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Rangeland Ecol Manage 67:440–454 | September 2014 | DOI: 10.2111/REM-D-13-00074.1
Resilience and Resistance of Sagebrush Ecosystems: Implications for State and
Transition Models and Management Treatments
Jeanne C. Chambers,
1
Richard F. Miller,
2
David I. Board,
3
David A. Pyke,
4
Bruce A. Roundy,
5
James B. Grace,
6
Eugene W. Schupp,
7
and Robin J. Tausch
8
Authors are
1
Research Ecologist,
3
Ecologist/Data Analyst, and
8
Range Scientist, US Forest Service, Rocky Mountain Research Station, Reno, NV 89512,
USA;
2
Professor, Rangeland Ecology & Management, Oregon State University, Corvallis, OR 97331, USA;
4
Research Ecologist, US Geological Survey,
Forest and Rangeland Ecosystem Science Center, Corvallis, OR 97331, USA;
5
Professor, Plant and Wildlife Sciences, Brigham Young University, UT
84602, USA;
6
Research Ecologist, US Geological Survey, National Wetlands Research Center, Lafayette, LA 70506, USA; and
7
Professor, Wildland
Resources, Utah State University, Logan, UT 84322, USA.
Abstract
In sagebrush ecosystems invasion of annual exotics and expansion of pi
˜
non (Pinus monophylla Torr. and Frem.) and juniper
(Juniperus occidentalis Hook., J. osteosperma [Torr.] Little) are altering fire regimes and resulting in large-scale ecosystem
transformations. Management treatments aim to increase resilience to disturbance and enhance resistance to invasive species by
reducing woody fuels and increasing native perennial herbaceous species. We used Sagebrush Steppe Treatment Evaluation
Project data to test predictions on effects of fire vs. mechanical treatments on resilience and resistance for three site types
exhibiting cheatgrass (Bromus tectorum L.) invasion and/or pi
˜
non and juniper expansion: 1) warm and dry Wyoming big
sagebrush (WY shrub); 2) warm and moist Wyoming big sagebrush (WY PJ); and 3) cool and moist mountain big sagebrush
(Mtn PJ). Warm and dry (mesic/aridic) WY shrub sites had lower resilience to fire (less shrub recruitment and native perennial
herbaceous response) than cooler and moister (frigid/xeric) WY PJ and Mtn PJ sites. Warm (mesic) WY Shrub and WY PJ sites
had lower resistance to annual exotics than cool (frigid to cool frigid) Mtn PJ sites. In WY shrub, fire and sagebrush mowing had
similar effects on shrub cover and, thus, on perennial native herbaceous and exotic cover. In WY PJ and Mtn PJ, effects were
greater for fire than cut-and-leave treatments and with high tree cover in general because most woody vegetation was removed
increasing resources for other functional groups. In WY shrub, about 20% pretreatment perennial native herb cover was
necessary to prevent increases in exotics after treatment. Cooler and moister WY PJ and especially Mtn PJ were more resistant to
annual exotics, but perennial native herb cover was still required for site recovery. We use our results to develop state and
transition models that illustrate how resilience and resistance influence vegetation dynamics and management options.
Key Words: Bromus tectorum invasion, ecological sites, environmental gradients, mechanical treatments, pi
˜
non and juniper
expansion, prescribed fire
INTRODUCTION
Sagebrush ecosystems are undergoing large scale transforma-
tions that are affecting a diversity of rangeland resources and
influencing both management plans and actions (Knick and
Connelly 2011; Miller et al. 2011). Invasion and expansion of
annual invaders, especially cheatgrass (Bromus tectorum L.), at
low to mid elevations are resulting in an annual grass fire cycle
and conversion of large areas to annual weed dominance
(D’Antonio and Vitousek 1992; Balch et al. 2013). Expansion
and infilling of pi
˜
non and juniper trees at middle to high
elevations are resulting in depletion of understory species and
increased risk of large and high-severity fires (Miller et al.
2013). Many sagebrush ecosystems are fragmented and
degraded, and multiple species associated with these ecosys-
tems are of conservation concern (Wisdom et al. 2005).
Researchers and managers alike have emphasized the need to
implement management actions that will increase resilience of
native ecosystems to stress and disturbance and/or enhance
resistance to invasion (Holling 1996; Briske et al. 2008; Brooks
and Chambers 2011; Chambers et al. 2014). We define
resilience as the capacity of an ecosystem to regain its
fundamental structure, processes, and functioning when altered
by stressors like drought and disturbances like overgrazing by
livestock and altered fire regimes (Holling 1973; Allen et al.
2005). We define resistance as the capacity of an ecosystem to
retain its fundamental structure, processes, and functioning
despite stresses, disturbances, or invasive species (Folke et al.
2004). Resistance to invasion by nonnative plants is increas-
ingly important in rangeland ecosystems; it is a function of the
abiotic and biotic attributes and ecological processes of an
ecosystem that limit the population growth of an invading
species (D’Antonio and Thomsen 2004).
To increase the resilience and resistance of sagebrush
ecosystems, management actions have focused on decreasing
woody species (shrubs and/or trees) to 1) reduce fuel loads and
continuity in order to decrease fire severity and extent, 2) lower
competitive suppression of perennial herbaceous species, which
This is Contribution Number 83 of the Sagebrush Steppe Treatment Evaluation Project,
funded by the Joint Fire Science Program (05-S-08), the Bureau of Land
Management, t he US Forest Service, the National Interagency Fire Center, and the
Great Northern Land Conservation Cooperative.
Correspondence: Jeanne C. Chambers, US Forest Service, Rocky Mountain Research
Station, Reno, NV 8 9512, USA. Em ail: jchambers@fs.fe d.us
Manuscript received 22 May 2013; manuscript accepted 1 December 2013.
ª 2014 The Society for Range Management
440 RANGELAND ECOLOGY & MANAGEMENT 67(5) September 2014
largely determine resilience to fire and resistance to invasion,
and 3) decrease longer term risk of cheatgrass dominance
(Miller et al. 2013, 2014b). Responses to management
treatments vary because sagebrush ecosystems occur over a
broad range of abiotic and biotic conditions and differ
significantly in resilience to stress and disturbance and
resistance to invasion (Chambers et al. 2014; Miller et al.
2013). A stronger understanding of factors that influence
resilience to management treatments and resistance to invaders
could increase the ability to prioritize treatment areas and to
select the most appropriate treatments for each area.
Recent research shows that resilience to disturbance changes
along environmental gradients in sagebrush ecosystems (Cham-
bers et al. 2014). Higher resilience is associated with greater
resource availability and more favorable environmental condi-
tions for plant growth and reproduction (Fig. 1A; Chambers
2005; Condon et al. 2011; Davies et al. 2012). Cooler and
moister sites at higher elevations characterized by mountain big
sagebrush (Artemisia tridentata Nutt. ssp. vaseyana [Rydb.]
