Forest

Service

United States Department of Agriculture

General Technical Report 

RMRS-GTR-388

Rocky Mountain

Research Station December 2018

Multispecies Mesocarnivore Monitoring: 

USDA Forest Service Multiregional Monitoring Approach

Jessie D. Golding

Michael K. Schwartz

Kevin S. McKelvey

John R. Squires

Scott D. Jackson

Cara Staab

Rema B. Sadak
Golding, Jessie D.; Schwartz, Michael K.; McKelvey, Kevin S.; Squires, John R.; Jackson, Scott D.; 

Staab, Cara; Sadak, Rema B. 2018. Multispecies mesocarnivore monitoring: USDA Forest 

Service multiregional monitoring approach. Gen. Tech. Rep. RMRS-GTR-388. Fort Collins, 

CO: U.S, Department of Agriculture, Forest Service, Rocky Mountain Research Station. 68 p.

Abstract 

Small- to mid-sized forest carnivores, also known as mesocarnivores, are an important part 

of the animal community within national forests. Many forest mesocarnivores are of conservation 

concern and are listed as threatened or endangered under the Endangered Species Act (ESA), 

have been petitioned for listing under the ESA multiple times, or have designations within the 

Forest Service that warrant consideration in decisions about planning, projects, or restoration. 

Mesocarnivores also receive heightened public attention and, as a result, are frequently the center 

of lawsuits brought against the Forest Service. However, there is no current monitoring framework 

in place to provide meaningful information about these species across larger scales. In addition, 

they are difficult to detect, occur in low densities, and have large home ranges. 

We propose an approach for monitoring fisher (Pekania pennanti), Canada lynx (Lynx 

canadensis), American marten (Martes americana), Pacific marten (M. caurina), montane red 

fox (Vulpes vulpes sspp.), and wolverine (Gulo gulo) across the western United States. This 

approach was developed with close collaboration between the Forest Service Rocky Mountain 

Research Station and the National Forest System (NFS), and focuses on answering three basic 

monitoring questions: (1) Is a species present? (2) Are multiple individuals of a single sex present? 

and (3) Are multiple individuals, including both sexes, present? To answer these questions we 

designed a goal efficient monitoring (GEM) framework with four occupancy states related to a rare 

species population. We developed a Bayesian multistate dynamic occupancy model to analyze 

this information over time and estimate the probability that a population is likely to remain in one 

of these four occupancy states or transition to a different state. This document elucidates the 

process that led to the decision to use this framework and outlines the conceptual basis for GEM. 

A practitioner’s guide with detailed GEM implementation instructions will follow in a subsequent 

publication. 

Keywords: fisher, lynx, marten, montane red fox, wolverine, multispecies monitoring, 

 forest carnivore, multistate model

Cover photos

From left, wolverine (photo: Jeff Copeland, The Wolverine Foundation, used with permission), lynx 

(photo: Milo Bircham, USDA Forest Service), and fisher (Michael Schwartz, USDA Forest Service). 

All Rocky Mountain Research Station publications are published by U.S. Forest Service employees and 

are in the public domain and available at no cost. Even though U.S. Forest Service publications are not 

copyrighted, they are formatted according to U.S. Department of Agriculture standards and research 

findings and formatting cannot be altered in reprints. Altering content or formatting, including the cover 

and title page, is strictly prohibited.
Authors

Jessie D. Golding is a Carnivore Research Associate with the USDA Forest Service, Rocky 

Mountain Research Station in Missoula, Montana. She received a B.A. degree in biology and 

environmental science from the University of California, Berkeley, and an M.S. degree in wildlife 

biology from the University of Montana.

Michael K. Schwartz is the Director of the National Genomics Center for Wildlife and Fish 

Conservation and Research in Missoula, Montana. He received a B.A. degree in psychology from 

Colby College, an M.S. degree in ecology and evolution from American University, and a Ph.D. in 

wildlife biology from the University of Montana.

Kevin S. McKelvey is a Research Ecologist with the Rocky Mountain Research Station in 

Missoula, Montana. He received a B.A. degree in history and an M.S. degree in forestry from the 

University of Montana, and a Ph.D. in forest ecology from the University of Florida.

John R. Squires is a Research Wildlife Biologist with the Rocky Mountain Research Station in 

Missoula, Montana. He received a B.S. degree in wildlife biology from Colorado State University, 

and an M.S. degree and Ph.D. in zoology from the University of Wyoming.

Scott D. Jackson is the USDA Forest Service’s National Carnivore Program Leader in Missoula, 

Montana. He received a B.A. degree in biology/environmental studies from Augustana College and 

an M.S. degree in fish and wildlife management from Montana State University.

Cara Staab is a Regional Wildlife Ecologist in the National Forest System, Northern Region 

(Region 1) in Missoula, Montana. She received a B.S. degree in biology from Northern Arizona 

University and an M.S. degree in wildlife and fisheries science from the University of Arizona.

Rema B. Sadak is a Regional Wildlife Ecologist in the National Forest System, Intermountain 

Region (Region 4) in Ogden, Utah. She received a B.S. degree in wildlife ecology from the 

University of Michigan and an M.S. degree in wildlife biology from Humboldt State University. 

Acknowledgments

We thank our colleagues Linh Hoang, Lydia Allan, Erin Swiader, Gary Hanvey, and Tammy 

Fletcher, all from the National Forest System, Northern Region (Region 1) office in Missoula, 

Montana, who provided extensive input in the early development of this plan; Katie Moriarty, Pacific 

Northwest Research Station, and Jody Tucker, Pacific Southwest Region (Region 5), for further 

conceptual development and insights into establishing and running long-term monitoring programs; 

Cory Davis, Scott Tomson, and Carly Lewis, from the Southwest Crown Collaborative Network for 

insight into running a mesocarnivore monitoring program; and National Forest biologists throughout 

the Northern Region (Region 1) and Intermountain Region (Region 4) who have provided feedback 

on this protocol as well as financial and logistical support.
Contents

Chapter 1: The Need for Mesocarnivore Monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Rationale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

 Primary Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 Secondary Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 Leadership and Partnerships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

 Monitoring Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 Monitoring Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 Statistical Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 Timeline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 Funding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Document Organization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Chapter 2: Background Information on Mesocarnivore Species . . . . . . . . . . . . . . . . . . 8

Primary Species Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 Fisher . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 Regulatory and Legal Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

 Benefits of Monitoring Fishers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 Canada Lynx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

 Regulatory and Legal Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 Benefits of Monitoring Lynx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 Wolverine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 Regulatory and Legal Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 Benefits of Monitoring Wolverines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

Secondary Species Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 American Marten and Pacific Marten . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 Regulatory and Legal Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

 Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

 Benefits of Monitoring Martens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

 Montane Red Fox . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

 Regulatory and Legal Status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

 Life History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

 Benefits of Monitoring Montane Red Foxes . . . . . . . . . . . . . . . . . . . . . . . . . 18

Chapter 3: Monitoring Questions for Primary Species . . . . . . . . . . . . . . . . . . . . . . . . 19

Monitoring Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Chapter 4: Goal Efficient Monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Rationale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 Rare Species Challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 Monitoring Design Challenge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

 Goal Efficient Monitoring Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Statistical Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Recommended Survey Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Implementation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Thresholds and Triggers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Centralized Database . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Appendix A: Regulatory and Legal Status of Primary and Secondary Species . . . . . 45

Appendix B: History of ESA Listing Petitions and Decisions for Primary Species . . . 46

Appendix C: Northern Rockies Lynx Management Direction 2006: 

 Occupied and Unoccupied Lynx Habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Appendix D: Mesocarnivore Information Needs in the Northern Region (Region 1) 

 and the Intermountain Region (Region 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Appendix E: Example Multispecies Mesocarnivore Monitoring Form . . . . . . . . . . . . . 60

U.S. Forest Service RMRS GTR-388. 2018. 1

Chapter 1: The Need for Mesocarnivore Monitoring

Rationale

This chapter describes the rationale for undertaking the monitoring 

effort and provides a brief overview of the monitoring plan.

The western United States is home to a diverse mesocarnivore 

community. Mesocarnivores, small to mid-sized (typically <15 

kilograms) carnivores, are ecologically important species. Many of these 

species have undergone significant distributional changes in the United 

States since European settlement began in the 19th century (Prugh et 

al. 2009; Roemer 2009). As a result, many mesocarnivore species are 

of conservation concern, as well as the focus of public interest and 

litigation. 

Long-term monitoring of mesocarnivores is important to the USDA 

Forest Service for many reasons. For example, monitoring plays a key 

role in assessing land management plans (i.e., Forest Plans), informing 

species conservation assessments, and conforming to species-specific 

monitoring requirements and agreements. 

The Forest Service is required by the National Forest Management 

Act of 1976 (NFMA) to provide for the diversity of plant and animal 

communities on National Forest System (NFS) land, including 

mesocarnivores and the habitats that support them. The Forest Service 

ensures diversity primarily through implementation of Forest Plans 

developed under current or historical planning regulations: the 2012 

Planning Rule (FR 2012a; USFS 2012) and 1982 Planning Rule 

(FR 1982), respectively. Monitoring is essential to testing relevant 

assumptions, tracking conditions over time, and measuring the 

management effectiveness of Forest Plans. Monitoring information is 

important for evaluating whether a change in management direction 

is needed. While the Forest Service generally monitors ecological 

conditions such as vegetation structure and composition (USFS 2012, 

2015), direct monitoring of wildlife populations can also be essential 

(Schultz et al. 2013). This is especially true for some mesocarnivores, 

such as those that were identified as management indicator species under 

the historical planning rule, those that may be identified as focal species 

under the current planning rule, or those for which there is conservation 

concern but precise habitat needs are unknown. Management indicator 

species are species whose population changes are “believed to indicate 

effects of management activities on other species of selected major 

biological communities or on water quality” (CFR 2012). Focal species 

provide information about the effectiveness of the Forest Plan “in 

providing the ecological conditions necessary to maintain the diversity of 

plant and animal communities and the persistence of native species in the 

plan area” (FR 2012a). 
U.S. Forest Service RMRS GTR-388. 2018. 2

The 2012 Planning Rule establishes a two-tiered monitoring 

approach for all Forest Plans, regardless of which planning rule 

originally guided the plan. One tier is for plan-level monitoring, and 

the other is for broader-scale questions that are best answered at larger 

geographic scales than a single plan area. Mesocarnivores with large 

home ranges, low population densities, and elevated conservation status 

across large landscapes are an excellent fit for broader-scale monitoring 

strategies. 

Mesocarnivore monitoring data can also help the Forest Service 

and others accurately evaluate species conservation status. Status 

designations include regional forester’s sensitive species and species 

of conservation concern (USFS 2012); species petitioned, proposed, 

or listed by the U.S. Fish and Wildlife Service (USFWS) as threated or 

endangered under the Federal Endangered Species Act (ESA) of 1973; 

and species evaluated by States for natural heritage rankings or wildlife 

action plans. Unbiased designations are important because the Forest 

Service ultimately must address species with elevated conservation status 

in project planning or land management planning efforts. 

Species monitoring, alone or in concert with habitat monitoring, 

may also be necessary to carry out specific commitments. For example, 

Canada lynx (hereafter, lynx) have unique monitoring requirements 

associated with the Northern Rockies Lynx Management Direction 

(hereafter Lynx Management Direction) (USFS 2007) and accompanying 

Biological Opinion (USFWS 2007). The Lynx Management Direction 

amended several Forest Plans to aid the recovery of lynx populations, 

and was predicated on certain assumptions that require monitoring. For 

example, it is important to validate over time which areas are occupied 

by lynx, so that the Lynx Management Direction can effectively promote 

species recovery. 

This document will focus on mesocarnivores in the Northern Region 

(Region 1) and Intermountain Region (Region 4) as example regions 

that have supported the development of the monitoring plan and will 

provide the first implementation of this approach. We recognize that the 

mesocarnivore species of interest will vary by region, but the challenges 

associated with monitoring these mesocarnivores will be similar across 

all regions. Therefore, the approach also provides a basis for more 

general mesocarnivore monitoring. In the western United States, other 

NFS Regions where this approach would be applicable include the 

Rocky Mountain Region (Region 2), Pacific Southwest Region (Region 

5), and Pacific Northwest Region (Region 6). 

Regions 1 and 4 and the Forest Service’s Rocky Mountain Research 

Station (RMRS) have worked collaboratively to identify three primary 

mesocarnivore species for monitoring. Primary species are species 

that have been identified in broad-scale monitoring priorities for 

mesocarnivores, have relevant designations under the 1982 or 2012 

Planning Rules (Appendix A), and have been petitioned for listing under 

the ESA multiple times (Appendix B). Within this monitoring structure, 
U.S. Forest Service RMRS GTR-388. 2018. 3

primary species will be the focus of surveys specifically designed to 

detect their population states with a priori statistical power. Other species 

not listed as being “primary” will be detected opportunistically within 

sampling designs associated with the primary species. 

Primary Species

1. Fisher (Pekania pennanti)

2. Canada lynx (Lynx canadensis)

3. Wolverine (Gulo gulo)

Additionally, we identified two secondary mesocarnivore species for 

opportunistic monitoring in conjunction with primary species monitoring. 

Secondary species are species with relevant designations under the 1982 

or 2012 Planning Rules (Appendix A) or native mesocarnivores where 

major gaps in knowledge about distribution exist. 

Secondary Species

4. American marten (Martes americana) 

5. Pacific marten (Martes caurina)

6. Montane red fox (Vulpes vulpes sspp.) 

Monitoring mesocarnivores is complicated by two overarching 

problems: (1) gaps in knowledge about the basic distribution and habitat 

requirements of mesocarnivores on Forest Service and other land 

ownerships; and (2) the lack of an appropriate, multiscale framework 

that can resolve short-term and long-term trend information about 

mesocarnivores while addressing the uncertainty related to general low 

detectability and low abundance of mesocarnivores. We hope that by 

having a well-articulated plan that meets the Forest Service needs, we 

can better engage with our partners to identify how to best work together 

on mesocarnivore monitoring in both the Rocky Mountain region and 

across the western United States. 

