (Young and Young, 1992).
Saskatoon serviceberry seeds should be sown in the fall, or prechilled seeds may be sown in the spring (Vines,
1960; Wasser, 1982; Young and Young, 1992). If dried berries are used (the entire fruit), the berries may need up to
one year of warm stratification (Dirr and Heuser, 1987).
Vines (1960) recommends drilling seed of Saskatoon serviceberry at a rate of 82 seeds per linear meter. Young
and Young (1992) recommend that seeds be sown in drill rows at a rate of about 270 seeds per linear meter and
covered with 05 cm of soil. These authors also suggest that Saskatoon serviceberry will benefit from half shade
during the first year of establishment. A sparse nurse crop of an appropriate species for shading the young
serviceberry seedlings might be useful for this purpose.
Kiger et al. (1987) seeded serviceberry in shrub seed mixtures on fresh subsoil material at a mine in Colorado at
a rate of 11 seeds m 2. Test plots were protected from browsing, and after 11 growing seasons there was good
establishment of serviceberry seedlings.
Container stock of Saskatoon serviceberry should be planted as early as possible in the spring (Wasser, 1982).
Mulching should improve establishment of transplants (USDI and USDA, 1988). Wasser (1982) recommends
using older nursery stock on difficult sites. Dietz et al. (1980) working on a burned and an open pine site in the
Black Hills of South Dakota found that spring planted bare root Saskatoon serviceberry survived 0 and 313 per
cent on the burn and open pine sites respectively after 10 years. Much better results were obtained by Fedkenheuer
et al. (1980) in Alberta, where survival of containerized transplants was between 78 and 100 per cent after two
years. Clark (1996) found that tubelings of serviceberry had higher survival when protected from browsing by deer
(40 versus 25 per cent after six years).
Transplanting wildings might also be an option for this species. Harris (1966) suggests that 15–30 cm tall
vigorous seedlings can be transplanted so long as the roots are not allowed to dry out during the process of
transplanting.
Transplanting mature Saskatoon serviceberry shrubs on a mine site in Colorado resulted in 100 per cent survival
after one year (Larson and Knudson, 1978). Long-term survival of serviceberry in these clumps has been good
(Agnew, 1992). Similar results (85 per cent survival after one year) have been reported for another Colorado mine
(Carlson, 1982), and long-term survival (11 years) of mature transplants at a third Colorado mine was reported at
87 per cent by Kiger et al. (1987).
Dirr and Heuser (1987) emphasize that the key to success with cuttings of Saskatoon serviceberry is timing. The
cuttings must be soft and taken well before the end bud has set. They suggest that late May through June is
optimum. Bishop and Nelson (1980) reported that softwood cuttings taken in late June rooted better than cuttings
taken two weeks earlier or semi-hardwood cuttings taken later. The cuttings should be between 75 and 15 cm long
(Harris, 1966). Cuttings dipped in 3000 ppm IBA-talc and kept in a mist chamber with bottom heat produce good
rooting (Dirr and Heuser, 1987).
Planting stock may be produced from root cuttings (Vines, 1960; Harris, 1966; Alberta Agriculture, 1980).
Pieces of root about 05 cm diameter make the best cuttings. These should be taken as early in the spring as
possible. The root cuttings should be planted with the stem end of the root up and only 05 cm below the surface
(Harris, 1966). Root sprouts (or suckers) may also be harvested in spring and transplanted. Harris (1966)
recommends harvesting the suckers in early spring with as many fine roots as possible and pruning the shoots off at
about 5 cm above the root.
Wasser (1982) recommends that Saskatoon serviceberry be planted separate from other more aggressive plant
species. This can be achieved by planting or seeding in alternate blocks, strips or rows. Meikle et al. (2000) found
that containerized transplants of Saskatoon serviceberry had significantly higher survival (62 per cent) on raw spoil
material as compared to areas receiving 21 cm of topsoil, where survival was 38 per cent. The authors
hypothesized that the lower bulk density of the topsoil allowed deer and elk to pull up entire plants rather than
simply nipping the tips of the shoots. In contrast, the heavier spoil substrate appeared to anchor seedlings more
securely and prevent removal of the roots (Meikle et al., 2000).
Black Chokecherry
Black chokecherry is a useful shrub for general revegetation in mountain shrubland habitats. It can be readily
established from direct seedings and from planting of stock. Success of seedings can vary greatly with seed source
(Plummer et al., 1968). Supplementary water may be needed on some sites during the first year of establishment.
Seeds of the cherry (Prunus) genus have embryo dormancy, which must be overcome with extended prechilling
(Young and Young, 1992). Lockley (1980) reported that the temperature at which germination will take place is
strongly related to stratification time. This study found that 16 weeks of chilling resulted in almost complete
germination of seed at 21–27 C, whereas, during a 24-week chilling the majority of the seed will begin radicle
growth at 3 C. Lockley concluded from his studies that fall planting of chokecherry might be best.
Untreated seed may be sown in the fall. However, for spring sowing, prechilled seeds should be used (Dirr and
Heuser, 1987). The pretreated seed should be sown when most seeds have cracked stones, but before radicles have
elongated to avoid damaging them (Wasser, 1982). Seed can be sown in drills spaced 02–03 m apart at a rate of
about 82 seeds per linear meter (Vines, 1960). Wasser (1982) recommends planting seeds 13 cm deep in moist,
firm seedbeds, or up to 25 cm deep in coarse soils or loose seedbeds, or when fall seeding.
Direct planting of black chokecherry seeds usually results in good establishment (Bay, 1975). Germination of
common chokecherry (Prunus virginiana), that was fall planted by Dietz et al. (1980) in South Dakota, was 19 and
28 per cent on two sites after three years. Long-term survival of these shrubs was 32 and 63 per cent after 10
years. Redente (1974) and Taylor (1977) both reported excellent to good establishment from seeded black
chokecherry on surface disturbed sites in Colorado.
The use of one- or two-year-old planting stock can result in good establishment (USDI and USDA, 1988;
Wasser, 1982). Watering may be required on drier sites. Containerized transplants of common chokecherry spring-
planted in the Black Hills by Dietz et al. (1980) did not survive after 10 years. Bare root transplants of chokecherry
in this experiment did much better, with survival rates of 28 and 37 per cent on two sites after 10 years. Kiger et al.
(1987) reported 21 per cent survival of chokecherry transplants protected from browsing after 11 growing seasons
on a Colorado mine. Mature shrub transplants on two additional mine sites in Colorado had 79 and 88 survival
after one year (Carlson, 1982; Larson and Knudson, 1978). Good long-term survival of these same chokecherry
transplants was subsequently reported (Agnew, 1992).