Beetle) and mountain brush types (i.e., mountain big sagebrush,
snowberry [Symphoricarpos spp.], serviceberry [Amelanchier
spp.]) typically have higher soil water and nutrient availability
and, consequently, higher net productivity than warmer and
drier sites at lower elevations characterized by Wyoming big
sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle
& Young) (West 1983a, 1983b; Tausch and Tueller 1990;
Alexander et al. 1993; Dahlgren et al. 1997; Miller et al. 2011).
These relationships are modified by effects of elevation,
landform, slope, aspect, and soil characteristics, and, thus,
vegetation composition and structure (Johnson and Miller
2006; Condon et al. 2011). Disturbance history differs along
environmental gradients, and because cooler and moister sites
with relatively high productivity had high fuel abundance and
continuity, they had more frequent presettlement fires (Miller et
al. 2011) and often have a greater abundance of fire-tolerant
species (Pyke et al. 2010). Higher mean biomass of perennial
grasses and forbs coupled with more rapid shrub recruitment
on productive, higher elevation sites can result in greater
resilience to disturbance through a smaller initial change in
community composition and more rapid recovery following fire
(Chambers 2005; Davies et al. 2012).
Research also indicates that resistance to invasive annual
grasses changes along environmental gradients in sagebrush
ecosystems (Fig. 1B). Resistance to an invasive species is a
function of the environmental conditions that define where a
species can establish and persist (i.e., fundamental niche) and of
the effects of interactions with the native community on its
actual occurrence (i.e., realized niche) (Chambers et al. 2014).
The fundamental niche of cheatgrass is determined by soil
temperature and water availability and, thus, is strongly
influenced by elevation, landform, slope, and aspect (Chambers
et al. 2007; Condon et al. 2011). In sagebrush ecosystems,
cheatgrass germination, growth, and/or reproduction appear to
be optimal in relatively warm and dry Wyoming big sagebrush
communities due to high climate suitability (Chambers et al.
2007; Roundy et al. 2007; Leger et al. 2009). Low and sporadic
precipitation limit cheatgrass establishment in warm and dry
salt desert communities at low elevations (Meyer et al. 2001),
while low soil temperatures constrain cheatgrass growth and
reproduction in mountain big sagebrush and mountain brush
communities at high elevations (Chambers et al. 2007). The
realized niche represents a subset of the fundamental niche and
is strongly mediated by resource availability and interactions
with the native plant community (Chambers et al. 2014).
Disturbances and stresses, including management treatments
that alter vegetation composition and/or structure, can increase
resource availability and affect resilience to disturbance and
resistance to invaders (Leffler and Ryel 2012). Effects of
vegetation management treatments designed to decrease woody
species depend on characteristics of the area to be treated,
treatment severity, and relative abundance of the primary
functional groups prior to treatment. For example, in wooded
shrublands prescribed fire removes fire-intolerant trees and
shrubs and can result in increased soil water and nutrient
availability (Rau et al. 2007; Roundy et al. 2014b), slow shrub
recovery, and lower resistance to cheatgrass (Miller et al.
2014b; Roundy et al. 2014a). In contrast, mechanical reduction
Figure 1. A, Resilience to disturbance and B, resistance to cheatgrass over
a typical temperature/precipitation gradient in the Great Basin. Predominant
ecological sites that occur along this continuum include Wyoming big
sagebrush on mesic/aridic sites to mountain big sagebrush on frigid/xeric
sites to mountain big sagebrush and root-sprouting shrubs on cryic/xeric
sites. Resilience increases along the temperature/precipitation gradient and
is influenced strongly by site characteristics like elevation and aspect.
Resistance also increases along the temperature/precipitation gradient and
is affected by disturbances and management treatments that alter
vegetation structure and composition and increase resource availability.
(adapted from Chambers et al. 2014)
67(5) September 2014 441
of trees has little effect on shrubs and at low to intermediate
tree covers can result in higher resistance to cheatgrass than
prescribed fire (Miller et al. 2014a; Roundy et al. 2014a).
Treatment outcomes are highly dependent on initial cover of
native perennial herbaceous species, but mechanical removal of
trees results in smaller initial change in community composition
and may result in more rapid recovery compared to burns.
State and transition models (STMs) are widely used in
rangeland management to describe changes in plant commu-
nities and associated soil properties, causes of change, and
effects of management interventions (Stringham et al. 2003;
Bestelmeyer et al. 2009). Incorporating information on the
resilience of rangeland ecosystems to disturbance into STMs
can improve our ability to understand and predict the outcomes
of management actions (Briske et al. 2008; Bagchi et al. 2012).
To date, a lack of information on the ecosystem attributes that
determine resilience and the uncertainties surrounding esti-
mates of these attributes has limited the development of STMs
that address ecosystem resilience (Briske et al. 2008; Bestel-
meyer et al. 2009). To our knowledge, there have been no
attempts to include information specifically related to resis-
tance to invasion in STMs. The Sagebrush Steppe Treatment
Evaluation Project (SageSTEP), a collaborative research and
management effort, provides the necessary data for evaluating
key elements of resilience and resistance to invasion in
sagebrush ecosystems (McIver et al. 2010; McIver and Brunson
2014) and incorporating this information into STMs.
Here we use SageSTEP data to examine effects of fire vs.
mechanical treatments on resilience and resistance to invasion
in sagebrush ecosystems. We compare sites that differ in
environmental characteristics and dominant sagebrush species
and that are exhibiting cheatgrass invasion and/or pi
˜
non and
juniper expansion. We predict that strong differences in
resilience and resistance to cheatgrass and other annual exotics
exist among sites that differ in environmental characteristics
and that these differences influence treatment outcomes. We
ask four related questions to address this prediction: 1) How do
environmental characteristics influence resilience to prescribed
fire and mechanical treatments? 2) How do environmental
characteristics influence resistance to cheatgrass and other
annual exotics after prescribed fire and mechanical treatments?
3) How do contingency effects of weather influence perennial
native herbaceous species and invaders without treatment? 4)
How does pretreatment cover of perennial native herbaceous
species influence resistance to invasion after prescribed fire and
mechanical treatments? We use our results to develop STMs for
sagebrush ecosystems that explicitly incorporate resilience and
resistance information.
METHODS
Study Sites
SageSTEP sites are arrayed across a broad geographical area
that encompasses a range of environmental conditions. For a
map of the SageSTEP site locations and description of the
overall study design, see McIver and Brunson (2014). In this
study, we examine the responses of three site types with
different dominant sagebrush species and soil temperature and
moisture regimes to prescribed fire and mechanical treatments.