This section outlines the details of how we in the Rocky Mountain 

region will accomplish the mesocarnivore monitoring strategy. 

Leadership and Partnerships 

The three primary species have large distributions and home range 

sizes. Patterns and trends will become clear, depending on species, 

at scales that range from hundreds to tens of thousands of square 

kilometers. We recognize that at these large scales, the most efficient 

and biologically relevant approach to mesocarnivore monitoring is a 

multiregional approach. Therefore, this strategy has been developed as a 

collaborative, multiregional effort led by RMRS in Missoula, Montana, 

the Northern Region (Region 1), and the Intermountain Region (Region 

4). We also recognize that at these scales, monitoring should involve 

many jurisdictions, including Federal, State, tribal, and large private 

landowners, which means that it is best accomplished as a cooperative 

partnership. Therefore, the ultimate goal of this monitoring strategy is to 

facilitate a larger, collaborative, multipartner, western United States-wide 

monitoring effort, with leadership shared across all partners. 

Approach
U.S. Forest Service RMRS GTR-388. 2018. 4

Monitoring Questions

To meet the Forest Service’s planning and management needs and to 

most effectively and efficiently monitor mesocarnivores, we propose the 

following multispecies, multiregional mesocarnivore monitoring strategy. 

The strategy will address: 

1. Gaps in knowledge about the distribution of mesocarnivores in 

the Rocky Mountain region;

2. Uncertainty related to low detectability, low abundance, and large 

home range sizes of mesocarnivores;

3. Simple, straightforward monitoring questions, that when repeated 

over time, will provide the Forest Service with appropriate 

information to determine both planning-scale (e.g., national 

forest) and broad-scale (e.g., region) distribution, population 

status, and trends of mesocarnivores; and 

4. Monitoring to support forest plans, regional broader-scale 

monitoring programs, and the Lynx Management Direction.

This plan uses a series of tiered monitoring questions that will 

maximize information efficiency so that, as each subsequent question is 

asked, it will answer all previous questions. The plan will address three 

questions about each of the primary species. Answers to these questions, 

when aggregated over time, will provide trend estimates that reflect 

the probability that an area remains occupied by either individuals or a 

reproducing population. These questions are:

1. Is the species present? 

2. Are multiple individuals of a single sex present?

3. Are multiple individuals, including both sexes, present? 

Not all of these questions will be answered in every location for 

every species. Rather, the plan allows for flexibility such that only 

appropriate questions for each area and species are addressed. Most past 

monitoring plans for carnivores are repeated grid-based surveys, where 

the landscape is divided into large (e.g., 5 km × 5 km) cells and selected 

cells are surveyed on a repeated basis (e.g., Kindberg et al. 2011; Proctor 

et al. 2010; Zielinski et al. 2013). Though this approach is effective, it 

is also expensive. Additionally, not all areas on the landscape have the 

same knowledge gaps or the same information needs. Thus, we need goal 

efficient monitoring (GEM), that is, the right survey design for the right 

location and knowledge level.

Monitoring Area

Broadly stated, the monitoring area is defined by the ranges, both 

current and historical, of the primary species of interest. As such, it 

incorporates areas managed and owned by many entities. Here, however, 

we will concentrate on Forest Service-administered lands in the Northern 

Region (Region 1) and Intermountain Region (Region 4), where 

implementation will begin (hereafter Monitoring Area) (fig. 1). Some 
U.S. Forest Service RMRS GTR-388. 2018. 5

Figure 1—The Monitoring Area (shown in dark green) and the Forest Service participating regions (Northern Region 

and Intermountain Region). 
U.S. Forest Service RMRS GTR-388. 2018. 6

national forests and national grasslands in Regions 1 and 4 do not contain 

habitat associated with any of the primary species and therefore are not 

included in the Monitoring Area. 

Statistical Approach

This monitoring strategy will use a multistate approach to address 

the monitoring questions for the primary species. The states each 

correspond to a monitoring question and are hierarchically structured 

so that information at one state provides information at all of the states 

below that state. This approach will be aided by the use of Bayesian 

statistics, which are revolutionizing the way managers can track animal 

populations (Betram et al. 2015; Gelman et al. 2014). Bayesian statistics 

provide multiple advantages for problems where information is scarce, 

because they can accomplish the following: (1) incorporate knowledge 

from previous efforts, (2) fill in missing data with previous efforts or 

from other species in the community, (3) combine multiple streams of 

data, (4) share information between species to inform estimates of rare 

species, and (5) quantify uncertainty from processes on the landscape as 

well as from sampling. Due to the difficulties in collecting data for these 

rare species, it is imperative that statistical analyses take full advantage 

of available information. Thus, the advantages conferred by Bayesian 

statistics are essential to a large-scale mesocarnivore monitoring strategy. 

Importantly, a Bayesian structure allows the Forest Service to 

be flexible concerning when and where the agency asks the three 

monitoring questions. These monitoring questions will change over time 

as levels of information and Forest needs change. To be both efficient 

and effective, the Forest Service must ask the right question in the 

right place and at the right time by using the most efficient and robust 

sampling design. Formally designing the monitoring within a multistate, 

Bayesian structure makes this possible. Refer to Chapter 4 for a more 

detailed description of this approach.

Timeline

Monitoring will be implemented with a pilot phase and a longterm implementation phase. The pilot phase began in 2017 and will run 

through 2020. This approach has worked well for other Forest Service 

mesocarnivore monitoring efforts. For example, the Southern Sierra 

Nevada Fisher Monitoring Program in the Pacific Southwest Region 

(Region 5) used a pilot phase to reduce uncertainty, which allowed the 

Forest Service to reduce effort and take advantage of new technology 

in the current implementation phase (ongoing) (Truex et al. 2016). The 

timeline of the implementation phase is tentative and will depend on 

a variety of factors, including funding. Figure 2 shows an overview 

of the focus of each phase of monitoring. A practitioner’s guide with 

detailed GEM implementation instructions will follow in a subsequent 

publication.
U.S. Forest Service RMRS GTR-388. 2018. 7

Figure 2—The focus of Pilot Phase and Implementation Phase of the monitoring effort.

Funding

The effort is currently funded through fiscal year 2018 by the 

Northern Region (Region 1) Resource Information Management (RIM) 

Board and the Intermountain Region (Region 4). Funding will continue 

to be sought from these sources, as well from partners and other sources.

The remainder of this document provides the details of how 

this strategy will be accomplished. Chapter 2 includes background 

information on each species, including life history information and why 

the species was selected. Chapter 3 outlines the process of developing 

the monitoring questions and determining where the Forest Service will 

ask each of the monitoring questions for each primary species. Chapter 4 

describes in detail the GEM framework, including the statistical and field 

approaches used to obtain information for monitoring questions.

Document 

Organization
U.S. Forest Service RMRS GTR-388. 2018. 8

Chapter 2: Background Information on Mesocarnivore Species

This chapter contains background information on the primary species 

(fishers, lynx, and wolverines) and secondary species (martens and 

montane red fox) considered in this plan. It includes why each species 

was selected, the specific legal and regulatory framework that dictates 

management of the species, life history information, and the benefits of 

monitoring the species.

Fisher

Fishers were selected as a primary species because of their small 

population size and because they are disjunct and genetically distinct 

from other fisher populations. In fact, the USFWS designated northern 

Rocky Mountain fishers as a distinct population segment (DPS) in 

2011. A DPS is a discrete population segment that is considered to 

be significant to the species as a whole (FR 2010b). The range of the 

northern Rocky Mountain DPS includes parts of east-central Idaho, 

southern Montana, and northwest Wyoming (FR 2011). Fishers in this 

DPS represent a distinct lineage that survived what was presumed 

to be extirpation by the 1960s (Schwartz 2007; Schwartz et al. 

2012; Vinkey et al. 2006). As a result, they have been identified as a 

management indicator species, regional forester’s sensitive species, or 

proposed species of conservation concern on multiple forests within 

the Monitoring Area. The species was not listed after the petition in 

2009 partially in response to results of a survey effort by the RMRS 

and partners, funded by the Northern Region (Region 1), to document 

fisher range in the northern Rocky Mountain region from 2004 through 

2011. Relatively little is known about their current distribution or status 

throughout the northern Rocky Mountains. 

Regulatory and Legal Status

Federal Endangered Species Act

Fishers in the northern Rocky Mountain region were petitioned both 

to be considered to be a DPS and to be listed under the ESA in 2009 (FR 

2010a). In 2011, the USFWS determined that the listing of the northern 

Rocky Mountain DPS as petitioned in 2009 was not warranted (FR 

2011) but concluded that this area did constitute a valid DPS. Fishers 

in the DPS were again petitioned for listing under the ESA in 2013 (FR 

2016a). In 2016, the USFWS determined that listing may be warranted 

due to factors B (overutilization) and E (other manmade factors) (FR 

2016a). Most recently, the USFWS found in 2017 (FR 2017) that the 

species is neither in danger of extinction now nor likely to become so in 

the foreseeable future due to climate change, development, poisoning, 

Primary Species 

Background
U.S. Forest Service RMRS GTR-388. 2018. 9

forestry, fire, trapping, or predation; thus, the species is not warranted 

for listing at this time under the ESA. Our monitoring approach will 

provide information helpful to determining whether listing is warranted. 

Appendix B outlines these petitions and decisions.

Forest Service

Fishers are considered a regional forester’s sensitive species in 

Regions 1 and 4 (USFS 2011, 2016). Fishers have been proposed as a 

species of conservation concern on the Nez Perce-Clearwater National 

Forest (USFS 2013b). 

Life History

Habitat Requirements

Fishers occupy forested landscapes that contain complex vertical 

(trees and snags) and horizontal (downed logs or debris) structure and 

dense canopies (Lofroth et al. 2010). These forests are typically found 

in low- to mid-elevation mixed-conifer or mixed-hardwood forests 

(Sauder and Rachlow 2014). These features, and thus fishers, are not 

necessarily exclusive to old-growth forests (Raley et al. 2012). At a 

home range scale, fishers in the northern Rocky Mountain region require 

large trees—the average maximum diameter at breast height [d.b.h.] in 

used stands was 107 cm—with cavities (Schwartz et al. 2013) and areas 

where vegetation heterogeneity is high (Sauder and Rachlow 2015). 

Females need cavities for reproduction and show a strong selection for 

such structures within their home range throughout the western United 

States (Paragi et al. 1996; Raley et al. 2012; Schwartz et al. 2013). 

Average female home-range size in the northern Rocky Mountains is 

approximately 25 km2

 and average male home-range size is substantially 

larger (Schwartz et al. 2006). Fishers exhibit seasonal habitat use in the 

northern Rocky Mountains. They favor old-growth subalpine fir (Abies 

lasiocarpa) and grand fir (Abies grandis) habitats in summer, but switch 

to using both young and old-growth forest types in the winter (Jones and 

Garton 1994). Fishers also avoid open and nonforested areas (Raley et al. 

2012) and in the northern Rocky Mountains avoid stands of ponderosa 

pine (Pinus ponderosa) and lodgepole pine (Pinus contorta), probably 

due to the lack of large-diameter trees or complex understory structure, 

or a combination thereof (Schwartz et al. 2013). 

Reproduction

Female fishers reproduce when they are 2 years old. Mating occurs 

in late spring. Females give birth to one to four kits the next year 

between mid-March and early April, following delayed implantation 

and a gestation period of approximately 36 days. Fishers mate again 

within approximately 10 days of giving birth. Parental care continues 

for approximately 10 months after birth. Kits are weaned at around 

10 weeks. At 4 months, juveniles become mobile and able to travel with 

their mother. At 7 months, juveniles are somewhat independent, but stay 

within their mother’s home range. Juveniles disperse at approximately 

10 months (Lofroth et al. 2010).
U.S. Forest Service RMRS GTR-388. 2018. 10

Historical Distribution

There is limited information on the historical distribution of fishers 

in the Rocky Mountains. Fishers may have historically occurred in eastcentral Idaho, southern Montana, northwest Wyoming, and northeastern 

Washington (Gibilisco 1994). By the early 1900s, populations of fisher 

in the United States had declined significantly because of unregulated 

harvest, poisoning, and habitat loss (Vinkey 2003). Fishers were 

probably extirpated from Montana by 1929 and considered extinct or 

extremely rare in Idaho by the 1950s (Vinkey 2003). In response, there 

were several reintroductions in north-central Idaho and western Montana 

in the early 1960s (Vinkey 2003; Vinkey et al. 2006). However, a novel 

mitochondrial haplotype was identified in 2006 in Montana and northcentral Idaho, suggesting that the species may not have been extirpated 

before the reintroductions, as previously thought (Schwartz 2007; 

Schwartz et al. 2012; Vinkey et al. 2006).

Current Distribution and Population Status

Currently fishers are present in northwestern Montana and central 

Idaho (Olson et al. 2014), but their exact distribution remains uncertain. 

Benefits of Monitoring Fishers

Monitoring fishers will provide the following benefits for the Forest 

Service: 

1. Improve understanding of current fisher distribution to inform 

project-level and planning-level NEPA analysis;

2. Improve understanding of long-term trends in fishers and 

identification of potential drivers of this trend;

3. Support Forest Plan monitoring programs, particularly on forests 

where fishers are considered a management indicator species, 

sensitive species, species of conservation concern, or focal species;

4. Support regional broader-scale monitoring programs; and 

5. Provide information that can inform future ESA listing decisions. 

If fishers are petitioned or listed under the ESA in the future, 

knowledge of current distribution on Forest Service lands will 

equip the Forest Service with the information necessary to comply 

with ESA. 