Semihardwood chokecherry cuttings taken in June were reported to root well by Fung (1984), especially when
treated with 08 per cent IBA. Chokecherry has been propagated from root sucker cuttings. Root cuttings taken in
late winter root well without hormone, but cuttings taken in May root better with a basal dip of 08 per cent IBA
(Allen and Farmer, 1976).
In Colorado, Meikle et al. (2000) found that the addition of 21 cm of topsoil significantly reduced the survival of
containerized chokecherry seedlings (31 per cent survival) relative to seedlings planted on unamended spoil
material (82 per cent survival).
Mountain Mahogany
Mountain mahogany is often successfully established by fall seedings on reclamation sites. However, success can
be limited in dry years. Seed dormancy of mountain mahogany is variable among populations, but seeds generally
require prechilling and or soaking in hydrogen peroxide solution (3–30 per cent v/v) (Young and Young, 1992)
prior to spring sowing. Untreated seed can be sown with success in the fall (Wasser, 1982). Piatt (1973) suggests
that larger seeds have better germination. However, considerable variation can occur in germination both within
(Piatt, 1973) and between seedlots (Smith and Bass, 1973). Seed should be planted 06–13 cm deep, or deeper on
dry or coarse textured substrates (Wasser, 1982). The seed of mountain mahogany is typically planted with
mixtures at a rate of 03–1 kg ha 1 drilled or twice this rate for broadcast seedings (Wasser, 1982).
Mountain mahogany seed sown in fall by Dietz et al. (1980) germinated 19–26 per cent in South Dakota. Survival
of this seed averaged between 36 and 42 per cent after 10 years on two sites that were tested. Establishment of
mountain mahogany seedlings in New Mexico was reported to be limited to shaded microsites by Woodmansee
(1969). This limitation was probably due to both reduced moisture and temperature stress in shade (Piatt, 1976).
Planting stock can be set out in spring after hardening off in cold frames or it may be planted in the summer, but
it requires two months of establishment prior to the first autumn frost (Wasser, 1982). Survival of containerized
mountain mahogany transplants at two sites in South Dakota was 67 and 182 per cent (Dietz et al., 1980).
The Plant Materials Handbook of the USDI Office of Surface Mining and Reclamation (USDI and USDA,
1988) suggests that cuttings of mountain mahogany can be used if they are hardened off prior to spring planting.
Rodents and rabbits often kill mountain mahogany seedlings, and deer and antelope can concentrate on, and
decimate small-scale plantings (Wasser, 1982). The control of wildlife populations may therefore be critical to
establishing mountain mahogany on reclamation sites.
Snowberry
Snowberry is an exceptionally useful species for establishing shrub cover on poor sites (Plummer et al., 1968).
Snowberry readily establishes from wildings and nursery stock. Once established, it can spread vegetatively by
layering and by seed. Plants are often established from rodent seed caches.
Nutlets of snowberries are very difficult to germinate and may require from 20 to 75 min of acid scarification
followed by both warm and then cold stratification (Vines, 1960; Young and Young, 1992). Due to the warm
stratification requirement, field sowing of untreated seeds will often not result in significant emergence until the
second year. Warm stratifying seeds for three or four months prior to fall sowing may overcome this problem
(USDI and USDA, 1988; Young and Young, 1992).
Seeds of snowberry should be planted 06–13 cm deep or deeper in drier, coarse-textured soils, loose seedbeds,
or when fall seeding (Wasser, 1982). Optimum rates of seeding for drilling or broadcasting snowberry seed are not
well established. The USDI Office of Surface Mining Reclamation and Enforcement’s Plant Materials Handbook
(USDI and USDA, 1988), recommends that mulch and N applications will enhance establishment. Kiger et al.
(1987) seeded snowberry in shrub seed mixtures on fresh subsoil material at a Colorado mine at a rate of 6 seeds
m 2. Test plots were protected from browsing and after 11 growing seasons there was good establishment of
snowberry seedlings.
Plummer et al. (1968) report that transplanting wildings of mountain snowberry pulled up by hand from thick
stands was very successful. They report that sufficient transplant material is a piece of stem with a short root
obtained early in the spring while the ground is wet, but before the plant has leafed out. The use of wilding material
may be especially useful for mined land reclamation (McArthur et al., 1974). Sindelar et al. (1973) reported
success in using a front-end loader to scoop up root pads of western snowberry in late May in Montana and
transplanting onto mine waste. The shrubs were mowed to 10 cm height prior to transplanting in this study. Similar
results using a front-end loader to transplant mature clumps of western snowberry were reported for surface mined
land in North Dakota (Friedlander, 1995). The author of the North Dakota study attributed high survival to an
unusually wet period. Long-term survival (11 years) of mature transplants at a mine in Colorado was reported as
100 per cent by Kiger et al. (1987).
Hardwood cuttings of all snowberry species are easy to root. Dirr and Heuser (1987) report that dormant
cuttings taken in December–January, dipped in 3000 ppm IBA-talc, and kept on an open bench with bottom heat
and cool tops rooted 90–100 per cent in four to six weeks. Vines (1960) suggests that plants may be produced from
root cuttings. Given the difficulties of establishing snowberry from seed, it may be more practical, as suggested by
Wasser (1982), to establish it from cuttings or wildings on mined lands. The use of irrigation in the early days of
stand establishment from cuttings has been recommended (USDI and USDA, 1988). However, Howard et al.
(1979) tested cuttings of mountain snowberry at six unirrigated surface mine reclamation sites in Wyoming and
Colorado. Survival of snowberry ranged from 25 to 100 per cent, except for a single mine in Wyoming, were
survival of snowberry and most other shrub cuttings was 0 per cent.
Root cuttings of mountain snowberry have been used with considerable success (Plummer et al., 1968). If a
donor thicket can be found, root segments, or root segments with shoots, can be pulled out of wet soil in the spring
before the shrubs leaf out. These root pieces can then be immediately planted on the reclamation site.
CONCLUSIONS
Based upon this survey of the published literature, there are several generalities that can be made regarding the
establishment of key shrub species on disturbances in mountain shrub zones of the Rocky Mountains. The first is
that most of the species described here show adaptations to local conditions. Therefore, it may be important to
utilize local ecotypes, varieties or subspecies in reclamation efforts. Second, these species are key browse species
for wildlife. Planting them on disturbed sites often turns out to be an expensive wildlife feeding operation.
Protection from browsing during the establishment phase is often the key to success with these species. Third,