We selected seven sites to examine effects of fire and
mechanical treatment on a Wyoming big sagebrush type (WY
shrub) exhibiting invasion by cheatgrass (Onaqui, Owyhee,
Roberts, Gray Butte, Moses Coulee, Rock Creek, and Saddle
Mountain) (see Pyke et al. 2014). Sites were located in five
states (Idaho, Nevada, Oregon, Utah, and Washington) and six
Major Land Resource Areas (Columbia Basin, Columbia
Plateau, Malheur High Plateau, Owyhee High Plateau, Snake
River Plains, and Great Salt Lake Area) (USDA-NRCS 2007).
Subdominants on these sites were bluebunch wheatgrass
(Pseudoroegneria spicata [Pursh] A. Love) or Thurber’s needle
grass (Achnatherum thurberianum [Piper] Barkworth), soils
were loams, soil temperature regimes were mesic to frigid, and
soil moisture regimes were aridic to xeric. Criteria used to select
sites were 1) the dominant shrub was Wyoming big sagebrush,
2) soils were loams, and 3) cheatgrass invasion had occurred
but native grasses and forbs were still present in the understory.
We selected nine sites to examine effects of fire and
mechanical treatment on Wyoming big sagebrush (Marking
Corral, South Ruby, Greenville Bench, Onaqui, and Scipio) and
mountain big sagebrush (Blue Mountain, Devine Ridge, Walker
Butte, and Seven Mile) types exhibiting pi
˜
non and juniper
expansion (see Miller et al. 2014b; Roundy et al. 2014a). Sites
were located in four states (Utah, Nevada, California, and
Oregon) and four Major Land Resource Areas (Malheur High
Plateau, Klamath Basin, Central Nevada Basin and Range, and
Salt Lake Area) (USDA-NRCS 2007). The Wyoming big
sagebrush type (WY PJ) had bluebunch wheatgrass and
needlegrasses in the understory, loamy and typically skeletal
soils, and mesic/aridic to xeric soil temperature/moisture
regimes. The mountain big sagebrush type (Mtn PJ) had Idaho
fescue (Festuca idahoensis Elmer) and bluebunch wheatgrass in
the understory, loamy soils or mollic epipedons, frigid/xeric soil
regimes, and little cheatgrass. Devine Ridge (DR) differed as
codominants were Sandberg bluegrass (Poa secunda J. Presl)
and Thurber needlegrass, soils were orthents, and it was on a
warm, west-facing slope. Criteria used to select sites were 1) the
dominant shrub was big sagebrush, 2) pi
˜
non and/or juniper was
currently expanding into the site, 3) there was no evidence that
stands were dominated previously by mature pi
˜
non or juniper,
4) soils were loams, 5) native grasses and forbs were present in
the understory, and 6) introduced species were present but not a
dominant component.
Treatments
Treatment-plot layout was a randomized complete block with
each of the 16 sites representing a block. The seven sagebrush
sites exhibiting cheatgrass invasion ranged in size from roughly
120 to 325 ha depending on the willingness of land managers
to remove sagebrush. Three treatment plots that were 20 to 81
ha in size were established at each sagebrush site. Each
treatment plot contained 6 to 9 measurement subplots that
were 0.1 ha (33330 m) in size and that were positioned to
cover a range of perennial grass cover. A one-time treatment—
control, prescribed fire, or mowing of sagebrush with a rotary
blade to a height of ~35 cm—was randomly assigned to each
treatment plot in the block. Treatments were applied in 2006,
2007, and 2008 using a stagger-start design (Loughin 2006) in
which all treatments at a given site were applied in the same
442 Rangeland Ecology & Management
year to form a statistical block, but start dates varied among
sites. A stagger-start design alleviates effects of starting an
experiment under the same set of climate conditions so that
results can be applied over a broader inference space.
Prescribed fire was applied in October on all sites except
Moses Coulee, which received a May fire, and mowing was
conducted in the same year as fire within a site. Shrub cover
was reduced from 26% to 4% by fire and from 20% to 8% by
mowing (Pyke et al. 2013).
For each of the nine sagebrush sites exhibiting tree
expansion, three 8–20 ha treatment plots were established
depending on site uniformity and topography. Each treatment
plot contained 15 measurement subplots that were 0.1 ha
(33330 m) in size and that were positioned to include the
range of tree covers that occurred on the site. One of three
treatments was randomly assigned to each treatment plot in the
block—control, cut-and-leave, and prescribed fire. In the cut-
and-leave treatment, all trees . 2 m tall were cut and left on the
ground. Treatments were applied in 2006, 2007, and 2009 for
the pi
˜
non and juniper sites in a stagger-start design (Loughin
2006). Burning and cut-and-leave treatments were applied at
each site in the same year to form a statistical block. Prescribed
fire was applied between August and early November, and tree
cutting was implemented between September and November.
Tree canopies were reduced to , 5% in burn plots and , 1% in
cut-and-leave plots (Miller et al. 2014b).
Measurements
Data were collected from each subplot on all 16 sites during the
growing season immediately prior to treatment application
(year 0) and for three growing seasons afterwards (Miller et al.
2014b; Pyke et al. 2014). Due to funding constraints, fewer
sites were sampled in year 4 posttreatment for all site types, and
the number of subplots sampled was decreased on six of the 11
sites that were sampled. Within each subplot, plant and ground
surface cover were sampled using the point-intercept method at
0.5-m intervals along five, 30-m transects (n¼300 points/
subplot) (Herrick et al. 2009). Plants were recorded by life
form. Native life forms included total shrubs, sagebrush
(Artemisia L.), perennial grasses, perennial forbs, annual forbs,
and soil crusts. Introduced plant life forms included exotic
grasses (primarily cheatgrass) and exotic forbs. Ground surface
cover consisted of bare ground and litter. Foliar cover of each
life form except shrubs was recorded as a single hit at each
point if the point came into contact with that life form. Shrub
canopy cover was measured by recording a hit as a direct
contact or the point falling within the live canopy perimeter.
More than one life form or ground cover class could be
recorded at a single point, but each had a maximum of one hit
per point. Bare ground was recorded when it was the only hit.
Analyses
Site and Treatment Effects. To examine resilience to fire and
resistance to invaders across the three site types, we performed
ANOVAs using generalized linear mixed effects models. We
examined fixed effects of site type, years since treatment,
treatment (fire and control), and their interactions for four
variables—shrub, perennial native herbaceous, all annual
exotic species, and only cheatgrass cover. Based on standard
graphical examinations of residuals, a negative binomial
distribution fit the data best when a log link and an offset of
the number of points sampled per plot was used. The overall
model design was based on a BACI (Before/After Control/
Impact) design in which sampling begins prior to treatment
(year 0) and continues posttreatment (years 1, 2, 3, and 4).