Canada Lynx

Lynx are one of the most high-profile mesocarnivores that occur on 

Forest Service lands in the Rocky Mountain region. They were selected 

as a primary species because of management concerns, their protection 

under the ESA, and ongoing litigation concerns related to proposed 

management actions. Not only are lynx the center of many legal 

challenges brought against the Forest Service, but because of their status 

as threatened under the ESA (since 2000) (FR 2000), they are addressed 

in almost every Forest Service planning document in the Monitoring 

Area. Lynx habitat in the Rocky Mountain region is patchily distributed 

(Squires et al. 2013). 
U.S. Forest Service RMRS GTR-388. 2018. 11

Although some of the current distribution of lynx is described 

(Squires et al. 2013), there is still no clear regionwide understanding of 

the status of lynx populations, and changes in distribution or populations 

cannot be tracked without a long-term monitoring structure. In addition, 

there are legal requirements from the Lynx Management Direction 

Biological Opinion to monitor “unoccupied habitat,” as defined by 

the 2006 amendment to the Lynx Conservation Agreement (USFS 

and USFWS 2006), on seven national forests within the Monitoring 

Area. These surveys are expensive and time consuming, and have 

not been conducted in a coordinated manner in the past to provide a 

comprehensive look at unoccupied lynx habitat. Therefore, monitoring 

lynx can provide the Forest Service with multiple benefits, most 

importantly compliance with regulatory and planning guidance including 

the Lynx Management Direction, and help inform management decisions 

both within and outside the Forest Service. 

Regulatory and Legal Status

Federal Endangered Species Act

Lynx are listed as threatened under the ESA (FR 2000). Prior to 

listing, lynx were petitioned for listing in 1994 and determined to be not 

warranted for listing (FR 1997). However, a 1997 court order remanded 

that decision and required the USFWS to reconsider the decision. The 

species remained a candidate under the ESA from May 27, 1997 (FR 

1997) until it was listed on March 24, 2000 as threatened under the 

ESA (FR 2000). In October 2017, the USFWS released a species status 

assessment for the lynx (USFWS 2017b). A subsequent review of that 

assessment in November 2017 concluded that the lynx may no longer 

warrant protection under the ESA, and should be considered for delisting 

(USFWS 2017b). However, a formal delisting proposal has not yet been 

issued. Appendix B outlines these petitions and decisions.

Forest Service

Because lynx are listed as threatened under the ESA, they do not 

have additional status within the Forest Service.

Life History

Habitat Requirements

At a landscape scale lynx require habitats that support dense 

populations of snowshoe hares (Lepus americanus) (Mowat et al. 2000). 

In the Rocky Mountain region, lynx habitat includes mature conifer 

and spruce-fir (Picea–Abies) forests, often in areas dominated by alpine 

fir, Engelmann spruce (Picea engelmannii), or lodgepole pine (Aubry 

et al. 2000; Squires et al. 2010), but such resources are often patchily 

distributed (Squires et al. 2013). Lynx select home ranges at midelevations (between 1,425 and 1,998 meters) in areas of high canopy 

cover and low topographic roughness with little open grassland (Squires 

et al. 2013). Females require mature spruce-fir forests with high cover, 

woody debris, and large-diameter trees for den sites, although studies 

have suggested that structure may play a more important role than 

forest age (Mowat et al. 2000; Squires et al. 2008). Dens are typically in 
U.S. Forest Service RMRS GTR-388. 2018. 12

drainage-like areas away from forest edges in sheltered areas. Females 

appear to show low fidelity to den sites (Squires et al. 2007). Female 

home-range size is approximately 90 km2

 and male home-range size is 

220 km2

 (Squires and Laurion 2000). Lynx generally exhibit seasonal 

shifts in vegetation preference, favoring mature, multistory forests in the 

winter—primarily spruce-fir forests—and a wider mixture of mature and 

young forests in the summer (Squires et al. 2013).

Reproduction

Female lynx typically reproduce when they are 2 years old, although 

they have been known to breed at 1 year of age when snowshoe hares 

are abundant. Lynx breed in March and April. This period is followed by 

a 60- to 70-day gestation period. Females give birth from late April to 

mid-June (Mowat et al. 2000). Maternal care continues for approximately 

12 months and kittens begin to disperse in April and May. Throughout 

their range, lynx reproduction is closely tied to snowshoe hare abundance 

(Mowat et al. 2000).

Historical Distribution

Lynx historically occurred throughout the northern Rocky Mountains 

in Idaho, Montana, Wyoming, and Colorado (McKelvey et al. 2000). In 

the 1960s and 1970s, lynx populations in the Rocky Mountain region 

followed an expansion pattern that closely tracked population eruptions 

in Canada with a 2-year time lag (McKelvey et al. 2000).

Current Distribution and Population Status

Northern portions of Montana and a small part of northern Idaho 

are thought to support the largest lynx population in the western United 

States (USFWS 2017b). Lynx distribution in the Rocky Mountains has 

declined since the early 1990s. In addition to persistent populations in 

the Clearwater Drainage and Purcell Range in western Montana (Squires 

et al. 2013), lynx were located previously in both the Garnet Range 

of Montana, and in the Greater Yellowstone Area of Montana, Idaho, 

and Wyoming, where reproduction was also documented (Squires and 

Laurion 2000). Recent surveys have failed to detect lynx in the Greater 

Yellowstone Area (USFWS 2017). Currently, there are no estimates of 

population status, although there have been efforts in the past to look 

at population growth in northern portions of the Rocky Mountains, 

including in the Clearwater Drainage (where populations were declining 

between 1999 and 2007) and the Purcell Mountains (where populations 

were increasing between 2003 and 2007) (USFWS 2017).

Benefits of Monitoring Lynx 

Monitoring lynx will provide the following benefits for the Forest 

Service:

1. Achieve compliance with Lynx Management Direction Biological 

Opinion term and condition 3 to monitor lynx unoccupied habitat 

(Appendix C), as defined by the 2006 amendment to the Lynx 

Conservation Agreement (USFS and USFWS 2006); 
U.S. Forest Service RMRS GTR-388. 2018. 13

2. Improve understanding of current lynx distribution and trend 

information that can be used to inform project-level to planninglevel NEPA analysis; and 

3. Support regional broader-scale monitoring programs that can 

inform future ESA decisions and provide the Forest Service 

with the information necessary to efficiently comply with ESA 

regulations, including recovery goals should a recovery plan be 

established. 

Wolverine

Wolverines were selected as a primary species because of small 

populations and the potential for wolverine to be listed under the ESA. 

Wolverines are considered a regional forester’s sensitive species in the 

Northern Region (Region 1) (USFS 2011). In addition, national forests 

within the Monitoring Area are incorporating wolverine monitoring 

questions into their revised forest plan monitoring requirements (e.g., 

Bridger-Teton [USFS 2016a]). Wolverines have been petitioned for 

listing under the ESA twice (1994 and 2000), and are currently proposed 

as threatened (FR 2016b). 

The range of wolverines in the United States has contracted 

significantly since the 1900s (McKelvey et al. 2014). However, the 

Rocky Mountain region is one of the few places in the contiguous United 

States where wolverines persist in a manner similar to their historical 

distribution (Aubry et al. 2007). They are an elusive species, such that 

even when present, they are difficult to detect because of low densities, 

and the harsh terrain that they inhabit, which often occurs in areas with 

difficult access (e.g., designated wilderness). As a result, relatively little 

is known about current population trends. 

Regulatory and Legal Status 

Federal Endangered Species Act 

Currently wolverines are proposed as threatened under the ESA (FR 

2016b). Wolverines were petitioned for listing under the ESA in 1994 

(FR 1995) and 2000 (FR 2003). The decision to list has been challenged 

multiple times. Appendix B outlines these petitions and decisions. 

Most recently, the February 4, 2013, proposed rule to list the species as 

threatened was reopened for public comment and closed on November 

17, 2016 (FR 2016b). A final listing decision is expected in 2018. 

Forest Service

Wolverines are considered a regional forester’s sensitive species in 

the Northern Region (Region 1) (USFS 2011). 

Life History 

Habitat Requirements

At a landscape scale, wolverines require persistent spring snow cover 

(Aubry et al. 2007; Copeland et al. 2010; Inman et al. 2013). Unlike most 

of the wolverine distribution in the western United States, wolverines in 
U.S. Forest Service RMRS GTR-388. 2018. 14

the Rocky Mountain region do not appear to be closely associated with 

alpine vegetation (Aubry et al. 2007). Thus, wolverines may be more 

closely associated with the climatic conditions that produce persistent 

spring snow cover in the Rocky Mountain region, which is essential 

for their denning (Copeland et al. 2010; Magoun and Copeland 1998). 

Wolverines den in structures that are either buried by snow (e.g., trees, 

boulders) or make dens within the snow itself (Magoun and Copeland 

1998). At a home range scale, wolverines need habitat that supports prey 

populations, including alpine and avalanche environments (Krebs et al. 

2007). Female home-range size in the Rocky Mountain region is between 

300 and 400 km2

 and male home-range size is between 797 and 

1,582 km2

 (Copeland 1996; Inman et al. 2012a). Wolverines exhibit 

seasonal shifts, often using low elevation forests in winter and midelevation forests in summer (Krebs et al. 2007). 

Reproduction

Female wolverines can reproduce when they are 2 years old, 

although average age of reproduction across their range is thought to be 

3 years old (Inman et al. 2007). Wolverines breed between May and July. 

Breeding is followed by a period of delayed implantation and a gestation 

period of approximately 30 to 40 days. Females give birth to one to two 

kits the following spring, usually between early February and midMarch (Inman et al. 2012b). Wolverines abandon their natal dens with 

weaned kits by late April or early May (Magoun and Copeland 1998) and 

move kits to secondary (maternal) den sites. Parental care continues for 

approximately 5 months (Copeland 1996). 

Historical Distribution

Wolverines were extirpated or driven to extremely low numbers in 

the lower 48 States by the 1900s (McKelvey et al. 2014). Previously, 

their range primarily included the western mountains of the United States 

and the Great Lakes region (Aubry et al. 2007). In the Rocky Mountain 

region, their range included Idaho, Montana, Wyoming, Utah, and 

Colorado. Evidence suggests that they were absent from Colorado and 

Utah by the 1920s, but persisting and increasing in the other States after 

the 1900s (Aubry et al. 2007). 

Current Distribution and Population Status

Wolverines now reoccupy much of their historical range in the 

Rocky Mountain region, including in Idaho, Montana, Wyoming, and 

Washington (McKelvey et al. 2014). More recently, wolverines were 

detected in 2014 and 2016 in Utah. In 2014 a wolverine was detected 

in the Uinta Mountains of Utah and in 2016 a wolverine was struck by 

a car in Rich County, Utah, near Bear Lake (Utah Division of Wildlife 

Resources 2016). Before those detections, the last detections in Utah 

were in 1979 (Utah Division of Wildlife Resources 2016). Genetic 

evidence indicates that the individuals that currently occupy these areas 

are descended from recent migrants from the north in Canada (McKelvey 

et al. 2014).
U.S. Forest Service RMRS GTR-388. 2018. 15

The most recent analyses of wolverine populations in the Rocky 

Mountain region suggest that populations are small and may have 

undergone recent reductions. Schwartz et al. (2009) estimated the 

effective population size, or the approximate number of breeders in 

a population, in Montana, Idaho, and Wyoming as 35 (95 percent 

confidence interval [CI]: 28–52). The majority of wolverines in the 

Rocky Mountain region persist within this area and this small effective 

population size is therefore of concern (Schwartz et al. 2009). Based 

on predictive habitat modeling, Inman et al. (2013) concluded that the 

western United States could support approximately 644 wolverines (95 

percent CI: 506–1881) and estimated that the current population, as of 

2013, was approximately 318 wolverines (95 percent CI: 249–926). 

Benefits of Monitoring Wolverines

Monitoring wolverines will provide the following benefits for the 

Forest Service:

1. Improve understanding of current wolverine distribution and trend 

information that can be used to inform project-level and planninglevel NEPA analysis; 

2. Support regional broader-scale monitoring programs; and 

3. Provide information that can inform future ESA decisions. 

American Marten and Pacific Marten

Currently, two species of martens are recognized in the United 

States, American martens (Martes americana) and the western martens 

(Martes caurina) (Dawson and Cook 2012). Historically, they have 

sometimes been treated as subspecies, but current genetic information 

indicates they are separate species (Dawson and Cook; Schwartz et al. 

2012). The taxonomic revision is recognized by the American Society 

of Mammalogists. Both species occur in the northern U.S. Rocky 

Mountains and form a narrow hybrid zone at their junction (Dawson 

and Cook 2012; Small et al. 2003). It is unknown whether the hybrids 

produce viable offspring. 

American martens were identified as a management indicator 

species on multiple national forests within the Monitoring Area, but 

with the new distinction, some might be more appropriately identified as 

Pacific marten. Martens were selected as a secondary species because of 

management concerns in some local areas and their potential to provide 

information about the community of carnivores in a Bayesian context. 

Due to the power of “borrowing strength,” which is increased precision 

and accuracy when multiple estimates use a common distribution in a 

Bayesian framework, information on marten occurrence patterns may be 

valuable for predicting abundance or occupancy of rare species (Iknayan 

et al. 2014). 

Secondary Species 

Background
U.S. Forest Service RMRS GTR-388. 2018. 16

Regulatory and Legal Status

Federal Endangered Species Act

Martens in the Rocky Mountain region are not protected under the 

ESA. However, a closely related subspecies, the Humboldt martens 

(M. caurina humboldtensis), was petitioned for listing in 2010 (FR 

2012b; called M. americana humboldtensis in the Federal Register). The 

USFWS withdrew Humboldt martens from listing consideration in 2014 

(FR 2015b), although this decision was partially overturned in 2017 

(Center for Biological Diversity et al. v. U.S. Fish and Wildlife Service 

2017). 