competition from weeds or other herbaceous species can greatly reduce the success of shrub establishment.
Implementing strategies to minimize herbaceous competitors will improve success. Lastly, all of these shrub
species are dependant upon soil organisms for proper mineral nutrition, especially in poor soils. Key soil
organisms may be present on raw spoil material, but probably only in small numbers. The use of topsoil in great
depths on mining disturbances may result in poor shrub establishment. However, small quantities of live-hauled
topsoil may provide important biological benefits. Specific recommendations regarding each of the species
covered by this review are summarized in Table III.
Table III. Recommended methods for establishing mountain shrub species on mined land in the US Rocky Mountain region
Species Best method Recommendations
Big sagebrush Seeding Use mountain big sagebrush subspecies where possible
Use seedlots with large seed size (less than 3500 seeds g 1 if possible)
Match seed to site characteristics
Broadcast seed in autumn on mechanically prepared seedbed with irregular
surfaces and surface litter
Seed with no, or low levels, of competing grass and forb species
Use shallow layer of direct-placed topsoil for soil biological benefits
Mulch on dry sites to prevent winter desiccation of seedlings
Antelope bitterbrush Seeding Drill seed with no-till drill in autumn at 25 cm depth, or use cased-hole
punch seeder
Use shallow layer of direct placed topsoil for soil biological benefits
Protect from browsing
Saskatoon serviceberry Seeding Drill seed in autumn at 06 cm depth, or broadcast seed and cover to similar depth
Black chokecherry Seeding Drill seed in autumn at 25 cm depth
Mountain mahogany Seeding Drill seed in autumn at 1 cm depth, or broadcast seed and cover to similar depth
Seed with no, or low levels, of competing grass and forb species
Snowberry Transplants Collect wildings in spring after soil thaw and before leaf-out
Prune or mow shoots to reduce shoot: root
Plant immediately on reclamation site
Whole shrub clumps or ‘pads’ can be transplanted mechanically
or Seeding Drill seed in autumn at 1 cm depth, or broadcast seed and cover to similar depth
REFERENCES
Agnew W. 1992. Woody plant establishment on reclaimed lands at Trapper Mine in northwest Colorado. In Reclamation with Trees and Woody
Shrubs. USDI Office of Surface Mining Reclamation and Enforcement: Duluth, MN; 611–615.
Alberta Agriculture. 1980. Propagation of Saskatoons. Alberta Tree Nursery and Horticulture Centre: Edmonton.
Alexander RR. 1985. Major habitat types, community types, and plant communities in the Rocky Mountains. Report No. RM-123. USDA
Forest Service, Rocky Mountain Forest and Range Experiment Station: Fort Collins, CO.
Alexander RR. 1987. Classification of the forest vegetation of Colorado by habitat type and community type. Report No. RM-478. Rocky
Mountain Forest and Range Experiment Station, USDA Forest Service: Fort Collins, CO.
Allen MF, Crisafulli CA, Friese CF, Jeakins SL. 1992. Re-formation of mycorrhizal symbiosis on Mount St. Helens, 1980–1990: interactions of
rodents and mycorrhizal fungi. Mycological Research 96: 447–453.
Allen R, Farmer RE, Jr. 1976. Mist propagation of chokecherry. Plant Propagator 22: 6–7.
Alvarez-Cordero E, McKell CM. 1979. Stem cutting propagation of big sagebrush (Artemisia tridentata Nutt.). Journal of Range Management
32: 141–143.
Anderson HG, Bailey AW. 1980. Effects of annual burning on grassland in the aspen parkland of east-central Alberta. Canadian Journal of
Botany 58: 985–996.
Anderson JE, Inouye RS. 2001. Landscape-scale changes in plant species abundance and biodiversity of a sagebrush steppe over 45 years.
Ecological Monographs 71: 531–556.
Austin DD, Urness PJ. 1983. Summer use of bitterbrush rangelands by mule deer. In Proceedings: Research and Management of Bitterbrush
and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). US Department of Agriculture Forest Service Intermountain
Forest and Range Experiment Station: Salt Lake City, UT; 203–212.
Austin DD, Urness PJ, King J. 1984. Late summer changes in mule deer diets with increasing use on bitterbrush rangeland. Great Basin
Naturalist 44: 572–574.
Banner RE. 1992. Vegetation types of Utah. Rangelands 14: 109–114.
Bay RR. 1975. Some Important Native Shrubs of the West. USDA Forest Service, Intermountain Forest and Range Experiment Station: Ogden,
UT.
Beauchamp H, Lang R, May M. 1975. Topsoil as a seed source for reseeding strip mine spoils. Wyoming Agricultural Experiment Station
Research Journal 90: 8.
Bilbrough CJ, Richards JH. 1993. Growth of sagebrush and bitterbrush following simulated winter browsing: Mechanisms of Tolerance.
Ecology 74: 481–492.
Bishop BH, Nelson SH. 1980. Propagation and transplanting of saskatoon (Amelanchier alnifolia Nutt.) softwood cuttings. Canadian Journal of
Plant Science 60: 883–890.
Bitterroot Consultants. 1998. Report to Seneca Coal Company: 1997 Reclamation Studies and Shrub Establishment Monitoring at Seneca II
and Seneca II-W mines. Bitterroot Consultants: Corvallis, MT.
Blaisdell JP. 1950. Effects of controlled burning of bitterbrush on the upper Snake River Plains. Report RP-20. USDA Forest Service,
Intermountain Forest and Range Experiment Station: Ogden, UT.
Blaisdell JP. 1953. Ecological effects of planned burning of sagebrush-grass range on the upper Snake River Plains. Report RP-1075. USDA:
Washington, DC.
Blaisdell JP, Murray RB, McArthur ED. 1982. Managing intermountain rangelands: sagebrush-grass ranges. Report No. INT-134. USDA Forest
Service Intermountain Forest and Range Experiment Station: Ogden, UT.
Boltz M. 1994. Factors influencing postfire sagebrush regeneration in south-central Idaho. In Proceedings: Ecology and Management of Annual
Rangelands, Monsen SB, Kitchen SG (eds). USDA Forest Service Intermountain Research Station No. INT-GTR-313: Boise, ID; 281–290.
Booth DT, Bai Y. 2000. Seeds and seedling establishment of Wyoming big sagebrush. In Billings Land Reclamation Symposium 2000: Striving
for Restoration, Fostering Technology and Policy for Reestablishing Ecological Function., Wagner L (ed.). Montana State University,
Reclamation Research Unit: Billings, MT; 282–291.
Bunting SC, Kilgore BM, Bushey CL. 1987. Guidelines for prescribed burning sagebrush-grass rangelands in the Northern Great Basin. Report
INT-231. USDA Forest Service Intermountain Research Station: Ogden, UT.
Carlson KE. 1982. Shrub and tree establishment by mature transplanting on strip mine spoils in northwest Colorado. MS thesis. Colorado State
University: Fort Collins, CO.
Carpenter AT, West NE. 1989. Water relations, aboveground growth, and phenology of Artemisia tridentata spp. vaseyana in relation to
dispersion pattern. In Symposium on Shrub Ecophysiology and Biotechnology, Wallace A, McArthur ED, Haferkamp MR (eds). USDA Forest
Service Intermountain Research Station: Logan, UT; 90–97.
Carpenter R. 1983. Artificial revegetation using antelope bitterbrush-a land managers view. In Proceedings Research and Management of
Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). No. GTR INT-152. USDA Forest Service,
Intermountain Forest and Experiment Station: Salt Lake City, UT; 118–125.
