(PDF) Manipulation of Rangeland Wildlife Habitats

Chapter 5

Manipulation of Rangeland Wildlife

Habitats

David A. Pyke and Chad S. Boyd

Abstract Rangeland manipulations have occurred for centuries. Those manipula-

tions may have positive or negative effects on multiple wildlife species and their

habitats. Some of these manipulations may result in landscape changes that frag-

ment wildlife habitat and isolate populations. Habitat degradation and subsequent

restoration may range from simple problems that are easy to restore to complex

problems that require multiple interventions at multiple scales to solve. In all cases,

knowledge of the wildlife species’ habitat needs throughout their life history, of

their population dynamics and habitat-related sensitivities, and of their temporal

and spatial scale for home ranges and genetic exchange will assist in determining

appropriate restoration options. Habitat restoration will begin with an understanding

of the vegetation’s successional recovery options and their time scales relative to

wildlife population declines. We discuss passive and active manipulations and their

application options. Passive manipulations focus on changes to current management.

Active manipulations may include removal of undesirable vegetation using manual

harvesting, mechanical, chemical, or biological methods while desirable vegetation

is enhanced through the reintroduction of desirable wildlife habitat structure and

function. These techniques will require monitoring of wildlife and their habitat at

both the landscape and site level in an adaptive management framework to learn from

our past and improve our future management.

Keywords Adaptive management ·Climate change ·Landscapes ·Monitoring ·

Passive versus active management ·State and transition models ·Wildlife habitat

management ·Vegetation manipulations

D. A. Pyke (B)

U.S. Geological Survey, Forest and Ecosystem Science Center, Corvallis, OR 97331, USA

e-mail: david_a_pyke@usgs.gov

C. S. Boyd

USDA Agricultural Research Service, Eastern Oregon Agricultural Research Center, Burns,

OR 97720, USA

L. B. McNew et al. (eds.), Rangeland Wildlife Ecology and Conservation,

https://doi.org/10.1007/978-3-031-34037-6_5

107

108 D. A. Pyke and C. S. Boyd

5.1 Introduction

Early hominins likely began manipulating their environment soon after they learned

to control ﬁre between about 1.5 and 0.4 million ybp (Gowlett 2016). They may

have noticed beneﬁts of improved hunting and gathering after wildﬁres thus leading

to intentional ﬁres to gain those beneﬁts. One of the earliest documented cases of

manipulating habitats for the beneﬁt of wildlife was during the thirteenth century

reign of Kublai Khan (Valdez 2013). Native Americans commonly used ﬁres to clear

lands for wildlife use and hunting (Lewis 1985). The classic example of ﬁre to control

woody plant encroachment onto the tall-grass prairie, beneﬁtted bison among other

ungulate wildlife (Lewis 1985). Europeans as they colonized the Americas applied

their previous experiences generally relying on only conservation on game reserves

and limited hunting controls while generally lacking knowledge on how to manipulate

habitat to beneﬁt wildlife (Leopold 1933).

The early 1900s began an awakening for information on how to actively manage

wildlife, as populations of some wildlife species were declining, and public lands

were being overused. Land improvement began with soil conservation, forest and

grazing management. Leopold (1933) argued these were tools for managing and

improving wildlife habitat. He advocated concepts of plant successional theory of

the day and recognized land manipulations via planting, livestock grazing use and

non-use, ﬁre use and prevention, and mechanical tools (e.g., plowing, mowing, etc.)

for manipulating vegetation in the context of improving or sustaining wildlife habi-

tats. Recent additions to this toolbox include chemical and microbiological treatments

(Pyke et al. 2017). More recently, animal monitoring technology has been useful in

detailing information on what plant communities wildlife species use seasonally.

When managers couple wildlife use locations with functional and structural forma-

tions of plants into communities within landscapes, managers begin to understand

how speciﬁc manipulations may improve or decrease a wildlife s pecies’ population.

However, manipulations geared to beneﬁt one species in the ecosystem, may be

detrimental to others with differing habitat requirements (Fulbright et al. 2018).

Understanding animal movements, life history, and habitat use has been greatly

improved by the use of remote sensing and geographic information systems (GIS) that

allows managers to depict animal spatial movements over time. These assist managers

in understanding spatial and temporal elements of wildlife population dynamics and

in understanding the scale at which manipulations to landscapes, whether intended

for the beneﬁt of wildlife or not, may ultimately impact how wildlife use or avoid

certain habitats over time. Depending on the wildlife species even small human

inﬂuences, such as power poles, may create roosts for predators and result in potential

prey avoiding surrounding lands, even if the vegetation community provides the

necessary plant species composition to become sufﬁcient habitat (Leu and Hanser

2011). Therefore, it is important for managers to understand landscape scale impacts

of habitat manipulations.

In this chapter, we address important concepts relating to wildlife habitat manage-

ment in rangeland settings through manipulations of plant communities within spatial

5 Manipulation of Rangeland Wildlife Habitats 109

and temporal contexts that align with wildlife habitat requirements. Initially, we

deﬁne wildlife habitat in a spatial and temporal context that impacts habitat quan-

tity and quality and discuss the applied ecology of rangeland plant communities.

Lastly, we address the various types of manipulations typically used in rangelands.

Because livestock grazing systems and ﬁre are presented elsewhere in this book

(Chaps. 4 and 6, respectively), we will limit our discussion of these tools to their

uses in manipulating habitat.

5.2 Concepts

Across the world, ecosystems have been fundamentally altered due to current and

historical anthropogenic activities, and the rate of change is increasing (Millen-

nium Ecosystem Assessment 2005). Over the last 100 years, policy making for and

management of wildlife habitat in the United States has seen dramatic change with

respect to both speciﬁc issues and the general nature of natural resource management

challenges. Historically, such challenges have related strongly to easily identiﬁable

disruptions of ecosystem pattern and process that were amenable to policy-based

solutions (Grier 1982;Boydetal.

2014). While many such policies continue to

play a deﬁning role in topical management of wildlife habitat, new factors s uch as

climate change and its indirect effects have been associated with broader disrup-

tions of ecosystem processes, creating strong impetus for a more expanded notion of

conserving not just habitats of individual species, but the ecosystems in which those

habitats exist (Benson 2012; Evans et al. 2013). In this section, we synthesize tradi-

tional concepts in conservation of wildlife habitat and explore how these concepts

are developing and changing to meet a new generation of challenges facing stewards

of rangeland wildlife habitat.

5.2.1 What is Rangeland Wildlife Habitat?

In its most basic form, the term “habitat” represents where an animal lives, and

resources it uses while there. Those basic resources fall under the categories of

food, water, and cover, which are collectively used by animals to meet basic needs

including survival in the face of predation, amelioration of thermal stress, and meeting

nutritional demands of metabolic maintenance, growth, and reproduction.

Habitat needs of wildlife species play out within spatially and temporally variable

rangeland environments. Because of this variability, wildlife species must not only

occupy a home range that is large enough to contain the habitat needs described

above, but the size of that home range may vary in accordance with yearly condi-

tions (Anderson et al. 2005). Within an animals’ home range, different habitats may

be better suited to speciﬁc life history needs (e.g., breeding, summer, or winter

110 D. A. Pyke and C. S. Boyd

habitat). The spatial dispersion of these seasonal habitats can create seasonal move-

ment patterns within the larger home range (Connelly et al. 2011). The existence of

seasonal habitats, and movement between these habitats may be related to weather

and climate extremes (e.g., summer vs winter habitat) but is often associated with

spatio-temporal variability in plant phenology and production, in association with

temperature gradients (e.g., elevation) and rainfall distribution patterns (Holdo et al.

2009; Le Corre et al. 2017; Pratt et al. 2017). Anthropogenic factors such as infrastruc-

tural development and hunting activities can have strong inﬂuence on the geography

of movements between seasonal habitats (Gates et al. 2012; Amor et al. 2019).

Wildlife habitat can be thought of as occurring across a range of conceptual scales,

from the geographical range of a species to the within-site habitat characteristics

important to that species. These scales collectively represent a hierarchy of needs

wherein the importance of smaller scale habitat characteristics is predicated on the

existence of sufﬁcient habitat elements at larger scales (Johnson 1980). At the largest

practical management scale for most rangeland managers, landscape cover refers to

the dominant overhead cover components expressed as a fractional percentage of

landscape area. These data are useful both in large scale management planning and

for assessing links between habitat properties and populations for species with large

home ranges (Aldridge et al. 2008). Generally, landscape cover is measured through

remote sensing where the reﬂectance of vegetation functional groups (e.g., shrubs,

perennial grasses, etc.) or prominent species dominate the wavelengths of pixels in

images and are used as cover attributes in landscape analyses (e.g., Jones et al. 2018).

These data can also be collated to more broadly determine cover of higher order biotic

and plant associations (e.g., Brown et al. 2007). In addition, contemporaneous tech-

nology surrounding remotely sensed landscape cover is developing rapidly, allowing

for higher resolution data to detect individual species and biological soil crusts (Karl

et al. 2017). Moreover, data storage and retrieval technology has advanced to the

point that retrospective fractional cover estimates are now available going back to

the late 1980s using historical Landsat imagery (Allred et al. 2021) providing the

ability to track temporal variation over larger scales. These data also create a broad

spectrum of opportunities for both managers and researchers to retrospectively assess

the effectiveness of habitat treatment practices and relationships between landscape

cover attributes and population dynamics of wildlife species.