Forest Service

Martens are considered a management indicator species on five 

national forests in the Monitoring Area: Bitterroot (USFS 1987a), 

Clearwater (USFS 1987b), Flathead (USFS 1986b), Custer-Gallatin 

(USFS 1986a), and Salmon-Challis (USFS 1988). 

Life History 

Habitat Requirements

Martens use a wide variety of forest types across their range. They 

are a forest obligate species that require dense canopy cover, woody 

debris that provides structure, and large trees for resting and denning 

sites (Cheveau et al. 2013; Thompson et al. 2012). Martens avoid open 

areas and, as a result, can be sensitive to forest harvest practices (e.g., 

Cushman et al. [2011]; Moriarty et al. [2016]; Potvin et al. [2000]) and 

recreation (Slauson et al. 2017). Mature and old mixed-hardwood and 

conifer forests appear to be important, although not exclusively used, 

throughout their range (Thompson et al. 2012). 

Reproduction

Female martens typically reproduce when they are 2 years old. 

Martens breed in July and August. Breeding is followed by a period of 

delayed implantation and a gestation period of approximately 27 days. 

Females give birth from late March to April to one to five kits. Martens 

abandon their natal dens when kits are 7 to 13 weeks old and move kits 

to secondary (maternal) den sites. Parental care continues through late 

summer (Buskirk and Ruggiero 1994).

Historical Distribution

Martens in the United States are not as widely distributed as they 

were before European settlement (Buskirk and Ruggiero 1994). In the 

Rocky Mountain region, however, martens are generally widespread 

and distribution is presumed to be similar to their historical distribution 

(Buskirk and Ruggiero 1994). 

Current Distribution and Population Status

Martens are currently thought to be widespread throughout the 

Rocky Mountain region (see map in Schwartz et al. 2012). Although 

local areas have reported some changes in populations, including a 
U.S. Forest Service RMRS GTR-388. 2018. 17

dramatic decrease reported in the Bitterroot National Forest as of 2013 

from track surveys (USFS 2013a), populations are generally thought to 

be stable and distribution unchanged. 

Benefits of Monitoring Martens 

Monitoring martens will provide the following benefits for the Forest 

Service:

1. Improve understanding of current marten distribution and 

populations that can be used to inform forest-plan requirements 

for monitoring martens as a management indicator species;

2. Support regional broader-scale monitoring programs; and

3. Provide information that can be used to inform rare species 

modeling.

Montane Red Fox 

Montane red foxes (Vulpes vulpes sspp.) were selected as a 

secondary species because little is known about the subspecies’ overall 

current distribution. Montane red foxes are part of the Nearctic genetic 

clade, one of two major genetic clades of native North American red 

foxes (Vulpes vulpes). The Nearctic clade is descended from populations 

that were isolated in the southern refugium during the last glacial 

maximum and currently occupies montane regions of the western United 

States and portions of southeastern Canada (see figure 1 in Aubry et 

al. 2009: 2675). Montane red foxes are a group of subspecies, several 

of which are of conservation concern, that have differentiated into 

subspecies in the western mountains of the United States. The Rocky 

Mountain red fox (Vulpes vulpes macroura) is the subspecies present in 

the Northern Region (Region 1) and Intermountain Region (Region 4), 

but does not have any conservation status currently. The Sierra Nevada 

red fox (Vulpes vulpes necator), which inhabits high elevation areas of 

the Sierra Nevada (Perrine et al. 2007), is considered endangered under 

the California Endangered Species Act (California Department of Fish 

and Wildlife 2018). A related valley subspecies, the Sacramento Valley 

red fox (Vulpes vulpes patwin) (Sacks et al. 2010), has a small genetic 

effective population size, causing concern, yet little is known about 

its contemporary abundance, demographic trajectory, or habitat use 

(Sacks et al. 2017). The Cascade red fox (Vulpes vulpes cascadensis), 

which occupies the crest and eastern slopes of the Cascade Mountains 

(Aubry 1983), is also considered a Washington State candidate species 

(Washington Department of Fish and Wildlife 2013) and a Washington 

Natural Heritage critically imperiled species (Washington Department 

of Natural Resources 2017). Because several subspecies are of conservation concern and little is known about the Rocky Mountain red fox, 

a proactive approach to understanding distribution of montane red 

foxes is likely to benefit the Forest Service by providing useful baseline 

information. 
U.S. Forest Service RMRS GTR-388. 2018. 18

Regulatory and Legal Status

Federal Endangered Species Act

Montane red foxes in the Rocky Mountain region are not protected 

under the ESA. A closely related subspecies, the Sierra Nevada red fox, 

was found warranted but precluded by other priorities for listing by the 

USFWS in 2015 (FR 2015a). 

Forest Service

Montane red foxes are not specifically addressed in any Forest 

Service directive in the Monitoring Area. See Chapter 1 for a summary 

of general Forest Service requirements that pertain to wildlife and 

monitoring of wildlife. 

Life History

Habitat Requirements

Montane red foxes occur in cooler, high elevation forests of the 

mountains in the western United States (Aubry et al. 2009). Little is 

known about their specific habitat requirements. A closely related 

subspecies, the Sierra Nevada red fox, occurs at elevations above 400 

meters (Sacks et al. 2010; Statham et al. 2012). 

Reproduction

Although little is known about the reproduction of montane red 

foxes, it is likely that it is similar to that of lowland red foxes (Vulpes 

vulpes). Red foxes are monogamous and breed from late winter to early 

spring. Similar to other related subspecies, montane red foxes may use a 

variety of den types, including a sheltered den in a structure, rock piles, 

or earthen dens (Aubry 1983). 

Historical Distribution

Prior to the last glaciation (Wisconsin glaciation), montane red 

foxes were distributed in mountain regions throughout the western 

United States. At that time, the tree line was 1,000 meters lower in the 

northwestern United States (Aubry et al. 2009). 

Current Distribution and Population Status

Little is known about the current distribution and population status 

of montane red foxes, although museum specimens indicate that the 

species was present in locations throughout the Rocky Mountain region 

approximately 80 to 100 years ago (Aubry et al. 2009).

Benefits of Monitoring Montane Red Foxes

Monitoring montane red foxes will provide the following benefits for 

the Forest Service:

1. Improve understanding of current montane red fox distribution 

and populations that are currently largely unknown; and 

2. Support regional broader-scale monitoring programs and provide 

baseline information on conservation status. 
U.S. Forest Service RMRS GTR-388. 2018. 19

Chapter 3: Monitoring Questions for Primary Species

This chapter contains a description of the monitoring questions for 

the primary species. It describes how those questions were developed and 

how they correspond to different states of a rare species population. 

We worked extensively with multiple parties across the Forest 

Service and potential external partners to gain a broad understanding of 

the information needs for the primary species. This process was started 

with a series of semistructured interviews conducted between June 

and August 2016 with Forest Service and potential external partners, 

including additional Federal agencies, State agencies, tribal governments, 

and nonprofit organizations. We ultimately focused this plan on meeting 

Forest Service information needs, although the information from this 

strategy is broadly usable to track the status of the primary species in the 

Rocky Mountain region. 

These efforts identified three important monitoring questions for the 

primary species: 

1. Is the species present? 

2. Are multiple individuals of a single sex present?

3. Are multiple individuals, including both sexes, present? 

The questions are tiered, such that answering any higher-tier question 

will provide an answer for that question and any questions below. The 

tiers are defined based on the amount of information they provide about 

the population. Thus, the first-tier question, “Is a species present?” 

provides the least amount of population-level information. Each of these 

questions corresponds to a different state of a population (table 1, fig. 3). 

Table 1—Possible population states for the primary species. Wolverine is used as an example species to illustrate 

the states.

Monitoring 

Questions
U.S. Forest Service RMRS GTR-388. 2018. 20

Figure 3—Monitoring questions, with tier indicated, and the population state 

for which the questions provide answers. Wolverines are used as an example 

species to illustrate the states.

To define which tier of question was appropriate for each primary 

species on each of the national forests, we conducted in-person meetings 

with the national forests in Regions 1 and 4 between July and October 

2017 and sent follow-up questionnaires to document the results of these 

meetings. Appendix D contains these questionnaires from the national 

forests that responded. We used national forests as an initial primary 

mapping unit because the laws and policies that guide the Forest Service 

are based on the national forest scale or larger. In addition, we reviewed 

past survey information for the primary species on Forest Service lands 

within the Monitoring Area as documented in

• Forest Service Forest Plan Monitoring Reports,

• Forest Service Survey Reports,

• Conference monitoring publication,

• Existing databases within the Forest Service and publicly 

available data sources, and

• Personal communication with Forest Service employees. 

Figures 4 through 6 show the highest-tier monitoring question for 

each of the primary species on a national forest scale. Figure 7 is a 

summary of the total number of questions for each primary species. 
U.S. Forest Service RMRS GTR-388. 2018. 21

Figure 4—Phase 1 monitoring questions for fishers in the Monitoring Area.
U.S. Forest Service RMRS GTR-388. 2018. 22

Figure 5—Phase 1 monitoring questions for lynx in the Monitoring Area.
U.S. Forest Service RMRS GTR-388. 2018. 23

Figure 6—Phase 1 monitoring questions for wolverines in the Monitoring Area.
U.S. Forest Service RMRS GTR-388. 2018. 24

R1 and R4 Monitoring Questions

3 questions - all primary species

2 questions - 2 primary species

1 question - 1 primary species

Jessie Golding 12/20/2017

0 50 100 150 200 25

Kilometers

Figure 7—Summary of the number of Phase 1 monitoring questions for primary species on a forest.
U.S. Forest Service RMRS GTR-388. 2018. 25

Chapter 4: Goal Efficient Monitoring 

This chapter describes the monitoring strategy: goal efficient 

monitoring. It includes the rationale for the approach, the statistical 

approach, recommended survey methods to collect the data necessary for 

each question, and an outline of how the monitoring information will be 

used. 

There are two main challenges to monitoring rare species. The first is 

that rare organisms are difficult to locate and surveys therefore tend to be 

expensive to implement. The second is, given the difficulties associated 

with rarity, a monitoring plan that is targeted to meet the needs of 

decisionmakers often does not scale appropriately to provide biologically 

relevant information about the species. Because of these challenges, 

effective long-term monitoring strategies for rare species are scarce. 

However, monitoring rare species is important because the status and 

trend of rare species affect legal, management, and societal decisions. In 

addition, there are legal mandates, such as the monitoring required for 

lynx under the Lynx Management Direction Biological Opinion (USFWS 

2007), to monitor rare species. Finally, monitoring may be the only 

source of information for rare species. Thus, we need an approach that 

robustly addresses the monitoring questions identified in Chapter 3 and 

that overcomes the existing challenges of rare species monitoring. 

Rare Species Challenges

Rare species are often difficult to detect, either because of low 

densities, secretive behavior, or lack of knowledge about when and 

where to find an organism. As a result, land and wildlife managers who 

need information about rare species are often faced with a tradeoff 

between selecting “omnibus” survey methods (Sauer et al. 2003), defined 

as methods capable of detecting multiple species, or targeted survey 

methods to get specific information on a species of interest. Information 

constraints typically limit the utility of omnibus methods for rare species 

(Manley et al. 2004). Because omnibus sampling approaches are, by 

definition, not specifically designed to collect sufficient data for any 

particular species, rare species occurrences may simply be too scarce to 

provide meaningful monitoring data. 

Recent statistical advances such as Bayesian multispecies models 

allow improved analyses of rare species data when collected during 

multispecies surveys (Iknayan et al. 2014). These advances in analytical 

capability can partially alleviate this problem, but the problem remains. 

There are, however, undeniable cost efficiencies associated with omnibus 

approaches; if one cares about monitoring more than one species, using 

omnibus approaches is generally more efficient than conducting multiple 

Rationale
U.S. Forest Service RMRS GTR-388. 2018. 26

independent surveys each targeted at an individual species (Manley et 

al. 2004). Further, individual species surveys cannot take advantage of 

efficiencies afforded by the use of Bayesian multispecies models; large 

sample sizes are needed to detect trends in rare species using traditional 

frequentist statistical analyses (e.g., Ellis et al. 2014). Thus, the ideal rare 

species monitoring plans must contain elements of both omnibus and 

targeted surveys. 

The key to efficiency in designing rare species surveys is to design 

the most cost-effective mixture of omnibus and targeted sampling 

approaches, such that all of the targeted species are adequately sampled. 

For rare carnivores, existing survey methods, including tracking to obtain 

genetic samples, remote cameras, or noninvasive hair snare methods, 

represent a mixture of omnibus and targeted efficiencies. These methods 

are inherently multispecies and therefore omnibus. However, they are 

also somewhat targeted in that they are better at detecting certain species. 

For example, small baited enclosures (cubbies) are effective for sampling 

mustelids, but other animals use these traps as well (Kendall and 

McKelvey 2008).

Monitoring Design Challenge

Most large-scale, long-term monitoring programs for wildlife are 

“surveillance” monitoring programs, a term defined by Nichols and 

Williams (2006), suggesting a generalized design. For instance, the 

Breeding Bird Survey (BBS) is a broad-scale survey that has been 

conducted from 1966 to the present across the United States to evaluate 

patterns of over 400 North American bird species (Sauer et al. 2017); 

data for all species are gleaned from a single omnibus sampling design. 

Surveillance monitoring often contains sophisticated statistical design 

elements and will provide reliable data for broad-scale trends, but there 

is little effort to ensure the scales are pertinent to specific management 

areas (e.g., watersheds) or that rare species are adequately sampled 

(Manley et al. 2004). 

Because surveillance monitoring is not targeted at answering specific 

questions for specific species, inferences tend to be weak and relative, 

and as such there is strong incentive to retain design and methods 

across time to produce more reliable indices. Consequently, surveillance 

monitoring designs are not responsive to current needs; design 

changes tend to be limited to localized increases in sampling density 

(densification) within a broader invariant design. Nichols and Williams 

(2006) noted that this type of surveillance monitoring efforts fails to 

address pertinent management questions and it is not designed to do so. 