Untreated plots (controls) are sampled to evaluate effects of
natural temporal variation. The spatial design of the experi-
ment was a randomized block. Blocks (random variable) were
sites in which plots (random variable represented by the
interaction of site by treatment) were randomly assigned
treatments. Measurements on the subplots were averaged to
the plot level. A staggered start design was used in which not all
sites were treated in the same year. This created the hierarchical
blocking (random) factors of ‘‘year of treatment.’’ To account
for temporal variation, calendar year, and calendar year by site
type were included as random effects in the model.
Because the two mechanical treatments were not equivalent
in the WY shrub and pi
˜
non-juniper sites, we used the same
method as above, but with two different models, to compare
effects of fire and mechanical treatments. For Mtn PJ and WY
PJ site types, we expanded the treatments to include cut-and-
leave, fire, and control. For WY shrub, we excluded site type
and included mowing, fire, and control. These and all other
analyses that included WY shrub sites excluded fire plots for 1)
the Roberts site due to an inadequate burn and 2) the Moses
Coulee site because the fire occurred in May instead of October.
As described above, year 4 posttreatment had fewer sites and
lower subsample numbers.
Annual Variation in Herbaceous Response Among Controls. To
evaluate the interannual variation during the period of the
experiment for each site type, we conducted ANOVAs in which
control plots were modeled as a function of calendar year, site
type, and their interaction. Response variables used were those
expected to exhibit high interannual variability: perennial
native herbaceous, total annual exotic, and cheatgrass cover.
Based on standard graphical examinations of residuals, a
negative binomial distribution fit the data best when a log link
and an offset of the number of points sampled per plot was
used.
Relationship of Perennial Native Herbaceous Species to Annual
Exotics and Cheatgrass. We examined relationships between
perennial native herbaceous cover in the year of treatment to
perennial native herbaceous, annual exotic, and cheatgrass
cover 3 yr after treatment. We conducted generalized linear
mixed effects ANOVAs for effects of treatment, site type,
perennial native herbaceous cover in the year of treatment, and
their interactions on perennial native herbaceous, total annual
exotic, and cheatgrass cover 3 yr after treatment. Based on
standard graphical examinations of residuals, a negative
binomial distribution fit the data best when a log link and an
offset of the number of points sampled per plot was used. We
maintained the randomized block spatial design of the
experiment and the staggered start design by including
treatment year as a random factor in the model.
We used simple linear regressions to help illustrate the nature
of these relationships. To take advantage of the BACI design,
we also conducted ANOVAs and simple linear regressions of
initial cover of perennial native herbaceous species related to
67(5) September 2014 443
total annual exotics and cheatgrass prior to treatment. These
analyses were based on individual subplot data.
RESULTS
Site and Treatment Effects
Shrub cover showed a three-way interaction among site type,
treatment, and year of response in the overall model
(F
8,39
¼ 4.01, P¼ 0.0015). Cover of shrubs was higher pretreat-
ment and in control plots on WY shrub sites than Mtn PJ sites,
but was higher on WY shrub than WY PJ sites only on fire plots
(Fig. 2; P , 0.05). Shrub cover did not differ between WY PJ
and Mtn PJ sites prior to treatment or in control plots. Fire
decreased shrub cover relative to pretreatment and to control
plots on all site types in all posttreatment years (P , 0.05).
Shrub cover increased from year 1 to years 3 and 4 in fire plots
on WY PJ and Mtn PJ sites (P , 0.05), but did not increase
posttreatment on WY shrub sites. In the model comparing fire
and mowing treatments for the WY shrub site, mowing
decreased shrub cover in all posttreatment years (Fig. 2;
P , 0.004), and shrub cover did not increase on mowed plots
over time. Shrub cover was lower on fire plots than mowed
plots (P , 0.004). In the model comparing fire and cut-and-
leave treatments, the WY PJ and Mtn PJ sites showed similar
responses to each other (Fig. 2). Shrub cover was higher on cut-
and-leave plots than pretreatment and on control plots only in
year 4 (P , 0.006). Fire plots had lower shrub cover than cut-
and-leave plots in all posttreatment years (P , 0.001).
Perennial native herbaceous cover showed significant differ-
ences only among treatments and years of response in the
overall model due to similar responses among sites (treat-
ment3year, F
4,39
¼ 18.30, P , 0.0001). Fire decreased perenni-
al native herbaceous cover relative to pretreatment and to
control plots in the first year after treatment on all sites (Fig. 3;
P , 0.002). However, cover on fire plots was higher in years 2,
3, and 4 relative to year 1 (P , 0.0009). In the model
comparing fire and mowing treatments for the WY shrub site,
perennial native herbaceous cover was higher in years 3 and 4
than pretreatment for both fire (Fig. 3; P , 0.03) and mowed
plots (P , 0.06), but did not differ for treatment and control
plots in years 2, 3, and 4. Fire plots differed from mowed plots
only in the first year after treatment when perennial native
herbaceous cover was lower on fire plots (P¼ 0.0001). In the
model comparing fire and cut-and-leave treatments for WY PJ
and Mtn PJ sites, perennial native herbaceous cover was higher
in cut-and-leave plots than pretreatment or in control plots in
years 2, 3, and 4 (Fig. 3; P , 0.05). Fire plots had lower
perennial native herbaceous cover than cut-and-leave plots in
years 1 and 2 (P , 0.006) but not in years 3 and 4.
Annual exotic cover differed among site types in the overall
model (Fig. 4; F
2,11
¼ 4.46, P¼ 0.0381). Cover of annual exotics
in Mtn PJ sites was less than in WY shrub sites (P , 0.01).
Annual exotic cover in WY PJ sites did not differ from that in
WY shrub sites, but was marginally higher than in Mtn PJ sites
(P , 0.10). A treatment by year interaction also occurred in the
overall model (F
4,39
¼ 14.6, P , 0.0001). Fire increased annual
exotic cover relative to pretreatment and to control plots in
years 3 and 4 (P , 0.01). In the model comparing fire and
mowing treatments for the WY shrub site, annual exotic cover
Figure 2. Shrub cover in control, fire, and mechanical plots on Mtn PJ, WY
PJ, and WY shrub sites the year before treatment and the first 4 yr after
treatment. Values are mean 6 SE.
444 Rangeland Ecology & Management
Figure 3. Perennial native herbaceous cover in control, fire, and
mechanical plots on Mtn PJ, WY PJ, and WY shrub sites the year before
treatment and the first 4 yr after treatment. Values are mean 6 SE.
Figure 4. Annual exotic cover in control, fire, and mechanical plots on Mtn
PJ, WY PJ, and WY shrub sites the year before treatment and the first 4 yr
after treatment. Values are mean 6 SE.
67(5) September 2014 445
was higher on mowed plots in years 3 and 4 compared to
pretreatment (Fig. 4; P , 0.02), but did not differ between fire
and mowed plots. In the model comparing fire and cut-and-
leave treatments for WY PJ and Mtn PJ sites, annual exotic
cover in years 3 and 4 was generally higher on cut-and-leave
plots than pretreatment or on control plots (Fig. 4; P , 0.06).
