# 84 FR 9648: Endangered and Threatened Wildlife and Plants; Removing the Gray Wolf (Canis lupus) From the List of Endangered and Threatened Wildlife

> Federal · Regulations · In force

URL: https://www.frixlaw.com/law-library/statutes/FR_PRORULE_2019-04420

## Section

- **Citation:** 84 FR 9648
- **Heading:** Endangered and Threatened Wildlife and Plants; Removing the Gray Wolf (Canis lupus) From the List of Endangered and Threatened Wildlife
- **Jurisdiction:** Federal
- **Kind:** Regulations
- **Status:** In force
- **Text as of:** August 14, 2026
- **Source:** Compiled text
- **Location:** Federal Register / Vol. 84 / 84 FR 9648

## Text

DEPARTMENT OF THE INTERIOR Fish and Wildlife Service 50 CFR Part 17 [Docket No. FWS-HQ-ES-2018-0097; FXES11130900000C2-189-FF09E32000] RIN 1018-BD60 Endangered and Threatened Wildlife and Plants; Removing the Gray Wolf ( Canis lupus ) From the List of Endangered and Threatened Wildlife AGENCY:
Fish and Wildlife Service, Interior.

ACTION:
Proposed rule.

SUMMARY:
We, the U.S. Fish and Wildlife Service (Service or USFWS), have evaluated the classification status of gray wolves ( Canis lupus ) currently listed in the contiguous United States and Mexico under the Endangered Species Act of 1973, as amended (Act). Based on our evaluation, we propose to remove the gray wolf from the List of Endangered and Threatened Wildlife. We propose this action because the best available scientific and commercial information indicates that the currently listed entities do not meet the definitions of a threatened species or endangered species under the Act due to recovery. The effect of this rulemaking action would be to remove the gray wolf from the Act's protections. This proposed rule does not have any effect on the separate listing of the Mexican wolf ( Canis lupus baileyi ) as endangered under the Act.

DATES:
Comment submission: We will accept comments received or postmarked on or before May 14, 2019.
Public hearings: We must receive requests for public hearings, in writing, at the address shown in FOR FURTHER INFORMATION CONTACT by April 29, 2019.

ADDRESSES:
You may submit comments by one of the following methods:
Mexican wolf ( Canis lupus baileyi ) as endangered under the Act.

DATES:
Comment submission: We will accept comments received or postmarked on or before May 14, 2019.
Public hearings: We must receive requests for public hearings, in writing, at the address shown in FOR FURTHER INFORMATION CONTACT by April 29, 2019.

ADDRESSES:
You may submit comments by one of the following methods:
(1) Electronically: Go to the Federal eRulemaking Portal: http://www.regulations.gov. In the Search box, enter Docket No. FWS-HQ-ES-2018-0097, which is the docket number for this rulemaking. Then, click on the Search button. On the resulting page, in the Search panel on the left side of the screen under the Document Type heading, click on the Proposed Rules link to locate this document. You may submit a comment by clicking on the blue “Comment Now!” box. If your comments will fit in the provided comment box, please use this feature of http://www.regulations.gov, as it is most compatible with our comment review procedures. If you attach your comments as a separate document, our preferred file format is Microsoft Word. If you attach multiple comments (such as form letters), our preferred format is a spreadsheet in Microsoft Excel.
(2) By hard copy: Submit by U.S. mail or hand-delivery to: Public Comments Processing, Attn: Docket No. FWS-HQ-ES-2018-0097; U.S. Fish & Wildlife Service Headquarters, MS: BPHC, 5275 Leesburg Pike, Falls Church, VA 22041-3803.
We request that you send comments only by the methods described above. We will post all comments on http://www.regulations.gov. This generally means that we will post any personal information you provide us (see Public Comments below for more information).
FOR FURTHER INFORMATION CONTACT:
Don Morgan, Chief, Branch of Delisting and Foreign Species, Ecological Services, U.S. Fish and Wildlife Service, Headquarters Office, MS: ES, 5275 Leesburg Pike, Falls Church, VA 22041-3803; telephone (703) 358-2444
w.regulations.gov. This generally means that we will post any personal information you provide us (see Public Comments below for more information).
FOR FURTHER INFORMATION CONTACT:
Don Morgan, Chief, Branch of Delisting and Foreign Species, Ecological Services, U.S. Fish and Wildlife Service, Headquarters Office, MS: ES, 5275 Leesburg Pike, Falls Church, VA 22041-3803; telephone (703) 358-2444. Persons who use a telecommunications device for the deaf (TDD) may call the Federal Relay Service at 800-877-8339.