They proposed instead that monitoring should emulate scientific studies 

and that a “targeted” approach to monitoring may be more appropriate 

(see also Holthausen et al. 2005). In a scientific study, a question is 

clearly posed with a matching sampling design to generate information 

that can be tested, with high confidence, and produce direct answers with 

adequate reliability. 
U.S. Forest Service RMRS GTR-388. 2018. 27

Nichols and Williams’ (2006) approach solves the problem of 

data production at suboptimal scales and with inadequate power by 

surveillance monitoring, but creates its own problems. Targeted questions 

are continuously fluctuating. Societal needs and priorities are constantly 

changing, and the acquisition of knowledge creates new questions. As an 

example that is simple but pertinent to rare carnivores, one often enters 

a particular land area in a state of ignorance: Whether the organism 

is present, or not, is unknown. So the first targeted question is: Does 

this area contain at least one organism? A monitoring approach can be 

designed to answer this question. 

Assuming that an organism has been found, the next series of 

questions is likely to revolve around its population status: Were there 

more than one, were there both females and males, and were the females 

reproductive? The necessary targeted information, and therefore the 

optimal targeted design, is different if the question is one of occurrence 

versus detecting individual males and females. Note also that though 

represented here as a linear chain of increasing information, the 

progression could easily vary: Reproductive females may be found in one 

survey but fail to be detected in subsequent surveys. Due to the vagaries 

of small population dynamics, populations may shrink or vanish. 

When we look across broader spatial extents, it is clear that both 

the local needs and information levels will be in a constant state of 

flux. Thus, there exists a conundrum for any broad-scale monitoring 

effort seeking to maintain continuity across space and time. Efficiency 

demands that the targeted goals and therefore efficient designs change 

across space and time. This contrasts with integrative needs of a broader 

survey, its continuity, and the ability to use and compare data collected in 

one place and time to inform data collected in different places and during 

either previous or subsequent times. In short, although Nichols and 

Williams (2006) lay out a compelling argument concerning why targeted 

monitoring is advantageous in a particular place and time, they do not 

offer a coherent approach to integrate these disparate efforts into a longterm, broad-scale monitoring effort. 

Goal Efficient Monitoring Approach

To address these challenges and provide the relevant management 

information from the monitoring questions presented in Chapter 3, 

we propose a new approach using a flexible framework we call goal 

efficient monitoring (GEM). GEM is a long-term monitoring framework 

that allows local flexibility and the ability to efficiently use data from 

seemingly disparate targeted local efforts that aggregate into a coherent 

whole. It is based on the ideas of focusing on states of a rare population 

rather than abundance, as was suggested by MacKenzie et al. (2005). 

It has four main components that make it uniquely able to solve rare 

species monitoring problems: 

1. Tiered monitoring questions that address the finest-scale local 

needs but can scale up appropriately because of their tiered 

nature;
U.S. Forest Service RMRS GTR-388. 2018. 28

2. Monitoring questions that correspond to well-defined and 

discernable population states of interest;

3. A well-developed understanding of the processes of transitioning 

between or staying within a population state; and 

4. Defined maximum scale at which each state is relevant.

In the context of multispecies mesocarnivore monitoring, this 

approach has the additional benefit of including sufficient elements of 

omnibus sampling to take advantage of efficiencies afforded through 

Bayesian multispecies surveys (Iknayan et al. 2014). 

The tiered monitoring questions for this plan were developed to 

meet local information needs but can provide relevant information, even 

if questions differ, across a larger area. For example, forest resource 

managers may want to know whether a rare species population on their 

forest has the potential to be a reproducing population, so they would ask 

the third-tier question: Are multiple individuals of a single sex present or 

are both sexes present? If the managers on the neighboring forest have 

only a question about whether the species is present, both efforts can be 

combined to determine species presence across the region because the 

third-tier question always answers the second- and first-tier questions. 

For the population states presented (table 1), we assumed, for the 

sake of clarity, that transitions follow a linear progression and that a 

population must transition through each state in a predictable manner. 

This is not strictly the case, and allowing transitions that jump states 

(e.g., both sexes present transitioning to not present) is acceptable within 

the modeling framework. The vast majority of transitions, however, will 

probably be those shown in figure 8: either remaining within a state or 

transitioning into an adjacent state. 

By tracking state transitions, answers to the tiered GEM questions, 

when aggregated over time, will provide monitoring trend estimates. For 

example, if upward transitions exceed downward transitions regionally, 

Figure 8—The four potential states (represented as circles) and the process of transitioning between the states 

(straight arrows) or staying within a state (curved arrows). Lynx are used as an example species to illustrate the 

states.
U.S. Forest Service RMRS GTR-388. 2018. 29

this will indicate an expanding population. One of the most powerful 

parts of GEM is that monitoring states over time gives us context to 

understand the biological importance of the current state. In fact, we find 

that the biological importance of the population state is best understood 

with knowledge of the previous transition or maintenance of a state 

(table 2). 

Within the GEM context, information on transitions or remaining 

within a state comes from repeatedly asking the tiered questions 

over time. Because the questions asked change with knowledge of 

the population state, surveys are conducted in a manner that always 

addresses the current information need. Thus, one has to constantly 

cycle through the questions, allowing for repeats of cycles as well as 

progression through the monitoring questions. Appendix E provides a 

hypothetical example of what a progression through 4 years of GEM 

would look like for the three primary species on the Bitterroot National 

Forest in the Northern Region (Region 1). 

The maximum scale at which the states for this plan (table 1) are 

relevant is based on the utility of knowing that a population exists in the 

highest state. That is, once the population becomes large enough that 

knowing there are multiple individuals of both sexes is trivial, the statebased approach ceases to be useful. At these higher population levels, 

meaningful information on population change takes the form of changes 

in numbers or in demographic parameters. In this effort we have chosen 

to delineate areas where population state will be estimated to areas no 

Table 2—Biological importance (inference that can be drawn from observation) of states based on observation and 

previous transition. Most likely importance in bold (if multiple possibilities shown). Wolverine is used as an example 

species to illustrate the states. 
U.S. Forest Service RMRS GTR-388. 2018. 30

larger than the equivalent of 50 home ranges of the species of interest. In 

most cases the actual population that inhabits these spaces will be much 

smaller than this theoretical carrying capacity.

We propose a multistate model to analyze the data in the GEM 

framework. The GEM analysis builds on the multistate dynamic 

occupancy models of Royle (2004), Royle and Link (2005), Nichols et 

al. (2007), MacKenzie et al. (2009), and Kéry and Schaub (2012).

 Figure 9 shows a schematic of the states and the associated statistical 

transition probabilities. Table 3a summarizes those probabilities and 

table 3b shows the probabilities associated with observing each state. 

The GEM multistate dynamic occupancy model will be most useful 

in a Bayesian hierarchical framework because these models can estimate 

transition or staying probabilities, even those with missing data, while 

explicitly dealing with observational uncertainty and imperfect detection 

(Dorazio 2016). Bayesian frameworks also allow for the incorporation 

of prior information (Dorazio 2016), meaning that surveys that were 

accomplished in the past can be used to inform estimates and design of 

surveys in the future. With the repeated asking of questions over time, 

Bayesian analysis provides a considerable advantage over traditional 

estimation methods. The previous survey efforts inform not only the 

next monitoring question, but also the information that the model has 

available to use for predictions. In fact, the estimation of all three of 

these questions can be accomplished in the single GEM model. Finally, 

Bayesian multispecies models may allow new insight into rare species 

by using information from the community of species that are detected by 

using omnibus survey methods (Iknayan et al. 2014). These advantages 

are essential for a large-scale, rare mesocarnivore monitoring program.

Statistical Approach

Figure 9—The four potential states (represented as circles) and the statistical transition probabilities (Ψ) between 

the states. Curved arrows represent the probability of not transitioning (1-Ψ) and the straight arrows represent 

the probability of transitioning (Ψ). Subscripts are as follows: np = not present; si = single individual; mi = multiple 

individuals (single sex); mb = multiple individuals (both sexes). Combined subscripts indicate a transition in the order 

that the subscripts appear (e.g., npsi = not present transitioning to single individual). 
U.S. Forest Service RMRS GTR-388. 2018. 31

In the GEM framework three types of information are required: 

detection/nondetection information, individual identification, and sex 

determination. Genetic sampling by noninvasive survey methods can 

provide all three of those pieces of information. In addition, identification 

reliability is high with genetic methods, which is particularly important 

for rare species. Therefore, for this strategy we recommend the use of 

three noninvasive survey methods to collect genetic samples: snow 

tracking (Squires et al. 2004), multispecies bait station, and targeted bait 

station (fisher – Schwartz et al. 2006; lynx – McKelvey et al. 1999).

Snow tracking should be used as the primary lynx detection method 

as it has proven to be the most reliable for winter surveys (Squires et al. 

2012). Snow tracking should be used in combination with multispecies 

bait station setups, as multispecies bait stations are more efficient for 

detecting a broad suite of species. Both snow track surveys (McKelvey 

et al. 2006) and bait stations allow the collection of genetic samples, 

Recommended 

Survey Methods

Table 3a—The transition probabilities (ψ) between the states. Subscripts are as follows: np = not present; si = single 

individual; mi = multiple individuals (single sex); mb = multiple individuals (both sexes). Combined subscripts indicate 

a transition in the order that the subscripts appear (e.g., npsi = not present transitioning to single individual). 

Table 3b—The probability of detecting (p) the states. Subscripts are as follows: np = not present; si = single 

individual; mi = multiple individuals; mb = multiple individuals (both sexes).
U.S. Forest Service RMRS GTR-388. 2018. 32

which enable verification of species and individual identification. We 

recommend the use of remote cameras at a portion of the multispecies 

and targeted bait stations to (1) provide multispecies omnibus survey 

information; (2) understand the interaction between animals and the bait 

station setup; and (3) obtain photos, videos, or both for promotional or 

educational materials. Table 4 provides an overview of how each survey 

method can be used to detect each state and what detection is needed to 

confirm presence of the state. 

National forests will have the ability to decide to participate in the 

survey effort. Ideally, national forests will work with the monitoring 

coordinator to design and implement multispecies surveys for the 

primary species. The monitoring coordinator will provide survey design 

assistance, including maps and survey locations. The coordinator 

will work closely with the national forests to ensure that the surveys 

maximize the potential to meet the needs of the forest and provide useful 

information in this framework. The monitoring coordinator will play an 

essential role in design, data storage, consistency, communication, and 

collaboration, all of which will help ensure the success of the effort.

Implementation

Table 4—Detection method for each state. Lynx is used as an example species to illustrate the states. 
U.S. Forest Service RMRS GTR-388. 2018. 33

Monitoring is most effective when data generated from the efforts are 

tied to thresholds or triggers (Nie and Schultz 2012; Schultz et al. 2013). 

Thresholds are predetermined signals that have been identified due to 

biological or management relevance. Triggers are predetermined signals 

that, when detected, require an action (Block et al. 2001). We encourage 

the development of thresholds and triggers as this monitoring plan is 

implemented. Similarly, triggers may be set to require committing to 

additional monitoring should populations of the target species diminish 

in occupancy by a certain percentage. Overall thresholds and triggers can 

help efficiently direct monitoring efforts by explicitly linking monitoring 

to the biology of the species or management (Schwartz et al. 2015). 

The approach described in this chapter can be accomplished only 

through centralized data storage. If the information collected to answer 

these monitoring questions is not stored in a central database, the 

information gained through repeating the questions over time could 

be lost. In a Bayesian system the power to identify current conditions 

depends on prior data collection, so access to these data represents 

an integral part of the statistical design. The central database will be 

maintained through RMRS and at a minimum will include monitoring 

location information, date, species detection information (including the 

method used to identify the species), and individual identification or 

genetic information (if applicable). Monitoring information will also 

be compatible with and stored in the Forest Service corporate wildlife 

database, NRM Wildlife, so that it is accessible to NFS employees. 

We will work with partners to ensure data formats are compatible 

with existing local databases, but data sharing will not occur without 

additional arrangements. This is especially important for rare species 

where location data may be sensitive.

Thresholds 

and Triggers

Centralized 

Database
U.S. Forest Service RMRS GTR-388. 2018. 34

References

Aubry, K.B. 1983. The Cascade red fox: Distribution, morphology, 

zoogeography, and ecology. Dissertation. Seattle, WA: University of 

Washington. 

Aubry, K.B.; Koehler, G.M.; Squires, J.R. 2000. Ecology of Canada lynx 

in southern boreal forests. In: Ruggiero, L.F.; Aubry, K.B.; Buskirk, 

S.W.; [et al.], eds. Ecology and conservation of lynx in the United 

States. Boulder, CO: University Press of Colorado; Fort Collins, CO: 

U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station.

Aubry, K.B.; McKelvey, K.S.; Copelands, J.P. 2007. Distribution and 

broad-scale habitat relations of the wolverine in the contiguous United 

States. Journal of Wildlife Management. 71: 2147–2158.

Aubry, K.B.; Statham, B.N.; Perrine, J.D.; [et al.]. 2009. Phylogeography 

of the North American red fox: Vicariance in Pleistocene forest 

refugia. Molecular Ecology. 18: 2668–2686.

Block, W.M.; Franklin, A.B.; Ward, J.P., Jr.; [et al.]. 2001. Design and 

implementation of monitoring studies to evaluate the success of 

ecological restoration on wildlife. Restoration Ecology. 9: 293–303.

Buskirk, S.W.; Ruggiero, L.F. 1994. Chapter 2: American marten. In: 

Ruggiero, L.F.; Aubry, K.B.; Buskirk, S.W.; Lyon, L.J.; Zielinski, 

W.J.; [et al.], eds. The scientific basis for conserving forest carnivores 

in the western United States: American marten, fisher, lynx, 

and wolverine. Gen.Tech. Rep. RM-254. Fort Collins, CO: U.S. 