Clark WJ. 1996. Enhancemant of shrub growth on an oil shale tract in the Piceance basin of Colorado. In High Altitude Revegetation Workshop
No. 12, Keammerer WR (ed.). Colorado Water Resources Research Institute, Colorado State University: Fort Collins, CO; 15–25.
Clements CD, Young JA. 2000. Antelope bitterbrush seedling transplant survival. Rangelands 22: 15–17.
Clements CD, Young JA. 2002. Restoring antelope bitterbrush. Rangelands 24: 3–6.
Cockrell JR, Schuman GE, Booth DT. 1995. Evaluation of cultural methods for establishing Wyoming big sagebrush on mined lands. In
Proceedings of the 2nd Annual National Meeting of the American Society for Surface Mining and Reclamation, Schuman GE, Vance GF
(eds). American Society for Surface Mining and Reclamation: Gillette, WY; 784–795.
Colorado Natural Areas Program. 1998. Native Plant Revegetation Guide for Colorado. Colorado Department of Natural Resources: Denver,
CO.
Cook CW. 1974. Surface rehabilitation of land disturbances resulting from oil shale development: final report, phase I, March 1, 1974. Report
No. 1. Environmental Resources Center, Colorado State University: Fort Collins, CO.
Cook CW, Hyde RM, Sims PL. 1974. Guidelines for revegetation and stabilization of surface mined areas in the Western States. Report 16.
Colorado State University Range Science Department: Fort Collins, CO.
Cotts NR, Redente EF, Schiller R. 1991. Restoration methods for abandoned roads at lower elevations in Grand Teton National Park, Wyoming.
Arid Soil Research and Rehabilitation 5: 235–249.
DePuit EJ, Coenenberg JG. 1979. Responses of revegetated coal strip mine spoils to variable fertilization rates, longevity of fertilization
program and season of seeding. Report No. 150. Reclamation Research Unit Montana Agricultural Experiment Station Montana State
University: Bozeman, MT.
Dietz DR, Uresk DW, Messner HE, McEwan LC. 1980. Establishment, survival, and growth of selected browse species in a Ponderosa pine
forest. Report RM-219. USDA forest Service, Rocky Mountain Forest and Range Experiment Station: Fort Collins, CO.
Dirr M, Heuser CW. 1987. The Reference Manual of Woody Plant Propagation: From Seed to Tissue Culture: A Practical Working Guide to the
Propagation of Over 1100 Species, Varieties, and Cultivars. Varsity Press: Athens, GA.
Draves RW, Berg WA. 1980. Shrub establishment on drastically disturbed land in the mountain shrub type. In Society for Range Management
Annual Meeting, Abstracts and Position Statements. Society for Range Management: San Diego, CA; 51.
Driver CH. 1983. Potentials for the management of bitterbrush habitats by the use of prescribed fire. Report INT-152. USDA Forest Service
Intermountain Forest and Range Experiment Station: Salt Lake City, UT.
Elmore FH. 1976. Shrubs and trees of the Southwest Uplands. Report No. 19. Southwest Parks and Monuments Association: Globe, AZ.
Evans RA, Young JA, Cluff GJ, McAdoo JK. 1983. Dynamics of antelope bitterbrush seed caches. In Proceedings Research and Management of
Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). GTR-INT-152. USDA Forest Service Intermountain
Forest and Range Experiment Station: Salt Lake City, UT; 195–202.
Everett RL, Meeuwig RO. 1975. Hydrogen peroxide and thiourea treatment of bitterbrush seed. Report No. INT-196. USDA Forest Service,
Intermountain Forest and Range Experiment Station: Salt Lake City, UT.
Fedkenheuer AW, Heacock HM, Lewis DL. 1980. Early performance of native shrubs and trees planted on amended Athabasca oil sand tailings.
Reclamation Review 3: 47–55.
Ferguson RB. 1983. Use of rosaceous shrubs for wildland plantings in the intermountain west. Report GTR-INT-157. USDA Forest Service
Friedlander JD. 1995. Transplanting and establishment of western snowberry (Symphoricarpos occidentalis Hook.) on reclaimed land. In
Proceedings of the 2nd Annual National Meeting of the American Society for Surface Mining and Reclamation, Schuman GE, Vance GF
(eds). American Society for Surface Mining and Reclamation: Gillette, WY; 779–783.
Frischknecht NC. 1977. Use of shrubs for mined land reclamation and wildlife habitat. In Reclamation for Wildlife Habitat: Proceedings
Reclamation Workshop II. Ecology Consultants: Fort Collins, CO; 113–126.
Fung MYP. 1984. Vegetative propagation of native shrubs in the Fort McMurray area, Alberta, Canada. Plant Propagator 30: 7–9.
Giunta BC, Christensen DR, Monsen SB. 1975. Interseeding shrubs in cheatgrass with a browse seeder-scalper. Journal of Range Management
28: 398–402.
Grant K. 1997. Colorado State Forest Service Nursery Conservation Seedling Tree Program: Species’ Wildlife Values. Colorado State Forest
Service: Fort Collins, CO.
Harding WSJ, Agnew WS. 1990. Shrub establishment and wildlife habitat improvement at Trapper Mine in northwest Colorado. In Fifth
Billings Symposium on Disturbed Land Rehabilitation, Munsshower FF, Fisher SE (eds). USDI Office of Surface Mining Reclamation and
Enforcement: Billings, MT.
Harris RE. 1966. The saskatoon. Report No. 1246. Canada Department of Agriculture: Ottawa.
Hess K, Alexander RR. 1986. Forest vegetation of the Arapaho and Roosevelt National Forests in central Colorado: a habitat type classification.
Report No. RM-266. USDA Forest Service Rocky Mountain Forest and Range Experiment Station: Fort Collins, CO.
Hess K, Wasser CH. 1982. Grassland, Shrubland, and Forested Habitat types of the White River-Arapaho National Forest. Report No. 53-82FT-
1-19. USDA Forest Service: Lakewood, CO.
Hironaka M, Fosberg MA, Winward AH. 1983. Sagebrush-grass habitat types of southern Idaho. Report 35. Forest Wildlife and Range
Experiment Station University of Idaho: Moscow, ID.
Hoffmann GR. 1979. Forest Vegetation of the Rout National Forest, Colorado: A Habitat Type Classification. University of South Dakota,
Department of Biology: Vermillion, SD.
Hoffman GR, Alexander RR. 1980. Forest vegetation of the Routt National Forest in northwestern Colorado: a habitat type classification.
Report No. RM-221. USDA Forest Service, Rocky Mountain Forest and Range Experiment Station: Fort Collins, CO.
Hoffman GR, Alexander RR. 1983. Forest vegetation of the White River National Forest in western Colorado: a habitat type classification.
Report No. RM-249. USDA Forest Service, Rocky Mountain Forest and Range Experiment Station.
Hormay AL. 1943. Bitterbrush in California. Report Research Note 34. USDA Forest Service, California Forest and Range Experiment Station.
Howard GS, Samuel MJ. 1979. The value of fresh-stripped topsoil as a source of useful plants for surface mine revegetation. Journal of Range
Management 32: 76–77.
Howard GS, Rauzi F, Schuman GE. 1979. Woody plant trials at six mine reclamation sites in Wyoming and Colorado. Report No. 177. USDA:
Washington, DC.
Ianson DC. 1996. Evaluation of Oust1 application rates & timing. Bitterroot Restoration: Corvalis, MT.
Jacobson TLC, Welch BL. 1987. Planting depth of ’Hobble Creek’ mountain big sagebrush seed. Great Basin Naturalist 47: 497–499.
Johnson KL, Anderson ES. 1980. Conservation Planting Guide for Wyoming and Colorado. University of Wyoming, Agricultural Extension
Service: Laramie, WY.
Jones RD. 1983. Rotomowing antelope bitterbrush: Preliminary report. In Proceedings: Research and Management of Bitterbrush and Cliffrose