At local scales, a key attribute of habitat is to provide cover associated with

a diversity of needs including nesting, brood-rearing, fawning/calving, breeding,

roosting, and thermal regulation. Cover, generally in the form of vegetation, must

occur in sufﬁcient amounts to allow for species’ survival and reproduction. Cover may

act to decrease visibility of animals and nests (Conover et al. 2010), but can also act

to disrupt air circulation patterns and reduce the ability of predators to ﬁnd prey using

olfactory cues (Fogarty et al. 2017). Cover also acts as a barrier to thermal extremes

that could otherwise result in decreased ﬁtness or death of wildlife species. For

example, Guthery et al. (2001, 2010) reported that heat stress can result in decreased

breeding activity and even death of northern bobwhite (Colinus virginianus), and

that these consequences can be abated by habitat that serves as thermal refugia. At

the other end of the spectrum, cover can also act to mitigate physiological stresses

5 Manipulation of Rangeland Wildlife Habitats 111

of winter thermal extremes for ungulate species such as mule deer (Odocoileus

hemionus; Webb et al. 2013).

Cover for wildlife comes in two basic structural forms: horizontal and vertical.

Horizontal cover (also known as “horizontal foliar density”) refers to the degree

of interception created by vegetation when habitat is viewed in a horizontal plane.

The degree of interception will vary by height from ground level and the cumulative

horizontal cover proﬁle at a site is often referred to as “vertical structure” (Nudds

1977). Vertical structure can be a good predictor of habitat use by prey species

(e.g., Holbrook et al. 2016) and is also an important determinant of habitat selection

and reproductive success of many avian species (Hagen et al. 2007; Kennedy et al.

2009). Measuring vertical structure is accomplished via the use of a photoboard or

pole painted in contrasting bands; vegetation obstruction of the board or pole (Grifﬁth

and Youtie 1988) is determined at a ﬁxed distance using either digital photography or

ﬁeld estimates (Nudds 1977; Limb et al. 2007). Vertical, canopy, or foliar cover refers

to the amount of land surface area obscured by vegetation when viewed from above.

Canopy cover shapes wildlife habitat suitability through its inﬂuence on shading,

which effects thermal properties of the habitat (Guthery et al. 2010), understory plant

dynamics (Boyd and Bidwell 2002) and microenvironments (Royer et al. 2012), and

is also the primary attribute impacting the ability of a habitat to protect prey species

from overhead predators (Matthews et al. 2011). Canopy cover is also applied to

both vegetation and non-vegetational components of habitat such as rock and bare

ground, which can be important in describing both the ecological context of a habitat,

as well as habitat suitability for some species (Conway et al. 2012; Pyke et al. 2014).

In practice, the thermal and hiding cover afforded by a habitat will be a function of

species requirements and the interactive effect of both horizontal and vertical cover

attributes (Culbert et al. 2013).

A major function of an animal’s habitat is to provide energy and nutrients neces-

sary for survival, growth, and reproduction. Energy and nutrient sufﬁciency is a

function of both the nutrient requirements of a species, which are subject to temporal

variation in association with life history stage, as well as the dynamics of plant

species composition, nutrient quality, and production in space and time (see discus-

sion of the latter below). Links between animal performance at a given life history

stage and the nutrients/energy provided by the habitat can be both direct and indirect.

Nutritional limitations may directly induce weight loss, result in impaired growth

and development, and decrease reproductive success (Boyd et al. 1996), particularly

during periods of thermal extremes (DelGiudice et al. 1990, 1991). Insufﬁciency of

nutrients/energy may indirectly affect individual animals and perhaps populations

by negatively impacting physiological status of affected individuals and increasing

the likelihood of mortality from disease or predation (Lochmiller 1996). Abiotic

factors, such as thermal extremes or drought conditions can interactively exacerbate

effects of nutritional limitations of habitats by reducing nutrient/energy availability

and inducing physiological stress that increases an animal’s nutrient/energy demand

(Lochmiller 1996; Dabbert et al. 1997).

112 D. A. Pyke and C. S. Boyd

5.2.2 Climate, Weather, and Soil Inﬂuences on Rangeland

Communities

Climate and weather factors are critically important in determining plant community

responses to disturbance factors, as well as a plant community’s potential for restora-

tion success. In fact, weather, and to some extent climate variability, are the most

frequent “it depends” caveats associated with generalizations of rangeland treatment

effects or recovery trajectories of associated plant communities. Re-establishment of

desired vegetation following disturbance often fails in rangeland ecosystems (Pyke

et al. 2013) and the likelihood of success has been strongly tied to precipitation

amount (Hardegree et al. 2011), timing, and frequency (Pyle et al. 2021) relative to

the needs of seeded or recovering species, and all of the preceding factors interact

with soil temperature (James et al. 2019) to determine recovery outcome.

Climate and weather have strong effects on rangeland productivity and compo-

sition, and by extension, the manipulation of rangeland wildlife habitats. The term

“climate” refers to the long-term (e.g., averaged across years) patterns of precipi-

tation, temperature, and other atmospheric properties for a given location. Climate

differs from “weather” in that the latter refers to short-term variation (e.g., within

year or shorter) in these same properties. At the continental scale, inter-annual to

multi-decadal oscillations in temperature and precipitation are strongly inﬂuenced

by recognizable ocean temperature patterns and circulation (Wang 2021). These

ocean–atmosphere phenomena include the Paciﬁc Decadal Oscillation, the El Niño

Southern Oscillation, and the Atlantic Multidecadal Oscillation (McCabe et al. 2004;

Guilyardi et al. 2009). While mechanics of how ocean temperature patterns inﬂuence

terrestrial climate and weather are beyond the scope this chapter, both the effects and

occurrence of these patterns are somewhat predictable and have been incorporated

into management decision making on rangelands (e.g., Raynor et al. 2020). Climate

is also changing in association with greenhouse gas emissions; predicted changes in

climate, including more frequent droughts and severe weather, suggest that the inﬂu-

ence of climate and weather on rangeland plant community dynamics will increase

over time (Polley et al. 2017) and portend future challenges for management of range-

land plant communities and wildlife habitats. The extent to which ongoing climate

change via greenhouse gas emission is inﬂuencing the occurrence of ocean–atmo-

sphere phenomena is not well understood at present. That said, it is likely that some

of the effects of climate change on rangelands (e.g., increased air temperatures) could

interact with ocean-atmospheric associated events such as drought to decrease range-

land plant productivity (Schlaepfer et al. 2017). Alternatively, the ongoing increase

in atmospheric CO2 may be differentially increasing the production potential for

some plant species, leading to altered successional dynamics and the potential for

increasing rangeland fuel loads (Ziska et al. 2005). The bottom line is that substantial

uncertainty exists regarding interrelationships between future climate and rangeland

plant communities, reinforcing the need for active and adaptive management of

rangeland wildlife habitats.

5 Manipulation of Rangeland Wildlife Habitats 113

While climate factors associated with ocean-atmospheric events have some degree

of predictability, the predictability of shorter-term weather conditions relevant to

restoration projects or recovery from disturbance has proven more difﬁcult and

the useful accuracy of m ost forecasting techniques does not extend beyond 7–

10 days (Hardegree et al. 2018). That said, current seasonal climate forecasts provide

some level of generalization of weather conditions for periods up to several months

(Doblas-Reyes et al. 2013) and new, more restoration-oriented products are emerging

(e.g., Bradford and Andrews 2021).

While short-term forecasting of weather can be difﬁcult, qualitative generaliza-

tions of site-associated temperature and moisture potential can be assessed using

abiotic characteristics such as soils, elevation, slope, and aspect. For some range-

lands, soil temperature and moisture regimes have been used by managers to assess

the capacity for plant communities to both recover from disturbances such as ﬁre or

grazing (i.e., resilience), as well as their capacity to resist biotic change due to stres-

sors such as invasive plant species (i.e., resistance; Chambers et al. 2014, 2016a, b).

While these classiﬁcations can be useful from a management planning standpoint,

site speciﬁc management should take into account current variability in climate and

weather factors as well as biotic conditions of a site (Miller et al. 2014).

Soils quite literally form the biogeochemical foundation upon which rangeland

wildlife habitats and other ecosystem services are built, and speciﬁc soil proper-

ties have strong inﬂuence on both plant community composition, and the resulting

habitat structure (Evans et al. 2017). Soil texture is a fundamental property of the

soil environment and has a strong role in inﬂuencing water availability for plants.

Inﬁltration of water into the soil proﬁle decreases as soil particle size goes from

coarse to ﬁne (i.e., in order of decreasing particle size: sand, silt, clay; Lowery et al.

1996). Water inﬁltration into the soil not only provides a supply of water to plants

but also helps to prevent overland ﬂow and surface soil erosion (Evans et al. 2017).