Department of Agriculture, Forest Service, Rocky Mountain Forest 

and Range Experiment Station: 7–37. 

California Department of Fish and Wildlife. 2018. Threatened and 

endangered species. [Database]. Sacramento, CA: California 

Department of Fish and Wildlife, Nongame Program. http://www.dfg.

ca.gov/wildlife/nongame/t_e_spp/. [Accessed May 31, 2018].

Cheveau, M.; Imbeau, L.; Drapeau, P.; Belanger, L. 2013. Marten space 

use and habitat selection in managed coniferous boreal forests of 

eastern Canada. Journal of Wildlife Management. 77: 749–760.

Code of Federal Regulations (CFR). 2012. National Forest System Land 

Management Planning, 36 CFR. Section 219 Subpart A.
U.S. Forest Service RMRS GTR-388. 2018. 35

Copeland, J.P. 1996. Biology of the wolverine in central Idaho. Thesis. 

Moscow, ID: University of Idaho. 

Copeland, J.P.; McKelvey, K.S.; Aubry, K.B.; [et al.]. 2010. The 

bioclimatic envelope of the wolverine (Gulo gulo): Do climatic 

constraints limit its geographic distribution? Canadian Journal of 

Zoology. 88: 233–246.

Cushman, S.A.; Raphael, M.G.; Ruggiero, L.F.; [et al.]. 2011. Limiting 

factors and landscape connectivity: The American marten in the 

Rocky Mountains. Landscape Ecology. 26:1137–1149.

Dawson, N.G.; Cook, J.A. 2012. Behind the genes: Diversification of 

North American martens (Martes americana and M. caurina). In: 

Aubry, K.B.; Zielinski, W.; Raphael, M.G.; [et al.], eds. Biology and 

conservation of martens, sables, and fishers: A new synthesis. Ithaca, 

NY: Cornell University Press: 23–38.

Dorazio, R.M. 2016. Bayesian data analysis in population ecology: 

Motivations, methods, and benefits. Population Ecology. 58: 31–44.

Ellis, M.M.; Ivan, J.S.; Schwartz, M.K. 2014. Spatially explicit power 

analyses for occupancy-based monitoring of wolverine in the U.S. 

Rocky Mountains. Conservation Biology. 28: 52–62.

Endangered Species Act of 1973 [ESA]; 16 U.S.C. 1531-1536, 1538-

1540. https://www.gpo.gov/fdsys/pkg/USCODE-2012-title16/html/

USCODE-2012-title16-chap35-sec1531.htm. [Accessed May 24, 

2018].

Federal Register (FR). 1982. Final rule: National Forest System land and 

resource management planning. Washington, DC: U.S. Department 

of Agriculture, Forest Service. 47(190): 43026–43052. https://cdn.

loc.gov/service/ll/fedreg/fr047/fr047190/fr047190.pdf. [Accessed 

December 22, 2017].

Federal Register (FR). 1995. Notice of 90-day petition finding; 

Endangered and Threatened wildlife and plants; 90-day Finding for 

a Petition to list as Endangered or Threatened the contiguous United 

States population of the North American wolverine. 60(75): 19567–

19568. https://www.gpo.gov/fdsys/pkg/FR-1995-04-19/pdf/95-9642.

pdf#page=1. [Accessed May 3, 2018].

Federal Register (FR). 1997. Notice of 12-month petition finding; 

Endangered and Threatened wildlife and plants; 12-month Finding 

for a Petition to list contiguous U.S. Distinct Population Segment of 

the Canada lynx. Washington, DC: U.S. Department of the Interior, 

Fish and Wildlife Service. 62(101): 28653–28657. https://www.gpo.

gov/fdsys/pkg/FR-1997-05-27/pdf/97-13808.pdf#page=1. [Accessed 

December 22, 2017].
U.S. Forest Service RMRS GTR-388. 2018. 36

Federal Register (FR). 2000. Final rule; Endangered and Threatened 

wildlife and plants; Determination of Threatened status for the 

contiguous U.S. Distinct Population Segment of the Canada lynx and 

related rule. Washington, DC: U.S. Department of the Interior, Fish 

and Wildlife Service. 65(58): 16052–16086. https://www.gpo.gov/

fdsys/pkg/FR-2000-03-24/pdf/00-7145.pdf#page=3. [Accessed 

May 3, 2018].

Federal Register (FR). 2003. Endangered and Threatened wildlife 

and plants; 90-day Finding for a Petition to list as Endangered or 

Threatened wolverine in the contiguous United States. 68(203): 

60112-60115. https://www.gpo.gov/fdsys/pkg/FR-2003-10-21/pdf/03-

26453.pdf#page=1. [Accessed May 3, 2018].

Federal Register (FR). 2010a. Notice of petition finding and initiation 

of status review; Endangered and Threatened wildlife and plants; 

90-day Finding on a Petition to list a Distinct Population Segment 

of the fisher in its United States Northern Rocky Mountain Range as 

Endangered or Threatened with critical habitat. Washington, DC: U.S. 

Department of the Interior, Fish and Wildlife Service. 75(73): 19925–

19935. https://www.gpo.gov/fdsys/pkg/FR-2010-04-16/pdf/2010-

8795.pdf#page=1. [Accessed December 22, 2017].

Federal Register (FR). 2010b. Notice of review; Endangered and 

Threatened Wildlife and Plants; Review of native species that are 

candidates for listing as Endangered or Threatened; Annual Notice of 

Findings on resubmitted petitions; Annual description of progress on 

listing actions. Washington, DC: U.S. Department of the Interior, Fish 

and Wildlife Service. 75(217): 69222–69294. https://www.fws.gov/

endangered/esa-library/pdf/CNOR%2011-10-2010.pdf. [Accessed 

December 22, 2017].

Federal Register (FR). 2011. Notice of 12-month petition finding; 

Endangered and Threatened wildlife and plants; 12-Month finding 

on a petition to list a Distinct Population Segment of the fisher in its 

United States Northern Rocky Mountain Range as Endangered or 

Threatened with critical habitat. Washington, DC: U.S. Department 

of the Interior, Fish and Wildlife Service. 76(126): 38504–38532. 

https://www.gpo.gov/fdsys/pkg/FR-2011-06-30/pdf/2011-16349.pdf. 

[Accessed December 22, 2017].

Federal Register (FR). 2012a. Final rule and record of decision; 

National Forest System Land Management Planning. Washington, 

DC: U.S. Department of Agriculture, Forest Service. 77(68): 

21162–21276. https://www.fs.usda.gov/Internet/FSE_DOCUMENTS/

stelprdb5362536.pdf. [Accessed December 22, 2017].
U.S. Forest Service RMRS GTR-388. 2018. 37

Federal Register (FR). 2012b. Notice of petition finding and initiation of 

status review. Endangered and Threatened wildlife and plants; 90-day 

Finding on a petition to list the Humboldt marten as Endangered or 

Threatened. 77(8): 1900–1908. https://www.gpo.gov/fdsys/pkg/FR2012-01-12/pdf/2012-479.pdf. [Accessed May 3, 2018].

Federal Register (FR). 2015a. Endangered and Threatened wildlife and 

plants; 12-Month Finding on a petition to list Sierra Nevada red fox 

as an Endangered or Threatened species. 80(195): 60990-61028. 

https://www.gpo.gov/fdsys/pkg/FR-2015-10-08/pdf/2015-25289.pdf. 

[Accessed May 3, 2018].

Federal Register (FR). 2015b. Notice of 12-month petition finding. 

Endangered and Threatened wildlife and plants; 12-month Finding on 

a petition to list Humboldt marten as an Endangered or Threatened 

species. 80(66): 18742-18772. https://www.gpo.gov/fdsys/pkg/FR2015-04-07/pdf/2015-07766.pdf. [Accessed May 3, 2018].

Federal Register (FR). 2016a. Petition findings and initiation of status 

review; Endangered and Threatened wildlife and plants; 90-day 

Findings on 17 petitions. Washington, DC: U.S. Department of the 

Interior, Fish and Wildlife Service. 81(7): 1373–1374. https://www.

gpo.gov/fdsys/pkg/FR-2016-01-12/pdf/2016-00157.pdf.

Federal Register (FR). 2016b. Proposed rule; reopening of comment 

period; Endangered and Threatened wildlife and plants; Proposed 

rule for the North American wolverine. 81(201): 71670–71671. 

https://www.gpo.gov/fdsys/pkg/FR-2016-10-18/pdf/2016-24929.pdf. 

[Accessed May 3, 2018].

Federal Register (FR). 2017. Notice of 12-month petition findings; 

Endangered and Threatened wildlife and plants; 12-month Findings 

on petitions to list 25 species as Endangered or Threatened species. 

Washington, DC: U.S. Department of the Interior, Fish and Wildlife 

Service. 82(192): 46618–46645. https://www.gpo.gov/fdsys/pkg/FR2017-10-05/pdf/2017-21352.pdf. [Accessed December 22, 2017].

Fish and Wildlife Service (USFWS). 2007. Biological opinion on the 

effects of the Northern Rockies lynx amendment on the Distinct 

Population Segment (DPS) of Canada lynx (Lynx canadensis) in 

the contiguous United States. Helena, MT: U.S. Department of the 

Interior, U.S. Fish and Wildlife Service, Montana Field Office. 85 pp.

Fish and Wildlife Service (USFWS) 2017a. Canada lynx (Lynx 

canadensis): 5-year review, summary and evaluation. Washington, 

DC: U.S. Department of the Interior, U.S. Fish and Wildlife Service.

Fish and Wildlife Service (USFWS) 2017b. Species status assessment for 

the Canada lynx (Lynx canadensis) contiguous United States Distinct 

Population Segment. Version 1.0, October, 2017. Lakewood, CO: U.S. 

Department of the Interior, U.S. Fish and Wildlife Service. 2 p.
U.S. Forest Service RMRS GTR-388. 2018. 38

Gelman, A.; Carlin, J.B.; Stern, H.S.; [et al.]. 2014. Bayesian data 

analysis. Vol. 2. Boca Raton, FL: CRC Press.

Gibilisco, C.J. 1994. Distributional dynamics of modern Martes in North 

America. In: Buskirk, S.W.; Harestad, A.S.; Raphael, M.G.; [et al.], 

eds. Martens, sables and fishers: Biology and conservation. Ithaca, 

NY: Cornell University Press: 59–71.

Holthausen, R.; Czaplewski, R.L.; DeLorenzo, D.; [et al.]. 2005. 

Strategies for monitoring terrestrial animals and habitats. Gen. 

Tech. Rep. RMRS-GTR-161. Fort Collins, CO: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 

Iknayan, K.J.; Tingley, M.W.; Furnas, B.J.; [et al.]. 2014. Detecting 

diversity: Emerging methods to estimate species diversity. Trends in 

Ecology and Evolution. 29: 97–106.

Inman, R.M.; Brock, B.L.; Inman, K.H.; [et al.]. 2013. Developing 

priorities for metapopulation conservation at the landscape scale: 

Wolverines in the western United States. Biological Conservation. 

166: 276–286.

Inman, R.M.; Inman, K.H.; Packila, M.L.; [et al.]. 2007. Chapter 

4: Wolverine reproductive rates and maternal habitat in Greater 

Yellowstone. In: Greater Yellowstone wolverine study, cumulative 

report, May 2007. Ennis, MT: Wildlife Conservation Society, North 

America Program: 65–84.

Inman, R.M.; Magoun, A.J.; Persson, J.; [et al.]. 2012b. The wolverine’s 

niche: Linking reproductive chronology, caching, competition, and 

climate. Journal of Mammalogy. 93: 634–644.

Inman, R.M.; Packila, M.L.; Inman, K.H.; McCue, A.J.; White, 

G.C.; Persson, J.; Aber, B.C.; Orme, M.L.; Alt, K.L.; Cain, S.L.; 

Frederick, J.A.; Oakleaf, B.J.; Sartorius, S.S. 2012a. Spatial ecology 

of wolverines at the southern periphery of distribution. Journal of 

Wildlife Management. 76: 778–792.

Jones, J.L.; Garton, E.O. 1994. Selection of successional stages by fisher 

in northcentral Idaho. In: Buskirk, S.W.; Harestad, A.; Raphael, M., 

eds. Martens, sables, and fishers: Biology and conservation. Ithaca, 

NY: Cornell University Press: 377–387. 

Kendall, K.C.; McKelvey, K.S. 2008. Chapter 6: Hair collection. In: 

Long, R.A.; MacKay, P.; Ray, J.C.; Zielinski, W.J., eds. Noninvasive 

survey methods for North American carnivores. Washington, DC: 

Island Press: 141–182.

Kéry, M.; Schaub, M. 2012. Bayesian population analysis using winbugs: 

A hierarchical perspective. Waltham, MA: Academic Press. 
U.S. Forest Service RMRS GTR-388. 2018. 39

Kindberg, J.; Swenson, J.E.; Ericsson, G.; [et al.]. 2011. Estimating 

population size and trends of the Swedish brown bear Ursus arctos

population. Wildlife Biology. 17(2): 114–123.

Krebs, J.; Lofroth, E.C.; Parfitt, I. 2007. Multiscale habitat use by 

wolverines in British Columbia, Canada. Journal of Wildlife 

Management. 71: 2180–2192.

Lofroth, E.C.; Raley, C.M.; Higley, J.M.; [et al.]. 2010. Conservation 

of fisher (Martes pennanti) in south-central British Columbia, 

western Washington, western Oregon, and California – volume 

I: Conservation assessment. Denver, CO: U.S. Department of the 

Interior, Bureau of Land Management.

MacKenzie, D.I.; Nichols, J.D.; Seamans, M.E.; [et al.]. 2009. Modeling 

species occurrence dynamics with multiple states and imperfect 

detection. Ecology. 90: 823–835.