Annual exotic cover was marginally higher on fire plots than on
cut-and-leave plots in years 3 and 4 (P , 0.07).
Cheatgrass cover exhibited a three-way interaction among
site type, treatment, and year in the overall model (Fig. 5;
F
8,39
¼ 2.11, P¼ 0.0501). Cheatgrass cover was higher on fire
plots in years 3 and 4 than pretreatment on WY shrub and WY
PJ sites (P , 0.03), but only in year 3 on Mtn PJ sites (P¼ 0.03).
However, on WY shrub sites, control and fire plots did not
differ in any year. Cheatgrass cover was higher in fire plots on
WY shrub than Mtn PJ sites in all years, but it did not differ for
WY shrub and WY PJ sites, and was marginally higher in WY
PJ than Mtn PJ sites only in year 4 (P , 0.08). In the model
comparing fire and mowing treatments for the WY shrub site,
cheatgrass cover was higher on mowed plots in years 3 and 4
than pretreatment (Fig. 5; P , 0.01). Cheatgrass cover on
mowed plots did not differ from that on fire plots or control
plots in any year. In the model comparing fire and cut-and-leave
treatments for WY PJ and Mtn PJ sites, cheatgrass cover was
higher on cut-and-leave than pretreatment and control plots in
years 3 and 4 (Fig. 5; P , 0.03). Cheatgrass cover did not differ
for fire vs. cut-and-leave plots, despite apparently higher cover
on fire than cut-and-leave plots in Mtn PJ.
Annual Variation in Herbaceous Cover Among Controls
Perennial native herbaceous cover differed only among years
(F
4,52
¼ 3.25, P ¼ 0.0187) and not among site types on control
plots (Fig. 6). Higher cover occurred in 2011 than all other
years (P , 0.05) on all site types, but no other differences
existed among years.
Annual exotic and cheatgrass cover showed strong differences
among sites types (F
2,52
¼ 28.06, P , 0.0001; F
2,52
¼ 27.01,
P , 0.0001, respectively) and weak differences among years
(F
2,52
¼ 2.33, P , 0.0685; F
2,52
¼ 2.265, P , 0.0751, respectively)
on control plots (Fig. 6). Cover of annual exotics and cheatgrass
decreased in the order: WY shrub . WY PJ . Mtn PJ. Higher
cover of annual exotics and cheatgrass occurred in 2011 than
2008 and 2010 (P , 0.02). Averaged over the 5 yr, cheatgrass
cover comprised 74%, 43%, and 75% of annual exotic cover on
WY shrub, WY PJ, and Mtn PJ control plots, respectively.
Relationship of Perennial Native Herbaceous Species t o
Annual Exotics and Cheatgrass
The relationship of initial perennial native herbaceous cover
before treatment (year 0) to perennial native herbaceous,
annual exotic, and cheatgrass cover in year 3 differed among
site types. Interactions among site type and initial perennial
native herbaceous cover occurred for year 3 cover of perennial
native herbaceous species (F
2,379
¼ 6.79, P¼ 0.0013), annual
exotics (F
2,379
¼ 7.28, P¼ 0.0008), and cheatgrass (F
2,379
¼ 3.68,
P , 0.0262). Also, differences among control and fire treat-
ments occurred in year 3 cover of perennial native herbaceous
species (F
1,11
¼ 5.70, P¼ 0.0361), annual exotics (F
1,11
¼ 10.85,
P¼ 0.0071), and cheatgrass (F
1,11
¼ 4.30, P¼ 0.0608).
Figure 5. Cheatgrass cover in control, fire, and mechanical plots on Mtn
PJ, WY PJ, and WY shrub sites the year before treatment and the first 4 yr
after treatment. Values are mean 6 SE.
446 Rangeland Ecology & Management
Linear regressions illustrate the nature of these relationships
for the different site types. In Mtn PJ sites, initial cover of
perennial native herbaceous species was positively related to
year 3 cover of these species in control (R
2
¼ 0.77, P , 0.0001),
fire (R
2
¼ 0.64, P , 0.0001) and mechanical plots (R
2
¼ 0.69,
P , 0.0001). Cheatgrass cover was generally low on Mtn PJ
sites and exhibited no relationship to initial perennial native
herbaceous species in year 3 on control, fire, or mechanical
plots. Annual exotic cover was comprised almost entirely of
cheatgrass cover on Mtn PJ sites and exhibited very similar
relationships to those for cheatgrass.
In WY PJ sites, cover of perennial native herbaceous species
was more variable among sites and the relationship between
initial and year 3 cover was not as strong as in Mtn PJ for
control (R
2
¼ 0.56, P , 0.0001), fire (R
2
¼ 0.32, P , 0.0001), or
mechanical plots (R
2
¼ 0.54, P , 0.0001). Initial perennial
herbaceous cover and cheatgrass cover in year 3 exhibited no
relationship in control plots, a weak negative relationship in
fire plots (R
2
¼ 0.05, P¼ 0.0421), and a moderate negative
relationship in mechanical plots (R
2
¼ 0.17, P , 0.0001).
Annual exotic cover in year 3 exhibited a weak negative
relationship to initial perennial native herbaceous species in
mechanical plots (R
2
¼ 0.0821, P , 0.01).
In WY shrub sites, the relationship between initial and year 3
perennial native herbaceous cover was strong in control
(R
2
¼ 0.57, P , 0.0001), fire (R
2
¼ 0.48, P , 0.0001), and
mechanical plots (R
2
¼ 0.51, P , 0.0001) (Fig. 7). Overall cover
of cheatgrass in WY shrub sites was high, and initial perennial
native herbaceous cover and year 3 cheatgrass cover were
negatively related in control plots (R
2
¼ 0.15, P¼ 0.0004).
Initial perennial native herbaceous cover and year 3 cheatgrass
cover also were negatively related in fire (R
2
¼ 0.11, P¼ 0.0068)
and especially mow plots (R
2
¼ 0.54, P , 0.0001). When the
Owyhee site, which had little to no cheatgrass cover, was
excluded from analysis the relationship was stronger for fire
plots (R
2
¼ 0.49, P , 0.0001) and similar for mow plots
(R
2
¼ 0.39, P , 0.0001). Similar relationships existed for initial
cover of perennial native herbaceous and year 3 cover of annual
exotics in control (R
2
¼ 0.098, P¼ 0.0062) and mow plots
(R
2
¼ 0.390, P , 0.0001). In fire plots initial perennial native
herbaceous cover and initial cover of annual exotics were
positively related (R
2
¼ 0.3604, P , 0.0001). Annual exotics
had generally high cover in fire plots in year 3 and showed no
relationship to initial perennial native herbaceous cover.