The relationship between water holding capacity, or the ability of soil to trap and

hold water, is inverse to that of water inﬁltration, with ﬁner textured soils being more

capable of retaining water. The impact of trading water inﬁltration potential for water

holding capacity is moderated by annual precipitation. In arid regions, coarse soils

can decrease evaporative loss, which off-sets reduced water holding capacity and

increases water available to plants. In more mesic areas with less evaporative loss,

the increased water holding capacity of ﬁner textured soils results in increased soil

water available for plants (Austin et al. 2004; Evans et al. 2017). Soil organic matter

content is correlated positively with water holding capacity and can, to some extent,

moderate the effects of particle size on soil water storage.

Plant species distributions within rangeland habitats are also inﬂuenced by soil

chemistry. For example, saline soils support halophytic plants to the exclusion of non-

salt tolerant species, while shinnery oak (Quercus havardii) mottes can create acidic

soil conditions that approximate the pH of forest soils (Wiedeman and Pendound

1960). Soil pH, along with particle size and organic matter, can also modulate

the persistence and efﬁcacy of herbicides; although the speciﬁc effects are depen-

dent on herbicide type (Duncan and Scifres 1983). Lastly, soil depth can inﬂu-

ence water storage capacity of a site as well as competitive interactions between

114 D. A. Pyke and C. S. Boyd

plants. In general, soil water storage decreases, and competition for belowground

resources increases as depth to restrictive layer (e.g., bedrock) decreases; this accen-

tuates the importance of understanding soil characteristics in predicting manage-

ment outcomes. For example, Miller et al. (2005) reported that with sufﬁcient

rooting depth, perennial bunchgrasses were maintained during juniper (Juniperus

spp.) woodland expansion in sagebrush (Artemisia spp.) steppe habitat, but in shal-

lower soils bunchgrasses declined dramatically or were entirely absent with juniper

expansion.

5.2.3 Rangeland Vegetation Dynamics

Understanding how and why rangeland plant communities and the associated wildlife

habitats change over time allows managers to infer impacts on constituent wildlife,

anticipate and act on opportunities for habitat improvement, and mitigate undesired

changes. Change in rangeland plant communities can be broadly classiﬁed in terms

of equilibrium and non-equilibrium succession. Under the non-equilibrium succes-

sion paradigm, vegetation dynamics are driven by stochastic, abiotic factors (e.g.,

precipitation) and herbivore density rarely reaches the level necessary to have strong

impact on successional change in habitat conditions (Vetter 2005). In contrast, equi-

librium succession refers to the idea that changes in plant community composition are

mediated via density-dependent biotic feedbacks between herbivores (i.e., wildlife

or livestock) and plant communities they utilize as habitat.

These paradigms have strong implications to policies relating to land use and

management, and recognizing these differences is more than just an academic exer-

cise. For example, biotic control of successional processes suggests that policies

that control herbivore density (e.g., grazing regulations or wildlife harvest regu-

lations) will stimulate desired changes in habitat conditions. Alternatively, abiotic

control of succession would argue for policies that promote preemptive management

to increase rangeland plant community resilience to episodically-stressful environ-

mental conditions. While equilibrium dynamics undoubtedly play a role in succes-

sional change in some rangeland systems (particularly at small spatio-temporal

scales), non-equilibrium dynamics are now recognized as the driving force behind

plant succession in most rangeland ecosystems (Briske 2017).

Management toward or maintenance of desired habitat conditions involves using

speciﬁc tools or processes to manipulate vegetation composition and structure. Equi-

librium and non-equilibrium dynamics can have strong inﬂuences on the types of

problems or challenges managers must overcome and implications these problems or

challenges create for management planning and actions. Simple habitat management

problems are those problems with solutions that are relatively invariant in space and

time. From a habitat management standpoint, these problems are typically associ-

ated with plant communities undergoing equilibrium succession (Boyd and Svejcar

2009). For these problems, generalized solutions have broad management utility.

An example of a simple problem might be reducing shrub fuels in an equilibrium

5 Manipulation of Rangeland Wildlife Habitats 115

system using a brush-beating technique. Results of brush beating are likely to be

both successful and predictable in space and time (due to the equilibrial nature of

the system) to the extent that treating 4 ha is synonymous with reducing the size of

the problem by 4 ha for the effective life of the treatment.

Complex habitat management problems are those where the nature of the problem,

and by extension appropriate management actions, will vary depending on the

location and when the action will occur (i.e., space and time; Boyd and Svejcar

2009). Complex problems are usually associated with non-equilibrium succession.

For example, restoration of perennial plants in arid or semi-arid rangeland systems

is typically a complex problem. Choice of management techniques (or whether to

even attempt restoration) in such systems is driven strongly by abiotic factors such

as precipitation and temperature patterns that vary strongly in space and time. In

this case, generalized solutions do not have broad management utility. Instead,

habitat manipulations involving complex problems in non-equilibrium systems

require a diversity of management techniques and tools to cope with a diversity

of abiotically-driven habitat management challenges.

The process of setting habitat management objectives and selecting appropriate

management actions to achieve or maintain those conditions in non-equilibrium

systems can be guided by using state-and-transition models. State and transition

models (Stringham et al. 2003) describe a range of potential plant community phases

that dynamically shift in plant dominance or habitat structure within a relatively stable

state (Fig. 5.1). Shifts, also known as pathways, among community phases within

a state are generally viewed as reversable and inﬂuenced by both management and

non-management factors. Movements between states are known as transitions and are

relatively irreversible. Additionally, some states are sufﬁciently persistent that their

existence represents what could be considered a new “novel ecosystem” (DiTomaso

et al. 2017). For example, the invasion of exotic annual grasses in the Great Basin

region of the United States has created vast areas of rangeland with near-monoculture

abundance of these species. Because these species promote, and can persist in the

presence of increased wildﬁre, these annual grass-dominated areas are extremely

stable; some consider such areas to be novel ecosystems and suggest a management

focus that recognizes the ecology (and management implications) of this alternative

state as a new reference state (Davies et al. 2021).

Putting it all together, state and transition models represent an organized frame-

work for managing plant communities and their associated wildlife habitats in an

ecologically based manner. In reality, a seemingly inﬁnite number of states could be

present for a plant community assemblage because community phases are repre-

sented as static plant composition, but are merely a gradation of shifts in plant

dominance that occur annually. Thus, the goal of constructing state-and-transition

models for management is to assign this variability into as few states and phases as

necessary so the model is sufﬁciently practical for management use, while retaining

sufﬁcient complexity to represent ecologically important plant community dynamics.

The utility of these models for managing rangeland plant communities and their asso-

ciated wildlife habitats can be increased by assigning values to states that are consis-

tent with either measured population densities of target wildlife species (Holmes and

116 D. A. Pyke and C. S. Boyd

Fig. 5.1 Generalized shrub-grassland community with ﬁve vegetation states (dotted rectangles)

with community phases (solid rectangles) within each state. Pathways (solid arrows) depict shifts

in habitat dominance or structure within a state driven by biotic and abiotic inﬂuences. Transitions

(dashed arrows) depict relatively irreversible changes in habitat dominance or structure

5 Manipulation of Rangeland Wildlife Habitats 117

Miller 2010) or assigning qualitative values that represent the likelihood that habitat

structure and plant composition of a state will service year-long or seasonal habitat

needs of target wildlife species (Boyd et al. 2014).

For non-equilibrium rangeland wildlife habitats, knowledge of the plant commu-

nity’s resilience and resistance to disturbance will help deﬁne and guide management

options. In this case resilience is deﬁned as the capacity of ecosystems to reorganize

and regain their fundamental structure, processes, and functioning (i.e., to recover)

when altered by stressors like drought and disturbances such as ﬁre or inappropriate

livestock grazing (Holling 1973; Chambers et al. 2016a). Resistance, in turn can be

deﬁned as the capacity of ecosystems to retain their fundamental structure, processes,

and functioning when exposed to stress (e.g., invasive species) or disturbance (e.g.,

ﬁre; Folke et al. 2004; Chambers et al. 2016a). Characterizing the resilience and

resistance of rangeland wildlife habitats involves examining both the abiotic and

biotic environments. There are a host of abiotic factors that inﬂuence resilience and

resistance of plant communities including temperature, precipitation, and a wide

variety of soil factors. In practice a useful index to abiotic resilience and resistance

can be created by characterizing soil temperature and moisture regimes across the

area of interest into descriptive categories. For example, Chambers et al. (2014)

characterized resilience and resistance of plant communities within the sagebrush

ecosystem along a gradient from warm and dry to cold and moist; resilience and

resistance increase along this gradient in accordance with increasing elevation and

plant community productivity. These categories can be combined with habitat needs

of a species or groups of species and geospatially depicted to help guide habitat

management at broad spatial scales. For example, Chambers et al. (2016a, b) created

a matrix that included all combinations of low, medium, and high resilience and

resistance, combined with low, moderate, and high landscape cover of sagebrush.

The resulting cells of the matrix create categories that can be geospatially depicted

to guide management planning for the greater sage-grouse at large spatial scales

(Fig. 5.2).