MacKenzie, D.I.; Nichols, J.D.; Sutton, N.; [et al.]. 2005. Improving 

inferences in population studies of rare species that are detected 

imperfectly. Ecology. 86: 1101–1113.

Magoun, A.J.; Copeland, J.P. 1998. Characteristics of wolverine 

reproductive den sites. Journal of Wildlife Management. 62: 

1313–1320. 

Manley, P.N.; Zielinski, W.J.; Schlesinger, M.D.; [et al.]. 2004. 

Evaluation of a multiple-species approach to monitoring species at the 

ecoregional scale. Ecological Applications. 14: 296–310.

McKelvey, K.S.; Aubry, K.B.; Anderson, N.J.; [et al.]. 2014. Recovery 

of wolverines in the western United States: Recent extirpation and 

recolonization or range retraction and expansion. Journal of Wildlife 

Management. 78: 325–334. 

McKelvey, K.S.; Aubry, K.B.; Ortega, Y.K. 2000. History and 

distribution of lynx in the contiguous United States. In: Ruggiero, 

L.F.; Aubry, K.B.; Buskirk, S.W.; [et al.], eds. Ecology and 

conservation of lynx in the United States. Boulder, CO: University 

Press of Colorado; Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station.

McKelvey, K.S.; Claar, J.J.; McDaniel, W.G.; [et al.]. 1999. National 

lynx detection protocol. Washington, DC: U.S. Department of 

Agriculture, Forest Service. 12 p.

McKelvey, K.S.; Kienast, J.V.; Aubry, K.B.; [et al.]. 2006. DNA analysis 

of hair and scat collected along snow tracks to document the presence 

of Canada lynx. Wildlife Society Bulletin. 34: 451–455.
U.S. Forest Service RMRS GTR-388. 2018. 40

Mowat, G.; Poole, K.G.; O’Donoghue, M. 2000. Ecology of lynx 

in Northern Canada and Alaska. In: Ruggiero, L.F.; Aubry, K.B.; 

Buskirk, S.W.; [et al.], eds. Ecology and conservation of Lynx in the 

United State. Boulder, CO: University Press of Colorado Fort Collins, 

CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain 

Research Station.

National Forest Management Act of 1976 [NFMA]; Act of October 22, 

1976; 16 U.S.C. 1600. https://www.gpo.gov/fdsys/granule/STATUTE90/STATUTE-90-Pg2949/content-detail.html. [Accessed May 24, 

2018]. 

Nichols, J.D.; Hines, J.E.; MacKenzie, D.I.; [et al.]. Occupancy 

estimation with multiple states and state uncertainty. Ecology. 88: 

1395–1400.

Nichols, J.D.; Williams, B.K. 2006. Monitoring for conservation. Trends 

in Ecology and Evolution. 21: 668–673.

Nie, M.A.; Schultz, C.A. 2012. Decision-making triggers in adaptive 

management. Conservation Biology. 26: 1137–1144.

Olson, L.E.; Sauder, J.D.; Albrecht, N.M.; [et al.]. 2014. Modeling 

the effects of dispersal and patch size on predicted fisher (Pekania 

[Martes] pennanti) distribution in the U.S. Rocky Mountains. 

Biological Conservation. 169: 89–98. 

Paragi, T.F.; Arthur, S.M.; Krohn, W.B. 1996. Importance of tree cavities 

as natal dens for fishers. National Journal of American Forestry. 13: 

79–83.

Perrine, J.D.; Pollinger, J.P.; Sacks, B.N.; [et al.]. 2007. Genetic evidence 

for the persistence of the critically endangered Sierra Nevada red fox 

in California. Conservation Genetics. 8(5): 1083–1095.

Potvin, F.; Belanger, L.; Lowell, K. 2000. Marten habitat selection in a 

clearcut boreal landscape. Conservation Biology. 14: 844–857.

Proctor, M.; McLellan, B.; Boulanger, J.; [et al.]. 2010. Ecological 

investigations of grizzly bears in Canada using DNA from hair, 1995–

2005: A review of methods and progress. Ursus. 21(2): 169–188.

Prugh, L.R.; Stoner, C.J.; Epps, C.W.; [et al.]. 2009. The rise of the 

mesopredator. BioScience. 59: 779–791.

Raley, C.M.; Lofroth, E.C.; Truex, R.L.; [et al.]. 2012. Habitat ecology of 

fishers in western North America: A new synthesis. In: Aubry, K.B.; 

Zielinski, W.J.; Raphael, M.G.; [et al.], eds. Biology and conservation 

of martens, sables, and fishers: A new synthesis. Ithaca, NY: Cornell 

University Press.

Roemer, G.W.; Gompper, M.E.; Van Valkenburgh, B.; [et al.]. 2009. The 

ecological role of the mammalian mesocarnivore. Bioscience. 59: 

165–173. 
U.S. Forest Service RMRS GTR-388. 2018. 41

Royle, J.A. 2004. Modeling abundance index data from anuran calling 

surveys. Conservation Biology. 18: 1378–1385.

Royle, J.A.; Link, W.A. 2005. A general class of multinomial mixture 

models for anuran calling survey data. Ecology. 86: 2505–2512.

Sacks, B.N.; Statham, M.J.; Perrine, J.D.; [et al.]. 2010. North American 

montane red foxes: Expansion, fragmentation, and the origin of the 

Sacramento Valley red fox. Conservation Genetics. 11:1523–1539.

Sacks, B.N.; Statham, M.J.; Wittmer, H.U. 2017. A preliminary rangewide distribution model for the Sacramento Valley red fox. Journal 

of Fish and Wildlife Management. 8(1): 28–38. e1944-687X. 

doi:10.3996/072016-JFWM-057.

Sauder, J.D.; Rachlow, J.L. 2014. Both forest composition and 

configuration influence landscape-scale habitat selection by fishers 

(Pekania pennanti) in mixed coniferous forests of the Northern Rocky 

Mountains. Forest Ecology and Management. 314: 75–84.

Sauder, J.D.; Rachlow, J.L. 2015. Forest heterogeneity influences habitat 

selection by fishers (Pekania pennanti) within home ranges. Forest 

Ecology and Management. 347: 49–56.

Sauer, J.R.; Fallon, J.E.; Johnson, R. 2003. Use of North American 

breeding bird survey to estimate population change for bird 

conservation regions. Journal of Wildlife Management. 67: 372–389.

Sauer, J.R.; Pardieck, K.L.; Ziolkowski, D.J., Jr.; [et al.]. 2017. The first 

50 years of the North American Breeding Bird Survey. The Condor. 

119: 576–593.

Schultz, C.A.; Sisk, T.D.; Noon, B.R.; [et al.]. 2013. Wildlife 

conservation planning under the United States Forest Service’s 2012 

planning rule. Journal of Wildlife Management. 77: 428–444.

Schwartz, M.K. 2007. Ancient DNA confirms native Rocky Mountain 

fisher (Martes pennanti) avoided early 20th century extinction. 

Journal of Mammalogy. 88: 921–925.

Schwartz, M.K.; Block, W.; Sanderlin, J. 2015. Manage habitat, monitor 

species. In: Morrison, M.L.; Mathewson, H.A. Wildlife habitat 

conservation: Concepts, challenges, and solutions. Baltimore, MD: 

John Hopkins University Press. 

Schwartz, M.K.; Copeland, J.P.; Anderson, N.J.; [et al.]. 2009. Wolverine 

gene flow across a narrow climatic niche. Ecology. 90: 3222–3232. 

Schwartz, M.K.; Ruiz-Gonzalez, A.;, Masuda, R.; [et al.]. 2012. 

Conservation genetics of the genus Martes: Assessing within-species 

movements, units to conserve, and connectivity across ecological and 

evolutionary time. In: Aubry, K.B.; Zielinski, W.J.; Raphael, M.G.; 

[et al.], eds. Biology and conservation of martens, sables, and fishers: 

A new synthesis. Ithaca, NY: Cornell University Press.
U.S. Forest Service RMRS GTR-388. 2018. 42

Schwartz, M.K.; Sanderlin, J.S.; Block, W.A. 2015. Manage habitat, 

monitor species. In: Morrison, M.L.; Mathewson, H.A., eds. Wildlife 

habitat conservation: Concepts, challenges, and solutions. Baltimore, 

MD: Johns Hopkins University Press.

Schwartz, M.K.; Ulizio, T.; Jimenez, B. 2006. U.S. Rocky Mountain 

fisher survey protocol. Missoula, MT: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 13 p.

Slauson, K.M.; Zielinski, W.; Schwartz, M.K. 2017. Ski areas affect 

Pacific marten movement, seasonal habitat use, and density. Journal 

of Wildlife Management. 81: 892–904.

Squires, J.R.; DeCesare, N.J.; Kolbe, J.A.; [et al.]. 2008. Hierarchical 

den selection of Canada lynx in western Montana. Journal of Wildlife 

Management. 72: 1497–1506.

Squires, J.R.; DeCesare, N.J.; Kolbe, J.A.; [et al.]. 2010. Seasonal 

resource selection of Canada lynx in managed forests of the Northern 

Rocky Mountains. Journal of Wildlife Management. 74: 1648–1660.

Squires, J.R.; DeCesare, N.J.; Olson, L.E.; [et al.]. 2013. Combining 

resource selection and movement behavior to predict corridors 

for Canada lynx at their southern range periphery? Biological 

Conservation. 157: 187–195.

Squires, J.R.; Laurion, T. 2000. Lynx home range and movements in 

Montana and Wyoming: Preliminary results. In: Ruggiero, L.F.; 

Aubry, K.B.; Buskirk, S.W.; [et al.]. Ecology and conservation of lynx 

in the United States. Boulder, CO: University Press of Colorado; Fort 

Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky 

Mountain Research Station.

Squires, J.R; McKelvey, K.S.; Ruggiero, L.F. 2004. A snow-tracking 

protocol used to delineate local lynx, Lynx canadensis, distributions. 

Canadian Field-Naturalist. 118: 583–589.

Squires, J.R.; Olson, L.E.; Turner, D.L.; [et al.]. 2012. Estimating 

detection probability for Canada lynx Lynx canadensis using snowtrack surveys in the northern Rocky Mountains, Montana, USA. 

Wildlife Biology.18: 215–224.

Statham, M.J.; Sacks, B.N.; Aubry, K.B.; [et al.]. 2012. The origin 

of recently established red fox populations in the United States: 

Translocations or natural range expansions? Journal of Mammalogy. 

93: 52–65.

Thompson, I.; Fryxell, J.; Harrison, D.; [et al.]. 2012. Improved insights 

into use of habitat by American martens. In: Aubry, K.B.; Zielinski, 

W.J.; Raphael, M.G.; [et al.], eds. Biology and conservation of 

martens, sables, and fishers: A new synthesis. Ithaca, NY: Cornell 

University Press: 209–230.
U.S. Forest Service RMRS GTR-388. 2018. 43

Truex, R.L.; Tucker, J.M.; Bolis, J.S.; [et al.]. 2016. Sierra Nevada Forest 

Plan Amendment (SNFPA) Adaptive Management Strategy: Forest 

Carnivore Monitoring Protocol Phase II. Washington, DC: U.S. 

Department of Agriculture, Forest Service.

USDA Forest Service (USFS). 1986a. Custer National Forest 

management plan. Billings, MT: U.S. Department of Agriculture, 

Forest Service, Custer National Forest.

USDA Forest Service (USFS). 1986b. Forest Plan, Flathead National 

Forest. Kalispell, MT: U.S. Department of Agriculture, Forest Service, 

Flathead National Forest.

USDA Forest Service (USFS). 1986c. Forest Plan, Gallatin National 

Forest. Bozeman, MT: U.S. Department of Agriculture, Forest 

Service, Gallatin National Forest. 

USDA Forest Service (USFS). 1987a. Forest Plan, Bitterroot National 

Forest. Hamilton, MT: U.S. Department of Agriculture, Forest 

Service, Bitterroot National Forest.

USDA Forest Service (USFS). 1987b. Forest Plan, Clearwater National 

Forest. Kamiah, ID: U.S. Department of Agriculture, Forest Service, 

Clearwater National Forest.

USDA Forest Service (USFS). 1988. Forest Plan, Salmon National 

Forest. Salmon, ID: U.S. Department of Agriculture, Forest Service, 

Salmon National Forest. 

USDA Forest Service (USFS). 2007. Northern Rockies lynx management 

direction: Record of decision. Washington, DC: U.S. Department of 

Agriculture, Forest Service.

USDA Forest Service (USFS). 2011. USFS Northern Region (R1) 

sensitive species list. Missoula, MT: U.S. Department of Agriculture, 

Forest Service, Northern Region. 

USDA Forest Service (USFS). 2012. Final programmatic environmental 

impact statement: National Forest System land management planning. 

Washington, DC: U.S. Department of Agriculture, Forest Service.

USDA Forest Service (USFS). 2013a. Forest plan monitoring and 

evaluation report: Fiscal years 2010–2013. Bitterroot National Forest. 

Hamilton, MT: U.S. Department of Agriculture, Forest Service, 

Bitterroot National Forest.

USDA Forest Service (USFS). 2013b. Nez Perce-Clearwater National 

Forest list of potential species of conservation concern. Kamiah, 

ID: U.S. Department of Agriculture, Forest Service, Nez PerceClearwater National Forest. https://www.fs.usda.gov/Internet/FSE_

DOCUMENTS/stelprd3807171.pdf. [Accessed March 3, 2017]. 
U.S. Forest Service RMRS GTR-388. 2018. 44

USDA Forest Service (USFS). 2015. Land management planning 

handbook, Chapter 30 monitoring. (FSH) 1909.12. Washington, DC: 

U.S. Department of Agriculture, Forest Service. 

USDA Forest Service (USFS). 2016. Intermountain Region (R4) 

threatened, endangered, proposed, and, sensitive species. Ogden, 

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

Region. 