in Western North America, Tiedemann AR, Johnson KL (eds). US Department of Agriculture Forest Service Intermountain Forest and Range
Experiment Station: Salt Lake City, UT 158–162.
Kay CE. 1995. Browsing by native ungulates: effects on shrub and seed production in the Greater Yellowstone ecosystem. In Wildland Shrub
and Arid Land Restoration Symposium, Roundy BA, McArthur ED, Haley JS, Mann DK (eds). INT-GTR-315. USDA Forest Service
Intermountain Research Station: Las Vegas, NV; 310–320.
Kiger RO, Berg WA, Herron JT, Phillips CM, Atkinson RG. 1987. Shrub establishment in the mountain shrub zone. In 4th Biennial Symposium
on Surface Mining and Reclamation of the Great Plains. Montana State University, Reclamation Research Unit 8704: Billings, MT.
Kituku VM, Laycock WA, Powell J, Beetle AA. 1995. Propagating bitterbrush twigs for restoring shrublands. Report No. INT-GTR-315. USDA
Forest Service, Intermountain Research Station: Salt Lake City, UT.
Kleinman LH, Richmond TC. 2000. Sagebrush and mine reclamation: what’s needed from here? In Billings Land Reclamation Symposium
2000. Striving for Restoration, Fostering Technology and Policy for Reestablishing Ecological Function, Wagner L (ed.). Montana State
University, Reclamation Research Unit: Billings, MT; 338–345.
Larson JE, Knudson RJ. 1978. A transplant system for revegetating surface mined land. Report 2630. USDA Forest Service, Equipment
Development Center: Missoula, MT.
Lockley GC. 1980. Germination of chokecherry (Prunus virginiana) seeds. Seed Science and Technology 8: 237–244.
Lyon LJ, Stickney PF. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In Proceedings of the Annual Tall
Timbers Fire Ecology Conference, Komarek E (ed.). University of Montana: Missoula, MT; 355–373.
Major J, Rejmanek M. 1992. Amelanchier-Alnifolia vegetation in Eastern Idaho, USA and its environmental relationships. Vegetatio 98:
141–156.
Martin RE. 1982. Antelope bitterbrush seedling establishment following prescribed burning in the pumice zone of the southern Cascade
Mountains. In Proceedings: Research and Management of Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL
(eds). USDA Forest Service Intermountain Forest and Range Experiment Station, GTR-INT-152: Salt Lake City, UT; 82–90.
Martin RE, Driver CH. 1983. Factors affecting antelope bitterbrush reestablishment following fire. In Proceedings: Research and Management
of Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service Intermountain Forest and
Range Experiment Station, GTR-INT-152: Salt Lake City, UT; 266–279.
Maser C, Maser Z, Molina R. 1988. Small-mammal mycophagy in rangelands of central and southeastern Oregon. Journal of Range
Mathews DT, Savage MS. 1990. Revegetation observations at surface coal mines in the Axial Basin, Yampa River Basin, and North Park,
Colorado. In Fifth Billings Symposium on Disturbed Land Rehabilitation, Munsshower FF, Fisher SE (eds). Montana State University:
Billings, MT; 219–234.
McArthur ED. 1983. Important shrubs for wildland plantings, compositae (Asteraceae). Report GTR INT 157. USDA Forest Service
Intermountain Forest and Range Experimental Station: Ogden, UT.
McArthur ED, Giunta BC, Plummer AP. 1974. Shrubs for restoration of depleted ranges and disturbed areas. Utah Science 35: 28–33.
McArthur ED, Pope CL, Davis JN. 1978. Rehabilitation of game range in the salt desert. In Seventh Wyoming Shrub Ecology Workshop,
Johnson KL (ed.). University of Wyoming: Rock Springs, WY; 23–50.
McArthur ED, Stevens R, Monsen SB. 1995. Adaptation and success of big sagebrush and rubber rabbitbrush on disturbed sites. In Decades
Later: A Time for Reassessment. Proceedings of the 12th annual meeting of the American Society for Surface Mining and Reclamation,
Schuman GR, Vance GF (eds). American Society for Surface Mining and Reclamation: Gillette, WY; 811–823.
McAtee WL. 1936. Groups of plants valuable for wildlife utilization and erosion control. Circular No. 412. USDA: Washington, DC.
McClanahan TR, Wolfe RW. 1993. Accelerating forest succession in a fragmented landscape: the role of birds and perches. Conservation
Biology 7: 279–288.
McDonough WT, Harniss RO. 1974a. Effects of temperature on germination in three subspecies of big sagebrush. Journal of Range
McDonough WT, Harniss RO. 1974b. Seed dormancy in Artemisia tridentata Nutt. subspecies vaseyana Rydb. Northwest Science 48: 17–20.
McDonough WT, Harniss RO. 1975. Know your sagebrush–and better your range. Utah Science 36: 99–103.
McIlveen WD, H. Cole J. 1976. Spore dispersal of Endogonaceae by worms, ants, wasps, and birds. Canadian Journal of Botany 54:
1486–1489.
McKell CM. 1950. A study of plant succession in the oak brush (Quercus gambelii) zone after fire. MS thesis. University of Utah: Salt Lake
City, UT.
McLendon T, Redente EF. 1991. Nitrogen and phosphorus effects on secondary succession dynamics on a semi-arid sagebrush site. Ecology 72:
2016–2024.
McLendon T, Redente EF. 1992. Effects of nitrogen limitation on species replacement dynamics during early secondary succession on a
semiarid sagebrush site. Oecologia 91: 312–317.
McLendon T, Redente EF. 1994. Role of nitrogen availability in the transition from annual-dominated to perennial-dominated seral
communities. In Proceedings Ecology and Management of Annual Rangelands, Monsen SB, Kitchen SG (eds). USDA Forest Service,
Intermountain Forest Range Experiment Station, No. INT-313; 352–362.
Medin DE, Ferguson RB. 1971. Shrub establishment on game ranges in the northwestern United States. In Wildland Shrubs, their Biology and
Utilization, McKell CM, Blaisdell JP, Goodin JR (eds). USDA Forest Service, Intermountain Forest and Range Experiment Station, GTR-
INT-1: Logan, UT; 359–368.
Medin DE, Ferguson RB. 1980. A pilot planting trial on a southwestern Idaho deer winter range. Report No. INT-261. USDA Forest Service,
Meikle TW. 2000. A design solution to big sagebrush establishment: seed production plots and facilitation beds. In Billings Land Reclamation
Symposium 2000—Striving for Restoration, Fostering Technology and Policy For Reestablishing Ecological Function, Wagner L (ed.).
Montana State University, Reclamation Research Unit: Billings, MT; 312–319.
Meikle TM, Ballek L, Briggs B, Noble J. 1995. Big sage (Artemisia tridentata var. wyominensis) an initial comparison of seedling survival
between direct seeding and planting contanerized stock. In Proceedings of the 2nd Annual National Meeting of the American Society for
Surface Mining and Reclamation, Schuman GE, Vance GF (eds). American Society for Surface Mining and Reclamation: Gillette, WY;
796–800.
Meikle TM, Lu S, McAdoo D. 1996. Report to Seneca II Mine: Final Results of Herbivory Study. Bitterroot Consultants: Corvallis, MT.
Meikle TM, Pfannenstiel V, Karo R. 2000. Alteration of topsoil depth for shrub establishment. In Billings Land Reclamation Symposium
2000—Striving for Restoration, Fostering Technology and Policy for Reestablishing Ecological Function, Wagner L (ed.). Montana State
University, Reclamation Research Unit: Billings, MT; 120–126.
Merrill LB. 1971. Selectivity of shrubs by various kinds of animals. In Wildland Shrubs, Their Biology and Utilization, McKell CM, Blaisdell