Utility of using resilience and resistance to inform habitat management will be

increased by supplementing knowledge of contributing abiotic factors with current

assessments of biotic properties, particularly at the project implementation scale.

These biotic properties relate to the abundance of plant species within a community

that have disproportionately strong inﬂuence on resilience and resistance. A good

example is the inﬂuence that native perennial bunchgrasses have on resilience and

resistance of sagebrush plant communities. These species effectively occupy space

and utilize resources within the soil proﬁle such that their abundance is highly and

inversely correlated with probability of invasion by exotic annual grass species that

are prevalent throughout the sagebrush biome (Chambers et al. 2007;Davies

2008).

Thus, the abundance of perennial bunchgrasses can be used as a metric to identify

and prioritize for management those areas within a landscape that are most likely to

experience undesired change following disturbance. Additionally, the pre-treatment

abundance of these species can be used to gauge the potential for unintended and

undesired effects of active management treatments such as prescribed ﬁre (Bates

et al. 2000). At larger scales, assessment of biotic properties important to resilience

118 D. A. Pyke and C. S. Boyd

Fig. 5.2 Matrix depicting plant community resistance and resilience combined with landscape

suitability for greater sage-grouse habitat. Rows indicate generalized recovery potential (resilience)

and resistance to change during stress (e.g. exotic annual grass invasion). Increasing dominance

of the landscape by sagebrush (depicted in columns) broadly suggests increasing suitability for

greater sage-grouse. Cells within the matrix can be geospatially depicted to broadly inform decisions

regarding management of greater sage-grouse habitat. Taken from Chambers et al. (2017)

5 Manipulation of Rangeland Wildlife Habitats 119

and resistance will beneﬁt greatly from emerging geospatial technologies such as the

Rangeland Analysis Platform (Allred et al. 2021). These technologies not only allow

managers and researchers to assess the abundance of vegetation functional groups

across broad geographies, but can also be used to retroactively explore how plant

communities responded to disturbances and management treatments.

5.2.4 Point-Based Versus Process-Based Habitat

Management

One of the most basic challenges for contemporary rangeland wildlife habitat

managers is to determine the relative priorities associated with management of

ecosystem dysfunction versus the needs of individual species or groups of species of

concern, and determining where those priorities do and do not intersect. As discussed

earlier, the term “wildlife habitat” and by extension, wildlife habitat management,

is an inherently species-speciﬁc, and often a life-history phase-speciﬁc premise; for

example, we might use prescribed ﬁre as a tool to create plant community structure

suitable for nesting needs of black-capped vireo (Vireo atricapilla). Such manage-

ment has generally been tied to speciﬁc micro-habitat requirements representing

point-in-time vegetation conditions. We refer to this as point-based management

(Table 5.1). In the case of vireos, ﬁre can be beneﬁcial to nesting habitats because

this shrub-nesting species is picky about the height of shrubs in which it nests. When

shrubs become higher than a desired height, the habitat is no longer suitable for

nesting (Grzybowski 1995) and ﬁre can be used as a tool to reduce shrub height.

We can therefore think of point-based management as practices applied to speciﬁc

geographies that are intended to result in the ﬂoristic composition, structure, or

spatial arrangement of plant communities needed to meet speciﬁc habitat needs of

a species at a particular moment.

Point-based activities deﬁne much of our history with wildlife habitat manage-

ment on rangelands and the attraction to this type of management is multi-fold.

For example, point-based management is easy to administer where land ownership

boundaries deﬁne project areas (e.g., on private lands), and knowledge of species

habitat requirements provides a clear picture of desired changes to habitats, which

in turn suggests appropriate tools for the job. That said, if prescribed ﬁre is needed

to maintain proper nesting habitat for black-capped vireos, then how did this species

successfully evolve (i.e., it successfully nested and reproduced) within these habi-

tats for millennia? The answer probably relates to the fact that ﬁre frequency in

black-capped vireo habitat has decreased in modern times, creating conditions that

favor sustained growth of woody plant species (Grzybowski 1995). While point-

based management using prescribed ﬁre may indeed create beneﬁt to geographi-

cally speciﬁc vireo nesting habitats, managers should consider whether point-based

habitat deﬁciencies are merely symptomatic of higher order issues such as declining

ﬁre frequency. This is an important distinction because if local habitat deﬁciencies

120 D. A. Pyke and C. S. Boyd

Table 5.1 Contrasts between form-based and process-based approaches to management of

rangeland wildlife habitats

Characteristic Management type

Point-based Process-based

Goal Modify habitat conditions to align

with species habitat requirements

Modify ecosystem processes to

create enabling conditions that

inﬂuence desired future outcomes

Success metrics Direct management effects on habitat

composition and structure

Indirect management effects on

ecosystem processes

Spatial focus Plant community Landscape

Temporal focus Short term change Long term change

Diversity of

impact

Individual species or small groups of

species

Groups of species to species guilds

Frequency of

management

inputs

Opportunistic Persistent

are associated with disruption of ecosystem processes like ﬁre, then point-based

treatments may be creating islands of source habitat within landscapes that can act

as habitat sinks and may also serve to obfuscate or even disincentivize manage-

ment of ongoing system-level dysfunction, ultimately leading to reduced ecosystem

resilience (Hiers et al. 2016). Evaluating the importance of local vs. landscape factors

can be guided by frameworks (e.g., Pyke et al. 2015, 2017) that consider the spatial

ecology of primary threats to plant communities and associated habitats, home range

of the target species, types and locations of seasonal habitats, and the likely response

of target habitats based on abiotic characteristics.

When fundamental ecosystem issues such as disruptions in ﬁre frequency are

driving undesired changes to habitat of desired wildlife species, a different manage-

ment paradigm is required. In contrast to point-based management, the goal of

process-based management is to modify ecosystem processes to create enabling

conditions that inﬂuence desired future habitat attributes (Table 5.1). Effects of

process-based management will differ from point-management in that they are indi-

rect, often play out at relatively larger temporal and spatial scales, are more likely

to impact a larger number of species, and are likely to require persistent manage-

ment inputs over time. The need for a process-based approach to management of

wildlife habitats is becoming increasingly wide-spread due to both direct effects of

anthropogenic disturbance on ecosystem processes, and through the indirect effects

of climate change (Walker and Salt 2006).

A good example of process-based management would be the conservation of

low to mid elevation sagebrush habitats in the western US. The range of sagebrush

plant communities has decreased dramatically since European arrival due to a variety

of factors including agricultural conversions, oil and gas development, and housing

development. Within sagebrush habitats the spatial footprint of wildﬁre has increased

dramatically in recent decades, in part due to the dramatic expansion of exotic annual

5 Manipulation of Rangeland Wildlife Habitats 121

grass species such as cheatgrass (Bromus tectorum), which can create near continuous

coverage of ﬁne fuels that desiccate earlier in the growing season than native grasses;

effectively lengthening the ﬁre season. Sagebrush (Artemisia sp.) species within

the region are easily killed by ﬁre (Young and Evans 1978) and are difﬁcult to

restore following ﬁre (Mietier et al. 2018), creating a conservation crisis for a host of

sagebrush dependent wildlife species including the greater sage-grouse (Boyd et al.

2014). Improving habitat conditions for sagebrush dependent wildlife beneﬁts from

a process-based approach to create enabling conditions, namely treatments aimed at

reducing ﬁre occurrence and size, which allow for both active and passive restoration

of degraded habitats, and maintenance of intact habits. As noted above, in the absence

of enabling conditions, point-based restorative treatments run the risk of creating sink

habitats within dysfunctional landscapes, and the beneﬁts of successful point-based

restoration attempts are time limited in accordance with ﬁre dynamics (Boyd et al.

2017). Once enabling conditions have been achieved via process-based management,

point-based treatments can then be used to impact habitats within the landscape to

beneﬁt sagebrush dependent wildlife species (Pyke et al. 2015). This sequencing of

management emphases is consistent with hierarchical habitat selection by wildlife

species (Johnson 1980) and can help bring clarity to management planning and

increase effectiveness of conservation efforts in a growing diversity of complex and

dysfunctional ecosystems.

5.3 Landscape Context for Wildlife Habitat Manipulations

5.3.1 Rangeland Loss and Fragmentation

Rangeland by deﬁnition is “land supporting indigenous vegetation that either is

grazed or that has the potential to be grazed, and is managed as a natural ecosystem”

(SRM 1998). Changes in land uses may modify vegetation to maintain a desired

plant community that will beneﬁt the new land use or they can completely replace

the natural ecosystem with a simpliﬁed community of plants based on human desires

(e.g., crops). Exurban, suburban, and urban development provide decreasing levels

of natural plant communities with increasing levels of buildings and human infras-

tructure. In northeastern Colorado, ground- and shrub-nesting bird species diversity

declined in density with movement from rangeland to exurban developments and

while domesticated cats and dogs increased along the same gradient (Maestas et al.