SUPPLEMENTARY INFORMATION:
Executive Summary
Purpose of the Regulatory Action
Why we need to publish a rule. Under the Act, if we determine that a species is no longer threatened or endangered throughout all or a significant portion of its range, we must publish in the Federal Register a proposed rule to remove the species from the Lists of Endangered and Threatened Wildlife and Plants in title 50 of the Code of Federal Regulations (50 CFR 17.11 and 17.12). We also must make a final determination on our proposal within 1 year thereafter. Removing a species from the List (“delisting” it) can only be completed by issuing a rule.
This document proposes delisting gray wolves in the lower 48 United States and Mexico. This proposed rule assesses the best available information regarding the status of and threats to the species, and replaces our June 13, 2013, proposed rule to delist the gray wolf in the lower 48 United States and Mexico (78 FR 35664). This proposed rule does not have any effect on the separate listing of the Mexican wolf as endangered under the Act (80 FR 2487, January 16, 2015).
The basis for our action
ed rule assesses the best available information regarding the status of and threats to the species, and replaces our June 13, 2013, proposed rule to delist the gray wolf in the lower 48 United States and Mexico (78 FR 35664). This proposed rule does not have any effect on the separate listing of the Mexican wolf as endangered under the Act (80 FR 2487, January 16, 2015).
The basis for our action. Under the Act, we determine whether a species is an endangered or threatened species based on any one or more of five factors or the cumulative effects thereof: (A) The present or threatened destruction, modification, or curtailment of its habitat or range; (B) Overutilization for commercial, recreational, scientific, or educational purposes; (C) Disease or predation; (D) The inadequacy of existing regulatory mechanisms; or (E) Other natural or manmade factors affecting its continued existence. We have determined that the gray wolf in the lower 48 United States and Mexico (except the Mexican wolf subspecies) no longer meets the definition of an endangered or threatened species under the Act.
Peer review. We will seek comments from independent specialists to ensure that our designation is based on scientifically sound data, assumptions, and analyses. We will invite these peer reviewers to comment on our listing proposal. Because we will consider all comments and information received during the comment period, our final determination may differ from this proposal.
Information Requested
Public Comments
We intend that any final action resulting from this proposal will be based on the best scientific and commercial data available and will be as accurate and as effective as possible. Therefore, we request comments or information from the public, concerned Tribal and governmental agencies, the scientific community, industry, or any other interested parties concerning this proposed rule. Comments should be as specific as possible
n resulting from this proposal will be based on the best scientific and commercial data available and will be as accurate and as effective as possible. Therefore, we request comments or information from the public, concerned Tribal and governmental agencies, the scientific community, industry, or any other interested parties concerning this proposed rule. Comments should be as specific as possible.
As this proposal replaces our June 13, 2013, proposal to delist gray wolves in the lower 48 United States and Mexico (78 FR 35663), we ask that any comments previously submitted that are relevant to the status of wolves currently listed in the contiguous United States and Mexico, as analyzed in this rule, be resubmitted at this time. Comments must be submitted during the comment period for this proposed rule to be considered.
Please include sufficient information with your submission (such as scientific journal articles or other publications) to allow us to verify any scientific or commercial information you include.
Please note that submissions merely stating support for, or opposition to, the action under consideration without providing supporting information, although noted, will not meet the standard of best available scientific and commercial data. Section 4(b)(1)(A) of the Act directs that determinations as to whether any species is threatened or endangered must be made “solely on the basis of the best scientific and commercial data available.”
You may submit your comments and materials by one of the methods listed in ADDRESSES . We request that you send comments only by the methods described in ADDRESSES .
If you submit information via http://www.regulations.gov, your entire submission—including your personal identifying information—will be posted on the website. If your submission is made via a hardcopy that includes personal identifying information, you may request at the top of your document that we withhold this information from public review
mments only by the methods described in ADDRESSES .
If you submit information via http://www.regulations.gov, your entire submission—including your personal identifying information—will be posted on the website. If your submission is made via a hardcopy that includes personal identifying information, you may request at the top of your document that we withhold this information from public review. However, we cannot guarantee that we will be able to do so. We will post all hardcopy submissions on http://www.regulations.gov.