JP, Goodin JR (eds). USDA Forest Service, Intermountain Forest and Range Experiment Station, GTR-INT-1: Logan, UT; 339–342.
Meyer SE. 1994. Germination and establishment ecology of big sagebrush [Artemisia tridentata]: implications for community restoration. In
Proceedings Ecology and Management of Annual Rangelands, Monsen SB, Kitchen SG (eds). USDA Forest Service, Intermountain Research
Station No. INT-313: Ogden, UT; 244–251.
Meyer SE, Monsen SB. 1990. Seed-source differences in initial establishment for big sagebrush and rubber rabbitbrush. In Proceedings:
Symposium on Cheatgrass Invasion, Shrub Die-off, and Other Aspects of Shrub Biology and Management, McArthur ED, Romney EM,
Smith SD, Tueller PT (eds). General Technical Report GTR-INT-276. USDA Forest Service Intermountain Research Station: Las Vegas, NV;
200–208.
Monsen SB. 1999. Establishment of big sagebrush (Artemisia tridentata) in semiarid environments. In Proceedings: Sagebrush
Steppe Ecosystems, Entwistle PG, Debolt AM, Kaltenecker JH, Steenhof K (eds). Bureau of Land Management: Boise State University:
Boise, ID.
Monsen SB, Christensen DR. 1975. Woody plants for rehabilitating rangelands in the intermountain region. In Wildland Shrubs: Proceedings,
Symposium and Workshop at the Dedication of the US Forest Service Shrub Sciences Laboratory, Stutz EC (ed.). USDA Forest Service, Shrub
Sciences Laboratory: Provo, UT; 72–119.
Monsen SB, Richardson BZ. 1984. Seeding shrubs with herbs on a semiarid mine site with and without topsoil. In Proceedings of the
Symposium on the Biology of Atriplex and Related Chenopods, Tiedemann AR, McArthur ED, Stutz HC, Stevens R, Johnson KL (eds).
USDA Forest Service Intermountain Forest and Range Experiment Station, GTR-INT-172: Provo, UT.
Monsen SB, Shaw NL. 1982. Seeding antelope bitterbrush with grasses on south-central Idaho rangelands: a 39-year response. In Proceedings:
Research and Management of Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service,
Intermountain Forest and Experiment Station: Salt Lake City, UT; 126–136.
Murray RB. 1983. Response of antelope bitterbrush to burning and spraying in southeastern Idaho. In Proceedings: Research and Management
of Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service Intermountain Forest and
Nicholas PJ. 1979. Use of topsoil in the revegetation of a coal strip mine in northwest Colorado. MS thesis. Colorado State University, Fort
Collins, CO.
Nord EC. 1965. Autecology of bitterbrush in California. Ecological Monographs 35: 307–334.
Noste NV, Reinhardt ED, Wilson RA, Jr. 1989. Fire effects on Amelanchier alnifolia shrubs during phenological development stages. In
Symposium on Shrub Ecophysiology and Biotechnology, Wallace A, McArthur ED, Haferkamp MR (eds). USDA Forest Service
Intermountain Research Station, No. INT-256: Logan, UT; 158–163.
Nydegger NC, Smith GW. 1986. Prey populations in relation to Artemisia vegetation types in southwestern Idaho. In Symposium on the Biology
of Artemisia and Chrysothamnus, McArthur ED, Welch BL (eds). USDA Forest Service Intermountain Research Station, No INT-200: Provo,
UT; 152–156.
Paschke MW. 1997. Actinorhizal plants in rangelands of the western United States. Journal of Range Management 50: 62–72.
Paschke MW, Dawson JO. 1993. Avian dispersal of Frankia. Canadian Journal of Botany 71: 1128–1131.
Pechanec JF, Blaisdell JP, Stewart G. 1948. Sagebrush burning: good and bad. Report No. 34. USDA: Washington, DC.
Pendleton RL, Pendleton BK, Harper KT. 1989. Breeding systems of woody plant species in Utah. In Symposium on Shrub Ecophysiology and
Biotechnology, Wallace A, McArthur ED, Haferkamp MR (eds). USDA Forest Service Intermountain Research Station GTR-INT-256:
Logan, UT; 5–22.
Perryman BL, Maier AM, Hild AL, Olson RA. 2000. Demographic characteristics of three Artemisia tridentata subspecies. In Billings Land
Reclamation Symposium 2000: Striving for Restoration, Fostering Technology and Policy for Reestablishing Ecological Function, Wagner L
(ed.). Montana State University, Reclamation Research Unit: Billings, MT; 320–329.
Pfannenstiel VR, Wendt GW. 1985. Enhancing shrub establishment by utilizing direct haul topsoil on mine spoils in western Colorado. In
Symposium on the Reclamation of Lands Disturbed by Surface Mining: A Cornerstone for Communication and Understanding. American
Society for Surface Mining and Reclamation: Owensboro, KY; 1–14.
Piatt JR. 1973. Seed size affects germination of true mountain mahogany [Cercocarpus montanus]. Journal of Range Management 26: 231–232.
Piatt JR. 1976. Effects of water stress and temperature on germination of true mountain mahogany [Cercocarpus montanus]. Journal of Range
Plummer AP, Christensen DR, Monsen SB. 1968. Restoring Big Game Range in Utah. Report Publication 68-3. Utah Division of Fish and
Game: Salt Lake City, UT.
Ponder F. 1980. Rabbits and grasshoppers: vectors of endomycorrhizal fungi on new coal mine spoil. Report NC-250. USDA Forest Service
North Central Forest Experiment Station: St. Paul, MN.
Pyke DA. 1999. Invasive exotic plants in sagebrush ecosystems of the intermountain west. In Proceedings: Sagebrush Steppe Ecosystems,
Entwistle PG, Debolt AM, Kaltenecker JH, Steenhof K (eds). Bureau of Land Management: Boise State University, Boise, ID; 43–54.
Quinney DL, McHenry M, Weaver J. 1996. Restoration of native shrubland in a military training area using hand-broadcasting of seed. In
Proceedings: Shrubland Ecosystem Dynamics in a Changing Environment, Barrow JR, et al. (eds). USDA Forest Service Intermountain
Research Station, INT-GTR-338: Las Cruces, NM; 156.
Reay SD, Norton DA. 1999. Assessing the success of restoration plantings in a temperate New Zealand forest. Restoration Ecology 7: 298–308.
Redente EF. 1974. Revegetation of oil shale lands in Colorado. MS thesis. Colorado State University: Fort Collins, CO.
Redente EF, McLendon T, Agnew W. 1997. Influence of topsoil depth on plant community dynamics of a seeded site in northwest Colorado.
Arid Soil Research & Rehabilitation 11: 139–149.
Rice CL. 1983. A literature review of the fire relationships of antelope bitterbrush. In Proceedings: Research and Management of Bitterbrush
and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service Intermountain Forest and Range
Experiment Station, GTR-INT-152: Salt Lake City, UT; 256–265.
Richardson BZ, Monsen SB, Bowers DM. 1986. Interseeding selected shrubs and herbs on mine disturbances in southeastern Idaho. Report
INT-200. USDA Forest Service Intermountain Forest and Range Experiment Station GTR-INT-200: Provo, UT.
Roberson JA. 1986. Sage grouse-sagebrush relationships: a review. Report INT-200. USDA Forest Service Intermountain Forest and Range
Experiment Station: Provo, UT.
Robinson WA. 1986. Effect of fruit ingestion on Amelanchier seed germination. Bulletin of the Torrey Botanical Club 113: 131–134.
Rominger EM, Dale AR, Bailey JA. 1988. Shrubs in the summer diet Rocky Mountain bighorn sheep. Journal of Wildlife Management 52:
47–50.
Sands AR, Signe-Sather B, Saab V. 1999. Sagebrush steppe wildlife: Historical and current perspectives. In Proceedings: Sagebrush Steppe
Ecosystems, Entwistle PG, Debolt AM, Kaltenecker JH, Steenhof K (eds). Bureau of Land Management: Boise State University: Boise, ID;