2003). In 2019, half of the top ten states in percent population growth were states

with non-federal rural lands dominated by rangelands (Table 5.2). Current range-

land watersheds with the greatest projected housing development through 2030 are

around southern California cities, Las Vegas, Nevada and Phoenix, Arizona and will

result from exurban development (Reeves et al. 2018). Some of this exurban devel-

opment will lead to conversion of farmland to ranchettes, whereas rangelands are

then converted nearly simultaneously to farmlands (Emili and Greene 2014).

122 D. A. Pyke and C. S. Boyd

Table 5.2 Top ten states in percent growth of population between 2018 and 2019 (US Census

Bureau 2019;USDA 2020)

Rank State 2018 2019 Percent growth (%) Percent of rural land

that is rangeland (%)

1Idaho 1,750,536 1,787,065 2.1 36.7

2Nevada 3,027,341 3,080,156 1.7 85.1

3Arizona 7,158,024 7,278,717 1.7 82.9

4Utah 3,153,550 3,205,958 1.7 64.7

5Tex as 28,628,666 28,995,881 1.3 59.2

6South Carolina 5,084,156 5,148,714 1.3 0.0

7Washington 7,523,869 7,614,893 1.2 21.9

8Colorado 5,691,287 5,758,736 1.2 61.1

9Florida 21,244,317 21,477,737 1.1 9.9

10 North Carolina 10,381,615 10,488,084 1.0 0.0

There is a ﬂux between rangeland and farmland area in some locations of the

US due to economic ﬂuctuations of crop prices, disaster payment, and conserva-

tion incentive policies (e.g., Conservation Reserve Program) to convert farmland

to rangeland and the reverse with consequences to wildlife habitat and populations

(Rashford et al. 2011; Drummond et al. 2012; Smith et al. 2016;Larketal. 2020).

Coupled with human development comes the need for roads, irrigation, power and

water lines, fences and often changes in the vegetation. These manipulations create

the potential for wildlife habitat fragmentation even when they do not directly impact

the majority of rangeland plant communities (Reeves et al. 2018). This human foot-

print can have substantial impacts on some wildlife species (Leu and Hanser 2011).

For wildlife with large landscape patches of habitat, synanthropic predators of these

wildlife species may increase with greater human activities or infrastructures on the

landscape and threaten population survival of prey species. An example is increased

Corvid predation on greater sage-grouse nests with increased human activity or struc-

tures (Coates et al. 2016). Alternatively, direct losses of habitat for these wildlife prey

species may reduce land available for critical life history stages or may isolate their

populations through removals of corridors between habitat patches.

Past, current, and future social and economic needs have and will continue to shape

land uses, while new technologies may allow spatial placement of land manipula-

tions in habitat-friendly locations minimizing habitat losses while allowing resource

extraction or land uses. For example, horizonal drilling for oil and gas with multi-

bore well pads located on or near existing roads or human infrastructure corridors

(Thompson et al. 2015; Germaine et al. 2020) may minimize wildlife habitat impacts.

Livestock grazing occurs throughout rangelands and infrastructures to manage and

promote livestock production may also impact wildlife. In southern Alberta, Canada,

there are 77% more km of fence than all roads combined including unimproved

roads. For example, fences intended to impede movement of sheep will also impede

movement of pronghorns (Gates et al. 2012) and modelling indicates that fences

5 Manipulation of Rangeland Wildlife Habitats 123

restrict habitat area available to pronghorns (Reinking et al. 2019). Mineral licks,

both natural and human-placed licks are common attractants for wildlife (Kreulen

1985; Robbins 1993). Seasonal gestational beneﬁts of mineral licks are suspected

for some wild ungulates (Ayotte et al. 2006), however recent information indicates

they are potential locations for transmission of wildlife diseases (Payne et al. 2016;

Plummer et al. 2018). Seeps and spring development is another livestock-related

development that has potential beneﬁcial and detrimental impacts for wildlife. Water

developments of springs or seeps that capture and pipe water to troughs may result in

dewatering of these areas and in reducing the wetland vegetation associated with these

sites impacting wetland-dependent wildlife and insects especially in arid rangelands

(Parker et al. 2021). Well-designed water developments that spread water across

landscapes and are available to wildlife may have beneﬁts to some wildlife (Bleich

et al. 2005; Gurrieri 2020).

5.3.2 Broad-Scale Decisions

A review of the literature indicated that only about 10% of terrestrial restoration

projects considered landscape characteristics in locating projects (Gilby et al. 2018).

Considering landscape requirements and threats for wildlife species at a broad scale,

usually greater than a typical size of a restoration project (tens to hundreds of

hectares) can increase effectiveness of vegetation manipulations for creating habitat

that beneﬁts one or more populations of the species.

It is important to recognize that all wildlife species have broad and site scale

habitat needs while simultaneously recognizing that multiple species may overlap

in landscapes and coexist during certain times while other species may use the same

geographic locations, but at different times or seasons. For example, Garcia and

Armbruster (1997) evaluated the U.S. Bureau of Reclamation, Lonetree Wildlife

Management Unit in North Dakota for four proposed habitat manipulations to

improve gadwall (Mareca strepera) habitat while maintaining sharp-tailed grouse

(Tympanuchus phasianellus) habitat. They modelled four scenarios and incorporated

economic costs of manipulations into their results on gadwall and sharp-tail grouse.

Their model was limited to a few populations found on the r efuge. Other models

use a regional approach with multiple populations and varying habitats, but these are

rare (Doherty et al. 2016). Rarer still are models that consider optimal locations for

restoration across broad scales (Ricca et al. 2018; Ricca and Coates 2020).

Creating vegetation goals that meet the animal’s vegetation community and struc-

tural needs alone may not create wildlife habitat without considering other landscape

factors that may restrict the animal’s use or movement. For example, the habitat

manipulation goal for a shrub-obligate animal might be to clear trees that are roosting

habitat for predators and to create shrub habitat through releasing understory shrubs

and herbaceous vegetation from competition with trees. But if this cleared patch

is not connected to an adjacent shrub habitat without trees, the animal may never

use the treated area because there is no connection to safe habitat. The vegetation

124 D. A. Pyke and C. S. Boyd

objective of clearing trees and releasing shrubs and herbaceous vegetation could be

achieved, but the wildlife objective would not because the manager failed to consider

the connecting landscape of treeless area necessary to provide the animal access to

the cleared patch. A decision framework for landscape-level habitat manipulations

may assist in providing managers with queries to consider for optimizing animal

beneﬁts from habitat manipulations.

5.3.2.1 Does the Animal’s Population Cover a Broad Scale of Land

Types?

Afﬁrmative answers to one or more of the following questions will indicate the

potential that an animal’s range covers a broad scale of land types that some people

refer to as a landscape species.

1. Does the animal depend seasonally on multiple vegetation communities for

population survival?

2. Does the animal migrate seasonally?

3. Is the animal’s seasonal or annual home range larger than the typical manipulation

project?

4. Will habitat use of a manipulated area depend on current use of adjacent areas?

5. Will spatial gradients of environmental variables impact the achievement of

manipulation goals?

5.3.2.2 Deﬁne Regional or Broad Scale Landscape Objectives

for Habitat Manipulations

Landscape objectives should be deﬁned with the knowledge of how to monitor to

determine movement toward or away from the objective over time. These objectives

will likely deal with metrics obtained over large spatial or temporal scales. For vege-

tation components, remotely-sensed data is often used to determine changes in vege-

tation dominance over time and vegetation patch inter-relationships with surrounding

habitat patches. Coupled with vegetation metrics, it would be optimal to determine

any animal population or use objectives related to vegetation manipulations within

treated regions or landscapes (Pilliod et al. 2022). Examples of objectives include:

1. Increase connectivity among populations or seasonal habitat by 5% within the

region in 10 years.

2. Develop a system of ﬁre breaks to protect priority habitat and to maintain no net

loss of habitat and population levels in the region for the next 10 years.

5.3.2.3 Identify Components Necessary to Meet Landscape Objectives

Generally, managers do not have the capacity (physical or ﬁnancial) to restore all

landscapes and sites that require restoration within region. A priority structure will

5 Manipulation of Rangeland Wildlife Habitats 125

aid planning and hopefully target manipulations to locations within the region where

the likelihood of achieving objectives will be the greatest. This identiﬁcation process

is a triage of the entire landscape. The ﬁrst step in this process is to identify data

layers that deﬁne landscape or regional objectives for the habitat and the animal.

These objectives and data layers may include, but are not limited to:

1. Increasing connectivity among seasonal habitats or among separated popula-

tions might require maps of existing habitats and barriers for movement between

habitat or populations.

2. Conserving high quality habitat from future threats through risk maps of known

threats (e.g., ﬁre, invasive species, development, climate change).

3. Mapping potential habitat locations for beneﬁcial manipulations.

5.3.2.4 Identify Existing Habitat, Potential Habitat, and Wildlife

Population Trends Associated with Those Habitats

This stage provides data on the current state of populations across the landscape and

the habitat quality of those populations. This information is useful in determining

population strongholds where habitat connections might create new avenues for

genetic exchange among separated populations. Maps of current vegetation relative

to potential vegetation can aid decisions on where manipulations may produce habitat

and create corridors for population interchange. Knowledge of state and transition

successional models and maps of current and potential vegetation and of soils and

their associated descriptions of ecological dynamics may be useful at this stage.