DISCUSSION
Resilience to Fire and Mechanical Treatments
This research confirmed our prediction that differences in
resilience to fire and mechanical treatments exist among site
types differing in environmental characteristics and sagebrush
species. We found that resilience was influenced by soil
temperature/moisture regimes and generally increased from
warm/dry (mesic/aridic) WY shrub to cool moist (frigid/xeric)
Mtn PJ as shown previously (Fig. 1A; Chambers 2005; Condon
et al. 2011; Davies et al. 2012; Chambers et al. 2014).
Although WY shrub sites exhibited no increases in shrub cover
after fire or mowing, WY PJ and Mtn PJ had significantly
higher shrub cover 3 to 4 yr after fire and cut-and-leave
Figure 6. Perennial native herbaceous, total annual exotic, and cheatgrass
cover on control plots in Mtn PJ, WY PJ, and WY shrub plots in 2007
through 2011. Values are mean 6 SE.
67(5) September 2014 447
treatments. The increases in shrub cover on WY PJ and Mtn PJ
sites occurred because of higher levels of sagebrush recruitment
(Miller et al. 2014b) and a greater percentage of root-sprouting
shrubs such as yellow rabbitbrush (Chrysothamnus viscidi-
florus [Hook.] Nutt.) and Saskatoon serviceberry (Amelanchier
alnifolia [Nutt.] Nutt ex M. Roem). All site types had more
perennial, native herbaceous species in years 3 and 4 after
treatment than pretreatment, but fire and mow plots did not
differ from control plots on WY shrub sites. The perennial,
native herbaceous species that showed the greatest response
were Sandberg bluegrass and squirreltail (Elymus elymoides
[Raf.] Swezey) in WY shrub sites, bluebunch wheatgrass in WY
PJ, and Idaho fescue in Mtn PJ sites. Our inability to detect
larger differences among the WY PJ and Mtn PJ types was
likely due to variability among individual sites in soil
temperature/moisture regimes and initial levels of cheatgrass
invasion and/or pi
˜
non and juniper expansion.
Resilience was influenced by treatment severity and initial
abundance of plant functional groups. In WY shrub sites, fire
and mowing both reduced shrub cover and had parallel effects
on posttreatment shrub and perennial native herbaceous cover.
An increase in soil water and nutrient availability, limited
native species response, and high propagule pressure from
annual exotics likely resulted in the observed increase in
cheatgrass and overall annual exotic cover on treated plots
(Leffler and Ryel 2012). Contingency effects of growing season
conditions and competition from cheatgrass and annual exotic
forbs likely caused a lack of a consistent positive response for
perennial native herbaceous and shrub cover in WY shrub sites
(Bakker et al. 2003; Hendrickson and Lund 2010). In WY PJ
and Mtn PJ sites, shrub cover was reduced about 85% by fire,
but was not reduced by the cut-and-leave treatment. Increases
in perennial native herbaceous cover on these sites appeared to
be proportional to available resources as greater increases
occurred on high tree cover plots (Roundy et al. 2014b).
However, posttreatment values reflected pretreatment values
and were lower on plots with high (40–75%) than low (0–
20%) initial tree cover (Roundy et al. 2014b). Also, increases in
cheatgrass and other annual exotics were greater on treated
plots with middle (20–40%) to high (40–75%) than low (0–
20%) initial tree cover (Roundy et al. unpublished data). These
results indicate that treatment severity is typically higher for 1)
fire because it removes most woody vegetation and 2) high tree
cover as most woody vegetation is removed regardless of
treatment and little perennial native herbaceous vegetation
remains to facilitate recovery. On relatively warm and dry sites,
cut-and-leave treatments at low to mid tree covers likely will
result in a higher probability of recovery to a desirable state
than prescribed fire or treating at higher tree covers.
Resistance to Cheatgrass and Other Annual Exotics
Resistance to cheatgrass and other annual exotics was strongly
influenced by soil temperature/moisture regimes in both fire
and mechanical treatments and increased from warm/dry
(mesic/aridic) WY shrub to cold moist (frigid to cool frigid/
xeric) Mtn PJ as predicted (Fig. 1B). WY shrub sites were less
resistant to cheatgrass following fire and mechanical treatment
than WY PJ and Mtn PJ sites, and WY shrub sites and WY PJ
sites were less resistant to annual exotic forbs than Mtn PJ sites.
In WY shrub, treatment outcomes were influenced by generally
greater climatic suitability to cheatgrass (Chambers et al. 2007)
and annual exotic forbs and higher initial cover of exotics. In
WY PJ and Mtn PJ, differences in soil temperature/moisture
regimes among individual sites influenced resistance to
cheatgrass and annual exotic forbs. Cheatgrass cover was
generally low (, 5%) on WY PJ sites, except for Scipio which
was a warm mesic/aridic site, and on Mtn PJ sites, except for
Devine Ridge which was characterized as having frigid/xeric
conditions but was dominated by species adapted to warmer
sites like Thurber’s needlegrass (Miller et al. 2014b). Annual
exotic forb cover was high on only two WY PJ sites, Scipio and
Onaquim which are characterized by warm mesic/aridic
conditions (Miller et al. 2014b). Sites with low cheatgrass
and annual exotic forb cover typically had cool mesic to cool
frigid classifications. SageSTEP sites were selected to have
sufficient perennial herb cover to minimize increases in annual
exotics and did not cover the entire range of environmental
gradients in the Great Basin, but these results are consistent
Figure 7. The relationships between perennial native herbaceous cover in
year 0 (PNH
0
) and perennial native herbaceous cover in year 3 (PNH
3
)
indicated by a dashed line, cheatgrass cover in year 0 (BRTE
0
) indicated by
a dashed and dotted line, and cheatgrass cover in year 3 (BRTE
3
) indicated
by a solid line in control, fire, and mechanical plots on WY shrub sites.
PNH
3
is shown as an x (3), BRTE
0
as an open circle (
*
), and BRTE
3
as a
closed circle (
). Significant regression equations are indicated by an
asterisk.
448 Rangeland Ecology & Management
with prior research in the region (Chambers et al. 2007;
Condon et al. 2011; Davies et al. 2012).
Annual exotic forbs had a significant effect on treatment
response in WY Shrub and WY PJ sites where they occurred.
Species like desert madwort (Alyssum desertorum Stapf.),
curveseed butterwort (Ceratocephala testiculata Crantz [Roth]),
herb sophia (Descurainia sophia [L.] Webb exPrantl), and tall
tumblemustard (Sisymbrium altissimum L.) comprised 45% and
24% of the increase in annual exotic cover on fire and mow
plots in WY shrub sites, and 58% and 40% of the increase on
fire and cut-and-leave plots in WY PJ sites. Annual exotic forbs
including spring verba (Draba verna L.), herb sophia, and yellow
salsify (Tragopogon dubius Scop.) were only 1.3% in both fire
and cut-and-leave plots in frigid Mtn PJ sites. Decreased
competition with cheatgrass and increased soil water availability
can cause increased establishment of annual forbs (Ducas et al.