27–35.
Scholten GC. 1983. Bitterbrush management on the Boise Wildlife Management area. In Proceedings: Research and Management of
Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service Intermountain Forest and
Schuman GE, Booth DT, Cockrell JR. 1998. Cultural methods for establishing Wyoming big sagebrush on mined lands. Journal of Range
Schuman GE, Booth DT, Olson RA. 2000. Enhancing Wyoming big sagebrush establishment with cultural practice. In Billings Land
Reclamation Symposium 2000: Striving for Restoration, Fostering Technology and Policy for Reestablishing Ecological Function., Wagner L
(ed.). Montana State University, Reclamation Research Unit: Billings, MT; 292–303.
Semmer CE, Mills RA. 1990. Woody plant establishment techniques at Colorado Yampa Coal Company. In Fifth Billings Symposium on
Disturbed Land Rehabilitation, Munsshower FF, Fisher SE (eds). USDI Office of Surface Mining Reclamation and Enforcement: Billings,
MT.
Sindelar BW, Hodder RL, Majerus ME. 1973. Surface mined land reclamation research in Montana. Progress Report, 1972–73. Montana
Agricultural Experiment Station: Bozeman, MT.
Smith DR, Bass LN. 1973. Germinability of true mountain mahogany achenes as influenced by soil and other environmental factors.
Proceedings of the Association of Official Seed Analysts 63: 126–134.
Stark JM, Redente EF. 1987. Production potential of stockpiled topsoil. Soil Science 144: 72–76.
Stubbendieck JL, Hatch SL, Butterfield CH, Jansen BP. 1992. North American Range Plants. University of Nebraska Press: Lincoln, NE.
Sydnor RS, Redente EF. 2000. Long-term plant community development on topsoil treatments overlying a phytotoxic growth medium. Journal
of Environmental Quality 29: 1778–1786.
Taylor PH. 1977. Species adaptability for revegetating surface disturbed lands in Colorado. MS thesis. Colorado State University: Fort Collins,
CO.
Terwilliger C, Tiedeman JA. 1978. Habitat types of the mule deer critical winter range and adjacent steppe region of Middle Park, Colorado.
Report No. 16-739-CA. USDA Forest Service Rocky Mountain Forest and Range Experiment Station: Fort Collins, CO.
Tiedeman JA, Terwilliger C. 1978. Phyto-edaphic Classification of the Piceance Basin. Science Series no. 31. Range Science Department
Colorado State University: Fort Collins, CO.
Torrey JG. 1992. Can plant productivity be increased by inoculation of tree roots with soil microorganisms? Canadian Journal of Forest
Research 22: 1815–1823.
Tueller PT, Eckert RE. 1987. Big sagebrush (Artemisia-Tridentata vaseyana) and longleaf snowberry (Symphoricarpos-Oreophilus) plant
associations in northeastern Nevada. Great Basin Naturalist 47: 117–131.
Updike DR, Loft ER, Hall FA. 1990. Wildfires on big sagebrush/antelope bitterbrush range in northeastern California: Implications for deer
populations. In Symposium on Cheatgrass Invasion, Shrub Die-off, and Other Aspects of Shrub Biology and Management, McArthur ED,
Romney EM, Smith SD, Tueller PT (eds). USDA Forest Service Intermountain Research Station, GTR-INT-276: Las Vegas, NV; 41–46.
Urness PJ, Jensen CH. 1983. Goat use in Fall increases bitterbrush browse and reduces sagebrush density. In Proceedings: Research and
Management of Bitterbrush and Cliffrose in Western North America, Tiedemann AR, Johnson KL (eds). USDA Forest Service Intermountain
Forest and Range Experiment Station, GTR-INT-152: Salt Lake City, UT; 186–194.
USDA and NRCS. 2001. The PLANTS Database, Version 31 (http://plants.usda.gov). National Plant Data Center: Baton Rouge, LA.
USDI and USDA. 1988. Plant Materials Handbook. USDI Office of Surface Mining Reclamation and Enforcement: Washington, DC.
Van Dersal WR. 1938. Native woody plants of the United States: their erosion-control and wildlife values. Report Miscellaneous Publication/
Department of Agriculture No. 303. US: Washington, DC.
Vanderwall SB. 1994. Seed fate pathways of antelope bitterbrush—dispersal by seed-caching yellow pine chipmunks. Ecology 75: 1911–1926.
Vanderwall SB. 1995. Dynamics of yellow pine chipmunk (Tamias-Amoenus) seed caches: underground traffic in bitterbrush seeds. Ecoscience
2: 261–266.
Vines RA. 1960. Trees, Shrubs and Woody Vines of the Southwest. University of Texas Press: Austin, TX.
Wallmo OC. 1973. Important game animals and important recreation in the arid shrublands of the United States. In Arid Shrublands, Third
Workshop, United States/Australian Rangelands Panel, Hyder DN (ed.). Society for Range Management: Tucson, AZ; 98–107.
Wallmo OC, Carpenter LH, Regelin WL, Gill RB, Baker DL. 1976. Evaluation of deer habitat on a nutritional basis. Journal of Range
Warner GM, French DW. 1970. Dissemination of fungi by migratory birds: survival and recovery of fungi from birds. Canadian Journal of
Botany 48: 907–910.
Warner NJ, Allen MF, MacMahon JA. 1987. Dispersal agents of vesicular-arbuscular mycorrhizal fungi in a disturbed arid ecosystem.
Mycologia 79: 721–730.
Wasser CH. 1982. Ecology and culture of selected species useful in revegetating disturbed lands in the West. Report FWS/OSB-82/56. US. Fish
and Wildlife Service: Washington, DC.
Weaver H. 1967. Fire and its relationship to ponderosa pine. Tall Timbers Fire Ecology Conference 7: 127–149.
West NE. 1989. Spatial pattern: Functional interactions in shrub-dominated plant communities. In The Biology and Utilization of Shrubs,
McKell CM (ed.). Academic Press: San Diego, CA; 283–305.
West NE, Tausch RJ, Rea KH, Tueller PT. 1978. Taxonomic determination, distribution, and ecological indicator values of sagebrush within the
pinyon-juniper woodlands of the Great Basin. Journal of Range Management 31: 87–92.
Whitford WG. 1986. The importance of soil ecology and the ecosystem perspective in surface-mine reclamation. In Principles and Methods of
Reclamation Science: With Case Studies from the Arid Southwest, Reith CC (ed.). University of New Mexico Press: Albuquerque, NM;
151–187.
Woodis SG. 1989. Dietary relationships of mule deer and cattle in northwestern Nevada. MS thesis. Colorado State University: Ft. Collins, CO.
Woodmansee RG. 1969. Natural reproduction of Eurotia lanata, Atriplex Canescens, Cercocarpus Montanus, and Cowania Mexicana in New
Mexico. MS thesis. University of New Mexico: Albuquerque, NM.
Wright HA. 1971. Shrub response to fire. In Wildland Shrubs, Their Biology and Utilization, McKell CM, Blaisdell JP, Goodin JR (eds). USDA
Forest Service Intermountain Forest and Range Experiment Station, GTR-INT-1: Logan, UT; 204–217.
Wright HA, Neuenschwander LF, Britton CM. 1979. The role and use of fire in sagebrush-grass and pinyon-juniper plant communities. A state-
of-the-art review. Report GTR-INT-58. Intermountain Forest and Range Experiment Station: Salt Lake City, UT.
Young JA, Evans RA. 1978. Population dynamics after wildfires in sagebrush grasslands. Journal of Range Management 31: 283–289.
Young JA, Evans RA. 1986. Seedling establishment of five sources of big sagebrush in reciprocal gardens. In Symposium on the Biology of
Artemisia and Chrysothamnus, McArthur ED (ed.). USDA Forest Service Intermountain Forest and Range Experiment Station: Provo, UT;
370–374.
Young JA, Young CG. 1992. Seeds of Woody Plants in North America. Dioscorides Press: Portland, OR.