5.3.2.5 Identify Landscapes with Locations that Best Meet Habitat

Criteria

This step is accomplished either through a series of map overlays to examine unions

of spatial criteria or through a series of models using these criteria. Typically, results

are a series of gradations illustrating locations where manipulations are likely to

beneﬁt populations on one end to those likely to negatively impact populations on

the other (Figs. 5.2 and 5.3). Information similar to Fig. 5.2 provides managers

with management options and potential outcomes, while Fig. 5.3 incorporates the

potential outcomes to the animal’s population given other factors that might regulate

the animal’s use of the landscape.

5.4 Site-Scale Habitat Manipulations

Rangeland manipulations conducted at speciﬁc sites may be intended for improving

habitat for wildlife or they may have other intended goals (e.g., livestock forage

production, fuel reduction, watershed health) that have wildlife consequences.

126 D. A. Pyke and C. S. Boyd

Fig. 5.3 The union of Greater Sage-grouse (GRSG) breeding habitat probabilities (A, B, C) with

sage-grouse habitat resilience and resistance (1, 2, 3) within each of the seven management zones

(MZ—dashed polygons) and Priority Conservation Areas (GRSG PAC) across the current range for

the GRSG in the USA. Taken from Chambers et al. (2017)

5 Manipulation of Rangeland Wildlife Habitats 127

These consequences can range from beneﬁcial to detrimental depending on the

type of manipulation, its extent and intensity, its location relative to other habitat

requirements, and the wildlife species.

Outcomes of rangeland manipulations will depend upon a variety of factors, such

as, treatment objectives, methods and conﬁguration, weather, climate, and post-

treatment management. Ideally, s ite-level habitat manipulations are formulated with

the idea of providing information useful for adaptive habitat management. We provide

six important considerations for a manipulation to be effective (Pyke et al. 2017).

5.4.1 Develop Site-Speciﬁc Management and Sampling

Objectives

Properly written objectives will provide the spatial and temporal elements of the

proposed effective habitat manipulation and will provide guidance for data collec-

tion and level of change necessary to determine manipulation success (effectiveness

monitoring). A properly written habitat manipulation objective typically includes the

following (Elzinga et al. 1998):

1. The target plant species, groups of species, or ecological conditions (e.g., a plant

species, all shrubs, or bare soil) that will be measured to determine success,

2. Location of the manipulation,

3. The measurement attribute (e.g., cover, density, height),

4. The action of change (e.g., increase, decrease, limit, or maintain),

5. The quantity or qualitative state of the anticipated change,

6. The time frame for success.

5.4.2 Consider Ecological Site Characteristics

Ecological sites comprise “a land classiﬁcation system that describes vege-

tation, ecological potential, and ecosystem dynamics of land areas” (https://

www.nrcs.usda.gov/wps/portal/nrcs/main/national/landuse/rangepasture/ Accessed

04/18/2021). This system of land classiﬁcation was developed by the U.S. Natural

Resources Conservation Service and has become standardized for use across multiple

Federal land management agencies (Caudle et al. 2013). An individual ecological site

is “a distinctive kind of land with speciﬁc soil and physical characteristics that differ

from other kinds of land in its ability to produce a distinctive kind and amount of vege-

tation and its ability to respond similarly to management actions and natural distur-

bances” (https://www.nrcs.usda.gov/wps/portal/nrcs/detail/national/landuse/rangep

asture/?cid=stelprdb1068392 Accessed 04/17/2021).

Ecological site data for a location are identiﬁed through the Web Soil Survey

(https://websoilsurvey.nrcs.usda.gov/ Accessed 04/17/2021) where an i nteractive

map allows the user to outline an area of interest for the habitat manipulation.

128 D. A. Pyke and C. S. Boyd

Speciﬁc ecological site descriptions (ESD) of individual ecological sites are found

at the Ecosystem Dynamics Interpretive Tool (EDIT; https://edit.jornada.nmsu.edu/

catalogs/esd Accessed 04/17/2021). Each ESD includes a state and transition model

for the ecological site that describes stressors that may shift vegetation dynamics to

alternative stable states.

State and transition models provide information on the vegetation community

dominance in plant community phases in the reference state and in alternative states.

The current vegetation at the manipulation site is compared to these ranges of plant

communities in the array of states in the state and transition models to determine the

potential for a habitat manipulation to achieve the habitat objective. Manipulations

that may drive the community to one of the phases in the references state, as opposed

to those in an alternative state, are most likely to provide the greatest resilience to

further disturbances and resistance to invasive plants (Chambers et al. 2017).

5.4.3 Determine Land Use and Disturbance History

Past, present, and future land uses and the previous disturbance history may provide

managers with information regarding the time period necessary for successfully

achieving habitat objectives. In some cases, previous disturbances or land uses

may have led to the current vegetation at the site and may require changes in

these uses to achieve the objective. Before implementing a manipulation to a

site, managers might consider if previous manipulations have been done to the

site and if those were successful. On Bureau of Land Management property, the

Land Treatment Digital Library provides available spatial information on historic

land manipulations and reports on their success in meeting objectives (https://ltdl.

wr.usgs.gov/ Accessed 04/17/2021). Some disturbances may have led to a loss

of ecological potential through the loss of soil erosion as an example. This loss

of potential may determine whether the proposed manipulation can create the

proposed habitat. Another GIS-based tool to assist managers in making decisions

to move forward with manipulations at a proposed site is the Land Treatment Explo-

ration Tool (https://www.usgs.gov/centers/forest-and-rangeland-ecosystem-science-

center/science/land-treatment-exploration-tool Accessed 04/17/2021).

In addition, Interpreting Indicators of Rangeland Health (Pellant et al. 2020)is

a fast on-site assessment of the soil, hydrology, and biotic potential that can assist

managers in determining if site potential has been lost. Ratings of departures from

reference conditions (the potential for the site) that are more severe than moderate,

especially for soil and site stability and hydrologic function, may provide an indica-

tion that attaining the ecological potential for this location may not be possible; even

with revegetation, the soil or water on the site may no longer function at a level that

can support the potential vegetation and managers may be left with alternative states

and with questions if desirable habitat can be created with ecological processes in

which the site contains.

5 Manipulation of Rangeland Wildlife Habitats 129

5.4.4 Consider the Role of Pre- and Post-treatment Weather

Weather is a critical element in regulating plant responses, but it is outside the control

of the manager attempting to modify or create wildlife habitat. The weather before

a habitat manipulation may dictate existing plant’s vigor which relates to nutrient

status of the plant and the storage of nutrients in stems and roots immediately before

a manipulation that may partially cut or damage plants requiring regrowth after the

disturbance. If the manipulation is intended to reduce the damaged plant for as long as

possible, then weather before the disturbance that reduces the plant’s vigor may delay

regrowth and extend the habitat objective, such as reducing woody plants. However,

if the objective is to increase a group of plants through growth or seed production

and establishment, but plants are in poor vigor, then the disturbance may not achieve

its objective (Hardegree et al. 2012). In the future, models may incorporate past

weather and future weather predictions to assist in projecting plant responses to

habitat manipulation (Hardegree et al. 2016).

5.4.5 Evaluate Plant Removal Methods and Associated

Effects

5.4.5.1 Passive Manipulations

Passive forms of manipulations generally involve changes in current land manage-

ment with an expectation that plant community dynamics will respond with changes

in plant dominance to create the desired wildlife habitat. For example, changes in

livestock management may include changes in stocking rates including elimination

of use, changes in livestock periods of use, distribution, or changes in the type of

livestock grazing the area.

Targeted grazing is a passive form of manipulation where a class of animal

grazes for a set season and duration at a set stocking rate to shift plant species or

lifeform dominance in an area (Frost and Launchbaugh 2003; Bailey et al. 2019).

Targeted grazing for fuel reductions typically require fencing or herding animals to

graze live and standing dead plants that may become fuels for wildﬁres (Fig. 5.4a).

In addition, animals can learn to feed on plants they may not prefer normally or to

avoid plants they may normally prefer through conscious and subconscious learning.

This type of targeted grazing requires diet conditioning (i.e., training). Conditioning

is a natural process that young animals learn from their mothers in utero or from

milk and then is reinforced by following their mothers and eating the same foods

while experiencing similar ﬂavors and nutritional responses (Nolte and Provenza

1992; Nolte et al. 1992). Diet conditioning can also be used to teach animals to avoid

certain plants (Lane et al. 1990) or novel plants that are previously unknown (Walker

et al. 1992; Dietz et al. 2010). Supplements with polyethylene glycol, protein, and

130 D. A. Pyke and C. S. Boyd

energy may increase the use of some plants by animals, but these are species- and

animal-speciﬁc (Bailey et al. 2019).