2011), and both of these conditions likely occurred in the initial
years after treatment (Pyke et al. 2014; Roundy et al. 2014b).
Efforts to control exotic plant species often result in secondary
invasion by nontarget exotic plant species (Rinella et al. 2009;
Larson and Larson 2010). On WY shrub and WY PJ sites,
treatments that decrease woody species competition may release
both cheatgrass and annual forbs and increase fine fuels if sites
lack sufficient native, perennial herbaceous species to effectively
compete for increased resources.
Contingency Effects of Weather on Herbaceous Species
Annual variation of herbaceous cover in sagebrush ecosystems
closely tracks annual precipitation (West and Yorks 2002;
Bradley and Mustard 2005). Annual differences in herbaceous
cover in control plots generally were most pronounced on sites
with high climate suitability for cheatgrass and annual exotic
forbs (i.e., mesic/aridic to xeric soil temperature/moisture
regime). In WY shrub sites, which had high initial cover of
annual exotic species, there was a twofold difference in both
annual exotic and perennial native herbaceous cover between
the lowest and highest average cover. In WY PJ sites, there was a
10-fold difference between the lowest and highest average
annual exotic cover and a threefold difference between the
highest and lowest perennial native herbaceous cover. Longer-
term monitoring will be required to determine resistance to
annual exotics on these sites, but precipitation will likely interact
with treatment effects to influence both native perennial and
annual exotic cover over time (West and Yorks 2002).
Pretreatment Cover of Perennial Native Herbaceous Species
and Resistance to Invasion
Perennial native herbaceous species are a primary determinant
of site resilience to disturbance and management treatments
and/or resistance to cheatgrass and annual exotic forbs
(Chambers et al. 2007; Miller et al. 2014a). In WY shrub
sites, perennial native herbaceous cover of about 20%
appeared necessary to maintain relatively low cover of
cheatgrass in control plots and prevent significant increases
after fire and mowing (Fig. 7). Portions of these sites likely will
exhibit increases in cheatgrass after treatment even with an
average herbaceous cover of 20% due to high variation in cover
across sites and the influence of distances between perennial
plants and biological soil crusts on cheatgrass cover (Reisner et
al. 2013). Annual exotic forbs exhibited a general increase on
WY shrub sites after treatment, especially on fire plots,
regardless of perennial native herbaceous cover. Previous
research indicates that abundance of annual exotic forbs tends
to decrease over time (Allen and Knight 1984; McLendon and
Redente 1990), and annual exotic forbs may return to
pretreatment values in the absence of repeated disturbance.
In WY PJ and Mtn PJ sites, perennial native herbaceous
cover of 20% also appeared necessary to prevent a large
increase in cheatgrass and other annual exotics (data not
shown). In cooler and moister WY PJ and especially Mtn PJ
sites with relatively high resistance, perennial native herbaceous
cover may be less important for preventing dominance by
annual invaders due to lower climate suitability (Chambers et
al. 2007). However, adequate cover of perennial native
herbaceous species and root-sprouting shrubs is still necessary
for soil stabilization and overall site recovery (Miller et al.
2014a). Sites differ in topography, soils characteristics, and
productivity as well as resistance to invaders and all of these
factors should be taken into account when determining
indicators of potential recovery (Miller et al. 2013).
State and Transition Models That Incorporate Resilience and
Resistance Inf ormation
We use our results to expand on previously published STMs for
sagebrush (Holmes and Miller 2010) and wooded shrublands
(Briske et al. 2008; Peterson et al. 2009) by explicitly
incorporating information on resilience to disturbance and
management treatments and resistance to invasion. We use the
interagency framework for developing STMs (Caudle et al.
2013). Our data show that Wyoming big sagebrush sites with
warm/dry (mesic/aridic) soil temperature/moisture regimes are
characterized by low to moderate resilience to fire and
management treatments and low resistance to cheatgrass and
other annual exotics (Fig. 8). The sagebrush mowing treatment
was intended to release perennial herbaceous species and
increase resistance to annual exotics (R2 and 3b in Fig. 8). Both
mowing and fire resulted in slight increases in perennial native
herbaceous cover in the invaded state. However, annual exotic
and cheatgrass cover also tended to increase and there was no
recruitment of sagebrush. Post-burn areas with low perennial
native herbaceous cover were converted to an annual state (T5
and T7 in Fig. 8). The only exceptions were sites with relatively
cool soil temperatures (e.g., Owyhee). Thus, management
treatments are unlikely to provide a restoration pathway to the
reference state once these ecosystems are invaded if insufficient
perennial herbaceous species exist prior to treatment, especially
on the warm end of the temperature/precipitation gradient.
Recent research indicates that seeding after fire may increase
perennial native grasses and sagebrush, but cheatgrass and
annual exotics are likely to persist (R6, R8, and R9 in Fig. 8;
Knutson et al. 2014).
We include both Wyoming and mountain big sagebrush in a
big sagebrush STM with potential for pi
˜
non and juniper
expansion based on the responses of our WY PJ sites and a Mtn
PJ site excluded from this analysis. Mountain big sagebrush
sites typically have cooler and wetter temperature/precipitation
regimes than Wyoming big sagebrush sites (Miller et al.
2013b), but both species have broad ecological amplitudes
67(5) September 2014 449
and can overlap and even hybridize in lower elevation
expansion woodland (Garrison et al. 2013) where management
treatments are often conducted. Our data show that big
sagebrush sites that are exhibiting pi
˜
non and juniper expansion
and that have relatively warm (cool mesic to warm frigid) and
moist (xeric) soil temperature/moisture regimes are character-
ized by moderate resilience to fire and management treatments
and moderately low resistance to cheatgrass and other annual
exotics (Fig. 9). Increases in perennial native herbaceous species
and shrub recruitment occurred over a range of tree covers in
the early to mid-phases of tree expansion (Phase I through
Phase II) for both the cut-and-leave (4b in Fig. 9) and fire
treatment (3 and 5 in Fig. 9). This is consistent with the
community phase pathways for the reference state illustrated
elsewhere for wooded shrublands (Briske et al. 2008; Peterson
et al. 2009). Prior research indicates that infilling of Phase II
woodlands can result in a biotic threshold to a wooded state
with increased risk of high severity crown fires (T6 in Fig. 9)
and, depending on soils, slope, and understory species, an
abiotic threshold to an eroded state (T7 in Fig. 9; Miller et al.
2014a). We found that on relatively warm and dry sites,
presence of cheatgrass and other annual exotics coupled with
low cover of perennial native herbaceous species resulted in a
wooded/invaded state following cut-and leave-treatments (T8
in Fig. 9) and an annual state following fire (T10 and T11 in
Fig. 9). The increase in cheatgrass and other annual exotics
cover was positively related to tree cover (Roundy et al.