USDA Forest Service (USFS and USFWS). 2006. Canada Lynx 

Conservation Agreement. USFS Agreement #00-MU-11015600-013. 

Missoula, MT: U.S. Department of Agriculture, Forest Service; U.S. 

Department of the Interior, U.S. Fish and Wildlife Service. 13 p.

U.S. Fish and Wildlife Service. 2017 (USFWS). Species status 

assessment for the Canada lynx (Lynx canadensis): Contiguous 

United States distinct population segment. Version 1.0. October, 

2017. Lakewood, CO: U.S. Department of the Interior, U.S. Fish and 

Wildlife Service. 292 p.

Utah Division of Wildlife Resources. 2016. Wolverine found in Rich 

County. Wildlife News. July 1, 2016. https://wildlife.utah.gov/

wildlife-news/1872-wolverine-found-in-rich-county.html. [Accessed 

May 25, 2017].

Vinkey, R.S. 2003. An evaluation of fisher (Martes pennanti) 

introductions in Montana. Thesis. Missoula, MT: University of 

Montana. 

Vinkey, R.S.; Schwartz, M.K.; McKelvey, K.S.; [et al.]. 2006. When 

reintroductions are augmentations: The genetic legacy of fishers 

(Martes pennanti) in Montana. Journal of Mammalogy. 87: 265–271.

Washington Department of Fish and Wildlife. 2013. Threatened and 

endangered wildlife in Washington: 2012 Annual Report. Olympia, 

WA: Washington Department of Fish and Wildlife, Wildlife Program, 

Listing and Recovery Section. 251 p.

Washington Department of Natural Resources. 2017. 2017 Endangered, 

threatened, and sensitive vascular plant list. Olympia, WA: 

Washington State Department of Natural Resources, Natural Heritage 

Program. 10 p. [April 2017].

Zielinski, W.J.; Baldwin, J.A.; Truex, R.L.; [et al.] 2013. Estimating 

trend in occupancy for the southern Sierra fisher Martes pennanti

population. Journal of Fish and Wildlife Management. 4(1): 3–19.
U.S. Forest Service RMRS GTR-388. 2018. 45

Appendix A: Regulatory and Legal Status 

of Primary and Secondary Species

Forest Service Regions mentioned are as follows: Region 1 = Northern; 

Region 2 = Rocky Mountain; Region 3 = Southwestern; Region 4 = 

Intermountain; Region 5 = Pacific Southwest; Region 6 = Pacific 

Northwest; Region 9 = Eastern.

Status: Regional forester’s sensitive species

Scale: USDA FS Region

USDA FS location: Region 1, Region 4

Status: Proposed species of conservation concern

Scale: National forest

USDA FS location: Nez Perce-Clearwater National Forest

Status: ESA threatened

Scale: Species’ known range

USDA FS location: Region 1, Region 2, Region 3, Region 4, Region 9

Status: Regional forester’s sensitive species

Scale: USDA FS Region

USDA FS location: Region 1

Status: ESA proposed threatened

Scale: Species’ known range

USDA FS location: Region 1, Region 4, Region 5

Status: Management indicator species

Scale: National forest

USDA FS location: Bitterroot National Forest, Clearwater National 

Forest, Custer-Gallatin National Forest, Flathead National Forest, 

Salmon-Challis National Forest 

Status: None

Scale: Not applicable

USDA FS location: Not applicable

Species: Fisher

Species: Lynx

Species: Wolverine

Species: Marten 

(American and Pacific)

Species: 

Montane red fox
U.S. Forest Service RMRS GTR-388. 2018. 46

Appendix B: History of ESA Listing Petitions 

and Decisions for Primary Species 

Date: February 24, 2009

Action: Petition for a rule to designate fishers in the northern Rocky 

Mountain region as a Distinct Population Segment (DPS) and add 

them to the list of endangered or threatened wildlife protected by the 

Endangered Species Act (ESA)

Date: April 9, 2009

Action: Response to 2009 petition stating that emergency listing was not 

warranted and a 90-day review would commence in 2011

Date: June 30, 2011

Action: 12-month finding that listing of the fisher in the northern Rocky 

Mountain region was not warranted 

Date: September 23, 2013

Action: Petition for a rule to add the northern Rocky Mountain fisher 

DPS to the list of endangered or threatened wildlife protected by the 

ESA

Date: October 31, 2013

Action: Response to 2013 petition stating that emergency listing was not 

warranted

Date: January 12, 2016

Action: 90-day finding that listing of the northern Rocky Mountain 

fisher DPS may be warranted under the ESA based on factors B 

(overutilization) and E (other factors)

Date: October 5, 2017

Action: 12-month finding that listing of the northern Rocky Mountain 

fisher DPS as threatened or endangered under the ESA is not 

warranted

Date: August 23, 1994

Action: Petition for a rule to add Canada lynx to the list of endangered or 

threatened wildlife protected by the ESA

Date: August 26, 1994

Action: 90-day finding that listing of Canada lynx may be warranted

Date: December 27, 1994

Action: 12-month finding that listing of Canada lynx was not warranted

Date: March 27, 1997

Action: Court order remanded the 1994 Canada lynx 12-month finding 

back to the USFWS for reconsideration

Species: Fisher

Species: Lynx
U.S. Forest Service RMRS GTR-388. 2018. 47

Date: May 27, 1997

Action: 12-month finding that listing of Canada lynx was warranted but 

precluded and would be added to the list of candidate species

Date: July 8, 1998

Action: Proposed rule to list the lynx as threatened under the ESA

Date: March 24, 2000

Action: Canada lynx in the contiguous United States considered a DPS 

and listed as a threatened species under the ESA based on factor D of 

the ESA (inadequacy of existing regulatory mechanisms)

Date: November 13, 2017

Action: USFWS published a 5-year review and recommendation to 

proceed with proposed rule to remove the Canada lynx DPS from the 

threatened and endangered species list

Date: August 3, 1994

Action: Petition for a rule to add wolverine in the contiguous United 

States to the list of endangered or threatened wildlife protected by 

the ESA

Date: April 19, 1995

Action: August 3, 1994 petition did not present substantial information 

indicating that listing was warranted

Date: July 4, 2000

Action: Petition for a rule to add wolverine in the contiguous United 

States to the list of endangered or threatened wildlife protected by 

the ESA and designate critical habitat 

Date: October 21, 2003

Action: 90-day finding that the July 4, 2000 petition did not present 

substantial scientific and commercial information indicating that 

listing was warranted

Date: September 29, 2006

Action: Ruled that the 90-day finding was in error and ordered U.S. Fish 

and Wildlife Service (USFWS) to make a 12-month finding for the 

wolverine

Date: April 6, 2007

Action: Extended deadline for 12-month finding to February 28, 2008

Date: June 5, 2007

Action: Notice of status review initiation 

Date: March 11, 2008

Action: 12-month finding that wolverine did not constitute a DPS and 

therefore was not a listable entity under the ESA

Date: July 8, 2008

Action: Complaint challenging 12-month finding

Species: Wolverine

Species: Lynx 

(continued)
U.S. Forest Service RMRS GTR-388. 2018. 48

Date: March 6, 2009

Action: Settlement from the July 8, 2008 challenge where the USFWS 

agreed to submit a new 12-month finding to the Federal Register by 

December 1, 2010

Date: April 15, 2010

Action: Initiation of status review for wolverine

Date: December 14, 2010

Action: 12-month finding that wolverine constitutes a DPS and was 

warranted for listing under the ESA but precluded by other higher 

priorities

Date: April 13, 2012

Action: Challenge of 12-month finding

Date: September 20, 2012

Action: Court granted stay to USFWS

Date: February 4, 2013

Action: Proposed rule to list wolverine as threatened

Date: October 31, 2013

Action: Reopened rule for public comment

Date: February 5, 2014

Action: 6-month extension of final determination

Date: August 13, 2014

Action: Withdrawal of rule to list “based on our conclusion that the 

factors affecting the DPS as identified in the proposed rule were 

not as significant as believed at the time of the proposed rule’s 

publication in 2013”

Date: August 13, 2014

Action: Challenge of withdrawal

Date: April 4, 2016

Action: USFWS ordered to reconsider the rule

Date: October 18, 2016

Action: Reopened comment period on February 4, 2013 rule (comment 

period closed November 17, 2016)

Species: Wolverine 

(continued)
U.S. Forest Service RMRS GTR-388. 2018. 49

Appendix C: Northern Rockies Lynx Management Direction 

2006: Occupied and Unoccupied Lynx Habitat

Reference USDA Forest Service (USFS and USFWS). 2006. Canada Lynx 

Conservation Agreement. USFS Agreement #00-MU-11015600-013. 

Missoula, MT: U.S. Department of Agriculture, Forest Service; U.S. 

Department of the Interior, U.S. Fish and Wildlife Service. 13 p.

Figure C1—Lynx habitat in the northern Rocky Mountains. Figure source: USFS and USFWS (2006).
U.S. Forest Service RMRS GTR-388. 2018. 50

Appendix D—Mesocarnivore Information Needs 

in the Northern Region (Region 1) 

and the Intermountain Region (Region 4)

Staff from RMRS, Region 1, and Region 4 held meetings in July, 

August, and September 2017 with each national forest, including 

forest biologists and ecological staff officers on each national forest, to 

determine the information needs of the forest, namely where the national 

forest wanted to ask each of the monitoring questions for the primary 

species. The information provided here are the follow-up questionnaires 

that were sent to the national forests in the Northern Region 

(Region 1) and Intermountain Region (Region 4) after these meetings. 

Not all questionnaires were returned, so only those that were returned 

are presented.
U.S. Forest Service RMRS GTR-388. 2018. 51
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U.S. Forest Service RMRS GTR-388. 2018. 56
U.S. Forest Service RMRS GTR-388. 2018. 57
U.S. Forest Service RMRS GTR-388. 2018. 58
U.S. Forest Service RMRS GTR-388. 2018. 59
U.S. Forest Service RMRS GTR-388. 2018. 60

Appendix E: Example Multispecies Mesocarnivore 

Monitoring Form

The following is a hypothetical example of monitoring on the 

Bitterroot National Forest over a 4-year period with the GEM monitoring 

approach for fishers, lynx, and wolverines. It does not reflect any 

real monitoring or species occurrence data and is not intended to be a 

representation of what is present on the Bitterroot National Forest. 

	 For all maps, habitat is grouped by areas of ≥0.25 value from Olson

et al. (2014) and areas within 2 km of those patches (half of a small 

fisher home range [16 km2

] radius). Only groups ≥16 km2 are included.

The 15 km × 15 km (225 km2) grid cells selected were based on cells 

with ≥25 percent or more wolverine habitat.

Copeland, J.P.; McKelvey, K.S.; Aubry, K.B.; [et al.]. 2010. The 

bioclimatic envelope of the wolverine (Gulo gulo): Do climatic 

constraints limit its geographic distribution? Canadian Journal of 

Zoology. 88: 233–246.

Olson, L.E.; Sauder, J.D.; Albrecht, N.M.; [et al.]. 2014. Modeling 

the effects of dispersal and patch size on predicted fisher (Pekania 

[Martes] pennanti) distribution in the U.S. Rocky Mountains. 

Biological Conservation. 169: 89–98. 

Schwartz, M.K.; Ulizio, T.; Jimenez, B. 2006. U.S. Rocky Mountain 

Fisher Survey Protocol. Missoula, MT: U.S. Department of 

Agriculture, Forest Service, Rocky Mountain Research Station. 13 p. 

Squires, J.R.; McKelvey, K.S.; and Ruggiero, L.F. 2004. A snow-tracking 

protocol used to delineate local lynx, Lynx canadensis, distributions. 

Canadian Field-Naturalist. 118: 583–589.

USDA Forest Service (USFS and USFWS). 2006. Canada Lynx 

Conservation Agreement. USFS Agreement #00-MU-11015600-013. 

Missoula, MT: U.S. Department of Agriculture, Forest Service; U.S. 

Department of the Interior, U.S. Fish and Wildlife Service. 13 p.

USDA Forest Service (USFS). 2014. Lynx monitoring protocol in 

unoccupied lynx habitat. Unpublished report on file with: U.S. 

Department of Agriculture, Forest Service, Northern Region, 

Missoula, MT. 2 p.

References
U.S. Forest Service RMRS GTR-388. 2018. 61
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U.S. Forest Service RMRS GTR-388. 2018. 65
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In accordance with Federal civil rights law and U.S. Department of Agriculture (USDA) 

civil rights regulations and policies, the USDA, its Agencies, offices, and employees, 

and institutions participating in or administering USDA programs are prohibited from 

discriminating based on race, color, national origin, religion, sex, gender identity (including 

gender expression), sexual orientation, disability, age, marital status, family/parental status, 

income derived from a public assistance program, political beliefs, or reprisal or retaliation 

for prior civil rights activity, in any program or activity conducted or funded by USDA (not 

all bases apply to all programs). Remedies and complaint filing deadlines vary by program 

or incident. 

Persons with disabilities who require alternative means of communication for program 

information (e.g., Braille, large print, audiotape, American Sign Language, etc.) should 

contact the responsible Agency or USDA’s TARGET Center at (202) 720-2600 (voice and 

TTY) or contact USDA through the Federal Relay Service at (800) 877-8339. Additionally, 

program information may be made available in languages other than English.

To file a program discrimination complaint, complete the USDA Program Discrimination 

Complaint Form, AD-3027, found online at http://www.ascr.usda.gov/complaint_filing_

cust.html and at any USDA office or write a letter addressed to USDA and provide in 

the letter all of the information requested in the form. To request a copy of the complaint 

form, call (866) 632-9992. Submit your completed form or letter to USDA by: (1) mail: 

U.S. Department of Agriculture, Office of the Assistant Secretary for Civil Rights, 

1400 Independence Avenue, SW, Washington, D.C. 20250-9410; (2) fax: (202) 690-7442; 

or (3) email: program.intake@usda.gov.

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