Some evidence suggests that livestock grazing before wildlife arrive to an area

may increase the wildlife forage use of the area. Bailey et al. (2019) document several

studies indicating that livestock grazing improves forage for wildlife. However, most

studies only documented the improved nutrient levels of the forage, not increased

wildlife use of these locations. Crane et al. (2016) provides an exception by demon-

strating increased elk use in areas previously grazed vs. ungrazed by cattle. There

are many hypotheses for creating habitats through restoration and manipulation of

the current environment, but the full set of ecosystem complexities are rarely tested

(Hilderbrand et al. 2005). When manipulating a community to create habitat for

5 Manipulation of Rangeland Wildlife Habitats 131

◄Fig. 5.4 a Targeted Grazing—Cattle being used to graze cheatgrass in Nebraska to reduce

cheatgrass seed production and population and help recovery of mid-grass prairie. b Prescribed

Fire—Ruby Lake National Wildlife Refuge uses prescribed ﬁre to reduce undesirable plants

and release desirable vegetation for waterfowl. Mechanical Removal—c Bull Hog masticating

juniper tree in Utah and d Cut, drop and leave is one form of woody plant removal practiced

in Oregon’s Bureau of Land Management lands. e Pelleted herbicide tebuthiuron being used to

thin shrubs in Washington. This same method is used to aerially broadcast seeds for restora-

tion of desirable plants. f Biocontrol—Salt Cedar (Tamarix chinensis) has infested many riparian

areas of the Southwestern US, but introductions of tamarisk leaf beetle (Diorhabda elongata)

(inset) often control this invasive plant. Photo Credits. 4A. Julie Kray, USDA ARS, Fort Collins

Colorado—Photo is in Scottsbluff, Nebraska. 4B. US Fish and Wildlife Service—Ruby Lake

National Wildlife Refuge, Ruby Valley, Nevada https://usfws.medium.com/using-prescribed-ﬁre-

to-improve-habitat-and-save-wildlife-c836453d51b0. 4C. Onaqui, Utah SageSTEP Project site—

Photo by Brad Jessop, Bureau of Land Management Utah 2006. 4D. Middle of Nevada—Photo

taken on June, 2011, Natural Resources Conservation Service media folder—https://www.nrcs.

usda.gov/Internet/FSE_MEDIA/nrcs144p2_036837.jpg. 4E. Moses Coulee SageSTEP project site,

Washington—Photo taken by Scott Shaff, U.S. Geological Survey—November 24, 2008. 4F.

Photo and inset photo from Glen Canyon National Recreation Area, Utah—Photo National Park

Service Photo—Date unknown for both photos. Main Photo—https://www.nps.gov/glca/learn/

nature/images/Tamarisk-Minimally-Impacted-by-TLB-web.jpg;inset—https://www.nps.gov/glca/

learn/nature/images/saltcedar-leaf-beetle.JPG

wildlife, monitoring for wildlife use and ultimately population trends would be

helpful for adaptive management (Pilliod et al. 2022).

5.4.5.2 Active Manipulations

Active manipulations are necessary when passive management changes and succes-

sional processes are inadequate to meet objectives, whether for wildlife or for

other reasons. Active manipulations include ﬁre-, mechanical-, and chemical/

biological/microbial-induced modiﬁcations to physical or biological components of

the ecosystem (Fig. 5.4b–f).

Prescribed ﬁres can be useful tools when they remove or reduce undesirable

vegetation and encourage growth and dominance of desirable plants while not making

the community vulnerable to undesirable physical or biological components of the

ecosystem (e.g., soil erosion, hydrophobic soils, invasive plants). Tolerance of and

susceptibility to ﬁre depends on whether the entire plant is consumed by ﬁre and can

regrow after a ﬁre (Pyke et al. 2010). The Fire Effects Information System (FEIS;

https://www.feis-crs.org/feis/ Accessed 21 April 2021) provides information on the

susceptibility of individual plant species to ﬁre; useful information for a manager

deciding whether to use ﬁre for creating habitat.

Prescribed ﬁres are modulated through adjustments in ﬁre: (1) intensity by manip-

ulating fuel amount and packing, (2) duration by the size and cellular density of fuel

or by the ﬁre type (e.g., surface vs. crown ﬁre or backing vs. head ﬁres), (3) extent

and patchiness of burned areas (Pyke et al. 2017). The heat created (intensity) by

ﬁre and duration of that heat will determine its effect on plants and seeds (Whelan

132 D. A. Pyke and C. S. Boyd

1995). Larger ﬁres that kill plants with limited seed banks or regrowth mechanisms

will increase the time required for those plants to disperse to the site and recover,

especially for plants with limited dispersal mechanisms. Consult with trained ﬁre

manager in developing ﬁre objectives to meet habitat objectives.

Mechanical and Chemical manipulations (e.g., Fig. 5.4c–e) use several potential

pieces of equipment to modify vegetation on rangelands (https://greatbasinﬁrescie

nce.org/revegetation-equipment-catalog-draft/ Accessed 21 April 2021). Methods of

habitat manipulations can range from those that remove all plants to those that are

more selective for removing or thinning species or lifeforms. Mechanical equipment

that operates entirely above the soil surface is intended to remove or reduce height and

cover of vegetation. Shrubs with limited resprouting ability or without adventitious or

perennating buds on remaining live, woody tissue will be reduced in dominance more

than those with these resilience mechanisms; similar to the effect of ﬁre (Pyke et al.

2010). The FEIS provides information on resprouting ability of plants. Mechanical

equipment that digs into the soil kills or reduces the dominance of all plant life forms

impacted with the exception of plants with strong adventitious buds on roots or

rhizomes. Some equipment, such as tractor-pulled anchor chains, not only removes

large trees and shrubs, but also remove some herbaceous plants ( grasses, grasslike and

forbs) when they dig into the ground. Plows and harrows cause similar effects. These

areas of soil exposure may result in soil erosion and invasive plant establishment and

spread, especially in years immediately after treatment. Before treatment, consider

if invasive plants already exist on the site and might increase and spread with soil

disturbing treatments. Seeding with desirable plants and using herbicides focused on

invasive plants may be necessary to limit invasive species and encourage desirable

plant establishment and growth.

Miller et al. (2014) suggests considering a series of questions to weigh the mone-

tary and ecological costs and beneﬁts of using mechanical treatments to manipu-

late plant communities. These include: (1) will equipment create unacceptable soil

compaction? Wet, ﬁne-textured soils are more susceptible to compaction than dry,

course-textured soils. Mechanical manipulations in the dry season or when soils are

frozen may reduce the severity of soil compaction. (2) Will the mechanical manip-

ulation create unacceptable amounts of mineral soil exposed to raindrop impact and

will these patches be on steep slopes? Bare soil is vulnerable to invasions of undesir-

able plants and to soil erosion. Larger patches of bare soil are susceptible to wind- or

water-induced erosion, whereas the steeper the land’s slope, the greater the potential

for water-induced soil erosion. (3) Will the manipulation disturb biological soil crusts

(biocrusts)? Biocrusts are soil surface lichens, mosses, algae, and cyanobacteria that

adhere to soil particles and protect soil from wind- and water-induced erosion. In

some arid and semi-arid environments, biocrusts can also ﬁx nitrogen for use by

other organisms in the ecosystem (Belnap and Lange 2003). (4) Will the mechan-

ical treatment damage existing perennial grasses and forbs? If the intention of the

mechanical treatment is to reduce woody plants, then the resilience of the remaining

plant community constituents and their resistance to invasive plants is important.

If the mechanical treatment impacts community components that are necessary for

community resilience and resistance, then the resulting community after the treatment

5 Manipulation of Rangeland Wildlife Habitats 133

may achieve its objective of reducing woody plants, but may ultimately degrade the

site through loss of soil or reduced hydrologic capacity. (5) Will the treatment provide

a seedbed for seedling establishment? If a mechanical treatment is accompanied by

a reseeding treatment, then a seedbed for seedling establishment is important, but

recognize that if the community already has invasive plants, the mechanical treatment

may enhance invasive plant establishment and create a competitive environment for

the reseeded desirable plants. (6) Will changing the timing of treatments inﬂuence

plant response positively or negatively? Consider what is the optimum manipulation

time to reduce potential negative and maximize positive outcomes.

Herbicides can be selective, affecting only certain plant life forms, or non-selective

(broad-spectrum) potentially affecting all plant life forms. Some broad-spectrum

herbicides can become selective for certain plant groups by manipulating the timing

or application rate. In addition, each herbicide is registered for uses on different

types of lands. Be certain when selecting an herbicide that it is registered for use

on rangelands and follow all label instructions. New herbicides are being tested and

released annually. Work closely with a licensed herbicide applicator in selecting,

planning, and applying an herbicide.

Herbicides rarely eradicate a target plant species or group, but they often reduce

targeted species for a period of time. The removal of a target plant will often leave

a void for other plants to ﬁll. If desirable plants do not ﬁll those vegetation gaps,

undesirable plants, even the original target plant, may re-establish and dominate the

site. Seeding chemically treated areas with desirable vegetation may be necessary in

environments where residual vegetation is not sufﬁcient to ﬁll voids left by removed

vegetation.

Biocontrols are sometimes used to reduce undesirable plants (McFadyen 1998).