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land degradation & development

Land Degrad. Develop. 14: 459–480 (2003)

Published online 9 July 2003 in Wiley InterScience (www.interscience.wiley.com). DOI: 10.1002/ldr.568

BIOLOGY AND ESTABLISHMENT OF MOUNTAIN SHRUBS ON

Received 28 January 2002; Revised 6 September 2002; Accepted 6 December 2002

Copyright # 2003 John Wiley & Sons, Ltd.

BIOLOGY OF IMPORTANT MOUNTAIN SHRUBLAND SPECIES

Copyright # 2003 John Wiley & Sons, Ltd.

Symphoricarpos vaccinioides Rydb.

LAND DEGRADATION & DEVELOPMENT, 14: 459–480 (2003)

C:N ratio Low Medium/low Medium High Medium/low Medium Medium

Copyright # 2003 John Wiley & Sons, Ltd.

Fertility requirement Low Medium Medium Medium Low Medium Low

LAND DEGRADATION & DEVELOPMENT, 14: 459–480 (2003)

Purshia tridentata (Pursh) DC. (Antelope Bitterbrush)

Amelanchier alnifolia (Nutt.) Nutt. ex M. Roemer. (Saskatoon Serviceberry)

Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg. (Black Chokecherry)

Cercocarpus montanus Raf. (Mountain Mahogany)

Symphoricarpos occidentalis Hook. (Western Snowberry) and Symphoricarpos oreophilus Gray

CONSIDERATIONS IN THE RESTORATION OF THESE SPECIES

Inoculating with Symbiotic Organisms

SPECIFIC RESTORATION TECHNIQUES FOR THESE SPECIES

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