2014a). Other research indicates that seeding can increase
perennial herbaceous species and sagebrush (R12, R13, R14,
and R15 in Fig. 9). Depending on seeding mix, grazing, and
weather conditions, cooler and wetter sites can return to the
Figure 8. A state and transition model for a Wyoming big sagebrush ecosystem with a mesic/aridic soil temperature/moisture regime that is characterized
by low to moderate resilience to disturbance and management treatments and low resistance to cheatgrass and other annual exotics. Large boxes illust rate
states that are comprised of community phases (smaller boxes) which interact with the environment to produce a characteristic composition of plant
species, functional and structural groups, soil functions, and range of variability. Transitions among states are shown with arrows starting with T;
restoration pathways are shown with arrows starting with R. The ‘‘at risk’’ community phase is most vulnerable to transition to an alternative state.
450 Rangeland Ecology & Management
reference state if an abiotic threshold has not been crossed (R16
in Fig. 9) (Pyke 2011).
Our data show that mountain big sagebrush sites that are
exhibiting pi
˜
non and juniper expansion and that have cool
and moist (cool frigid/xeric) soil temperature/moisture re-
gimes are characterized by moderately high resilience to
disturbance and resistance to invasives (Fig. 10). As for the big
sagebrush type, increases in perennial native herbaceous
species and shrub recruitment occurred across a range of tree
densities (Phase I through Phase II) for cut-and-leave (4b in
Fig. 10) and fire treatments (3 and 5 in Fig. 10). Annual
exotics had generally low abundance on these sites. As for WY
PJ, in the absence of treatment, infilling of phase II woodlands
can result in a biotic threshold to a wooded state and,
depending on soils and topography, an abiotic threshold
crossing to an eroded state (T7 in Fig. 10; Miller et al. 2014a).
Seeding of Phase III woodlands (closed wooded shrublands)
with depleted understories after fire can increase perennial
herbaceousspeciesandsagebrush(R8andR9inFig.10)
(Pyke 2011). Depending on seed mix and grazing, return to
the reference state may be possible if an irreversible threshold
has not been crossed.
Figure 9. A state and transition model for a big sagebrush ecosystem with a cool mesic to warm frigid/xeric soil temperature/moisture regime that is
exhibiting pi
˜
non and juniper expansion. This type is characterized by moderate resilience to disturbance and management treatments and moderately low
resistance to cheatgrass and other annual exotics. Large boxes illustrate states that are comprised of community phases (smaller boxes) which interact
with the environment to produce a characteristic composition of plant species, functional and structural groups, soil functions, and range of variability.
Transitions among states are shown with arrows starting with T; restoration pathways are shown with arrows starting with R. The ‘‘at risk’’ community
phase is most vulnerable to transition to an alternative state.
67(5) September 2014 451
MANAGEMENT IMPLICATIONS
An understanding of the relative differences in resilience to
disturbance and management treatments and resistance to
cheatgrass and other annual exotics can be used to prioritize
areas for treatment and select the most appropriate treatments.
Resilience to management treatments and resistance to annual
exotic species is influenced by soil temperature/moisture
regimes and generally increases from warm/dry (mesic/aridic)
Wyoming big sagebrush to cool/moist (frigid/xeric) mountain
big sagebrush sites. We found that warm/dry Wyoming big
sagebrush sites had low to moderate resilience to fire and
Figure 10. A state and transition model for a mountain big sagebrush ecosystem with a cool frigid/xeric soil temperature/moisture regime that is exhibiting
pi
˜
non and juniper expansion. This type is characterized by moderately high resilience to disturbance and management treatments and resistance to
cheatgrass and other annual exotics. Large boxes illustrate states that are comprised of community phases (smaller boxes) which interact with the
environment to produce a characteristic composition of plant species, functional and structural groups, soil functions, and range of variability. Transitions
among states are shown with arrows starting with T; restoration pathways are shown with arrows starting with R. The ‘‘at risk’’ community phase is most
vulnerable to transition to an alternative state.
452 Rangeland Ecology & Management
mowing treatments and generally low resistance to annual
exotics. In the first 4 yr after treatment, we found that fire and
mowing resulted in only small increases in perennial native
grasses and forbs, little to no shrub recruitment, and significant
increases in annual exotics. Wyoming and mountain big
sagebrush sites exhibiting pi
˜
non and juniper expansion with
intermediate soil temperature regimes (cool mesic to warm
frigid/xeric) had moderate resilience to fire and cut-and-leave
treatments but moderately low resistance to annual exotics.
Fire and cut-and-leave treatments increased both perennial
native herbaceous species and shrub recruitment, but large
increases in annual exotics occurred on some sites. In relatively
warm big sagebrush sites in general (warm mesic to warm
frigid), perennial native grass and forb cover of about 20%
prior to treatments appeared necessary to prevent significant
increases in cheatgrass and other exotic annuals posttreatment.
Mountain big sagebrush with cool and moist (cool frigid/xeric)
soil temperature/moisture regimes had moderately high resil-
ience to fire and cut-and-leave treatments and resistance to
annual exotics. Fire and cut-and-leave treatments increased
both perennial native herbaceous species and shrub recruitment
and annual exotics were a minor component of these sites.
Treatment severity increased in big sagebrush sites exhibiting
tree expansion with 1) fire, because it removed all woody
vegetation, and 2) high tree cover, as most woody vegetation
was removed regardless of treatment and little perennial native
herbaceous vegetation remained to facilitate recovery. On sites
with relatively low resistance to annual exotics, cut-and-leave
treatments at low to middle tree covers result in a higher
likelihood of recovery to a desirable state than prescribed fire.
On sites with relatively high resistance to annual exotics, either
fire or cut-and-leave treatments at low to middle tree covers can
result in a desirable state. STMs that incorporate information
on resilience to management treatments and resistance to
annual exotics can be used as aids in the planning process.
Sagebrush sites occur over continuums of ecological conditions,
such as temperature and precipitation, and careful assessment
of site conditions always will be necessary to determine the
relevance of a particular STM, the suitability of a site for
treatment, and the most appropriate treatment(s) (Pyke 2011;
Chambers et al. 2014; Miller et al. 2013, 2014a).
ACKNOWLEDGMENTS
We thank Dave Turner for statistical advice, and Jim McIver, Peter
Weisberg, Tanya Skurski, and three anonymous reviewers for helpful
comments on the manuscript, and Mike Pellant for assistance with
development of the STMs. We acknowledge and thank our additional
SageSTEP co-PI’s who contributed to the design of this study. This is
Contribution Number 83 of the Sagebrush Steppe Treatment Evaluation
Project, funded by the Joint Fire Science Program (05-S-08), the Bureau of
Land Management, the US Forest Service, the National Interagency Fire
Center, and the Great Northern Land Conservation Cooperative.
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