Targeted grazing is a form of biological control, but insects are the most common

form of biocontrol of weedy plants. In addition, biocontrols can include microbial

pathogens (e.g., fungi, bacteria, and viruses; Harding and Raizada 2015). Insects are

generally released by hand at a site, while microbes are often applied using methods

similar to herbicide applications since they can be mixed with water, pelletized, or

coated on seeds or degradable inert biological forms such as rice hulls.

Effectiveness of biocontrols has been variable. Effective biocontrols generally do

not eradicate the target plant. Complete elimination of the target would likely erad-

icate the biocontrol agent too. Therefore, biocontrols may reduce undesirable plant

species to low levels and should the target plant increase, the biocontrol’s population

would ideally increase as their food source increases. Provided the biocontrol agent

reduces the target plant, a concomitant objective should be for desirable vegetation

to increase to ﬁll the void left through the death of the undesirable plant.

Revegetation (Figs. 5.4e and 5.5a–d) is used when desirable vegetation popula-

tions are insufﬁcient to provide propagules to ﬁll the void in an adequate timeframe

after undesirable plants are removed. The timeframe will vary depending on the

site’s resilience and resistance; sites with low values often need propagules to estab-

lish and dominate in less than ten years and those with high values having larger

timeframes. Managers may consider whether to seed or plant juvenile plants. Plant

species selected for creating wildlife habitat through revegetation is a union of the

134 D. A. Pyke and C. S. Boyd

group of plants deﬁned as habitat species and plants that have the potential for

existing and successfully reproducing on the site. The best source of information for

selecting native s pecies is the ecological site description for the site. Examine the

plant community phases found in the state and transition model and select the plant

community phase that matches the ideal life-forms to provide habitat composition

and structure for the target wildlife species (e.g., trees, shrubs, grasses and grass-like

and forbs). Include in the revegetation mixture plant species that would dominate the

site and are currently in insufﬁcient numbers for the site.

The geographic source of the propagule used in a revegetation project is important

for establishment and for sustaining future generations of plants on the site. Foresters

have known for decades that seed source is important for matching a tree’s genetics

to the environment where it will be grown (Johnson et al. 2004). They use seed

zones for collecting and planting reforestation projects. Rangeland provisional seed

zones are proposed for some r egions (Bower et al. 2014; https://www.fs.fed.us/wwe

tac/threat-map/TRMSeedZoneData.php Accessed 04/23/2021) and when used may

improve revegetation success. Climate change has sparked considerations for using

assisted migration techniques to move species or ecotypes within species from lower

to higher elevations or latitudes (Loss et al. 2011). Although these approaches have

been considered hypothetically, they are mostly in the testing phases (Wang et al.

2019).

Plantings and Seedings After selecting the species and propagule source, the type

of revegetation method is determined. Seedings are either broadcasted (Fig. 5.4e;

aerial or ground-based) or drilled (Fig. 5.5a, b). Plantings can come in several forms

(Shaw 2004) and are most often conducted with woody species. Small container-

grown plants are started in greenhouses, hardened to the environment, and trans-

planted at the site with their roots contained within a potting soil. Bare-root plantings

are initially grown in gardens in a loose compost soils, then the plant and roots are

extracted from the soil immediately before planting at the revegetation site. Cuttings

of shrub branches are taken from live plants and the cut branch is planted in the soil

and allowed to root. This is a common technique for shrubs in riparian areas because

branches can produce adventitious roots in moist soil. Wildings are small plants

extracted, with their soil, from an existing site and planted at a new location. This is

a good approach for salvaging plants that might be destroyed where human devel-

opment would require plant removal before development. Planting techniques are

often labor intensive, but may provide greater establishment than plants germinating

and establishing from seeds.

Seeding projects are the most common form of revegetation (Hardegree et al.

2011; Pilliod et al. 2017). Drill seeding is generally considered the most successful

seeding method because the seed drill places seeds at the appropriate depth in the

soil for germination and emergence of the seedling. Broadcasting seeds, when used

alone without other soil disturbing techniques (e.g., anchor chains, or harrows),

places seeds on or slightly above (if litter exists) the soil surface where they are

vulnerable to predation or displacement by wind and water (Stevens and Monsen

2004). Drill seeding often requires some site preparation (e.g., ﬁre) to remove any

larger woody vegetation that would limit the use of a tractor or would bind in the

5 Manipulation of Rangeland Wildlife Habitats 135

Fig. 5.5 Rangeland drills are designed to seed multiple species at different depths of soil. Tradi-

tional rangeland drill (a) that places seeds in furrows (c). In contrast, minimum-till rangeland drill

(b)leavesthe soil ﬂat(d) after placing seeds. Photo credits. a, b, c and d. Location likely Mountain

Home, Idaho in 2006. Photo by US Forest Service. Image currently on Great Basin Fire Science

Exchange, Revegetation Equipment Catalog, but originally in Joint Fire Science Final Report,

Project #07-1-3-12 by Dr. Nancy Shaw, USFS, https://www.fs.usda.gov/rm/pubs_other/rmrs_2011_

shaw_n003.pdf. Photo now found on https://revegetation.greatbasinﬁrescience.org/wp-content/upl

oads/2021/01/LRangelandDrillRightMinTillDrill_SoilDisturbance_USFS-294x300.jpg

seed drill. If tractors and drills are limited by obstacles or terrain, aerial seeding is

the best seeding method.

Emerging seeding technologies are being tested and may prove helpful in

increasing seedling emergence, establishment, and competition with invasive species

and decreasing seed predation. Coating seeds with hormones to hasten or delay germi-

nation may insure that germination occurs at the ideal time of the season or may

allow a bet-hedging strategy with seeds germinating over a longer timeframe than

136 D. A. Pyke and C. S. Boyd

normal (Madsen et al. 2016, 2018; Davies et al. 2018). Seeds encompassed in pellets

with activated carbon may allow simultaneous herbicide applications of preemergent

herbicides to reduce invasive plants while the pellet absorbs and retains the herbi-

cide allowing safe germination of desired species (Brown et al. 2019). Coating seeds

with materials that prevent animals from eating seeds may alleviate seed predation

common with broadcast seeds (Pearson et al. 2019).

Restoration of biocrusts is another emerging ﬁeld that may become common for

arid and semiarid environments where biocrusts are an important ecosystem compo-

nent for rangeland health. Biocrust production and application are most common for

cyanobacteria that can be commercially increased for applications, whereas research

for moss and lichen restoration is in its infancy (Antoninka et al. 2020).

5.4.6 Effectiveness Monitoring for Adaptive Resource

Management

Adaptive resource management (ARM) is an evolutionary process where the best

management decisions are enacted to achieve desired outcomes (i.e. objectives) and

the outcomes are tested (i.e., monitored) along with environmental variables that

may inﬂuence outcomes to determine their effectiveness at one or more timeframes

and across numerous similar sites. If objectives were not met and an alternative

management action is suspected to improve achieving objectives, then the alterna-

tive is enacted and the process is repeated (Fig. 5.6; Reever-Morghan et al. 2006;

Williams et al. 2009; Pilliod et al. 2021). Rangeland manipulations applied to lands

with a goal of improving wildlife habitat should incorporate monitoring the habitat

and the associated wildlife populations to determine if the predicted habitat was

achieved and if wildlife populations are responding in the predicted manner (Pilliod

et al. 2021, 2022). This is not a trivial component of manipulations and often is an

expensive, time-consuming component that requires adequate planning and funds to

accomplish. When done correctly, ARM will incorporate data from multiple sites

using compatible methods and producing adjustments to the previous manipulation

model or to formulate alternative models to improve effectiveness of manipulations

to produce wildlife habitat.

5.5 Conclusions

Manipulations of rangeland ecosystems in the twenty-ﬁrst century should not be

viewed as impacting singular resources, but rather consider the complexity of the

resource being manipulated and multiple physical and biological components that

may respond to manipulations. Wildlife species may use multiple types of habitats

across large landscapes or they may limit use to a narrow range of conditions in

5 Manipulation of Rangeland Wildlife Habitats 137

Fig. 5.6 Adaptive management begins with assessing the ecological status of current plant commu-

nities as well as factors that limit succession to a more desirable state. In management planning,

objectives are formulated, and important ecological processes determined. These processes suggest

speciﬁc management tactics. This information is translated into a spatially explicit plan (Plan Imple-

mentation) that indicates what will be done and where it will occur on the landscape. Following

plan implementation, research and monitoring are used to evaluate management impacts and assess

the validity of assumptions made in the planning process. This adaptive management process links

different conservation elements into an iterative cycle of planning, doing, and learning that allows

for management in the face of uncertainty and is necessary when managing complex problems in

non-equilibrium rangeland ecosystems

small isolated environments. Regardless, rangeland manipulations at a local scale

may inﬂuence wildlife at larger spatial extents, therefore, it is important to consider

broad-scale responses even when manipulations are focused at local levels to prevent

unintended consequences to wildlife species from the applied manipulation. Range-

land managers have many tools for planning and implement manipulations with more

tools arriving in the future.

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Manipulation of Rangeland Wildlife Habitats

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