Comments and materials we receive, as well as supporting documentation we used in preparing this proposed rule, will be available for public inspection on http://www.regulations.gov at Docket No. FWS-HQ-ES-2018-0097, or by appointment, during normal business hours at U.S. Fish and Wildlife Service Headquarters (see FOR FURTHER INFORMATION CONTACT ).
Peer Review
In accordance with our joint policy on peer review published in the Federal Register on July 1, 1994 (59 FR 34270), we will seek the expert opinions of at least three appropriate and independent specialists regarding scientific data and interpretations contained in this proposed rule. The purpose of peer review is to ensure that our decisions are based on scientifically sound data, assumptions, and analyses. We will invite these peer reviewers to comment during the public comment period on our proposed action; these comments will be available along with other public comments in the docket for this proposed rule.
We will consider all comments and information we receive during this comment period during our preparation of the final determination. Accordingly, the final decision may differ from this proposal.
Table of Contents
Previous Federal Actions General Background The 1978 Reclassification National Wolf Strategy Approach for this Proposed Rule The Entities Addressed in this Rule How We Address the C
e will consider all comments and information we receive during this comment period during our preparation of the final determination. Accordingly, the final decision may differ from this proposal.
Table of Contents
Previous Federal Actions General Background The 1978 Reclassification National Wolf Strategy Approach for this Proposed Rule The Entities Addressed in this Rule How We Address the C. lupus Entities in this Rule How We Address Taxonomic Uncertainties in this Rule Summary of Our Approach Species Information Biology and Ecology Taxonomy of Gray Wolves in North America Range and Population Trends Prior to 1978 Reclassification Historical Range of the Gray Wolf Entity Historical Abundance of the Gray Wolf Entity Historical Trends in Range and Abundance for the Gray Wolf Entity Distribution, and Abundance of the Gray Wolf Entity at the Time of the 1978 Reclassification Current Distribution and Abundance of the Gray Wolf Entity Gray Wolf Recovery Plans and Recovery Implementation Recovery Criteria Recovery Progress Historical Context of Our Analysis Summary of Factors Affecting the Species Human-caused Mortality Effects on Wolf Social Structure The Role of Public Attitudes Human-caused Mortality Summary Habitat and Prey Availability Great Lakes Area: Suitable Habitat Great Lakes Area: Prey Availability West Coast States: Suitable Habitat West Coast States: Prey Availability Habitat and Prey Availability Summary Disease and Parasites Effects of Climate Change Cumulative Effects Post-delisting Management State Management Post-delisting Management in Minnesota, Wisconsin, and Michigan The Minnesota Wolf Management Plan Depredation Control in Minnesota Post-delisting Depredation Control in Minnesota Post-delisting Regulated Harvest in Minnesota The Wisconsin Wolf Management Plan Depredation Control in Wisconsin Post-delisting Depredation Control in Wisconsin Post-delisting Regulated Harvest in Wisconsin The Michigan Wolf Management Plan Depredation Control in Michigan Post-delisting Depredation Control in Mic
lan Depredation Control in Minnesota Post-delisting Depredation Control in Minnesota Post-delisting Regulated Harvest in Minnesota The Wisconsin Wolf Management Plan Depredation Control in Wisconsin Post-delisting Depredation Control in Wisconsin Post-delisting Regulated Harvest in Wisconsin The Michigan Wolf Management Plan Depredation Control in Michigan Post-delisting Depredation Control in Michigan Post-delisting Regulated Harvest in Michigan Post-delisting Management in the West Coast States The Oregon Wolf Management Plan The Washington Wolf Management Plan The California Wolf Management Plan Tribal Management and Conservation of Wolves Management on Federal Lands Great Lakes Area West Coast States Summary of Post-delisting Management Determination of Species Status Summary and Conclusion of Our Analysis Determination of Status Throughout All of its Range Determination of Status Throughout a Significant Portion of its Range Proposed Determination Effects of This Rule Post-delisting Monitoring Required Determinations Clarity of This Proposed Rule National Environmental Policy Act Government-to-Government Relationship With Tribes Previous Federal Actions
Gray wolves were originally listed as subspecies or as regional populations of subspecies in the contiguous United States and Mexico. Early listings were under legislative predecessors of the Act—the Endangered Species Preservation Act of 1966 and the Endangered Species Conservation Act of 1969. Later listings were under the Endangered Species Act of 1973. The Federal Register citations for all the rulemaking actions described in the following paragraphs are provided in table 1, below.
In 1978, we published a rule reclassifying the gray wolf as an endangered population at the taxonomic species level ( C. lupus ) throughout the contiguous United States and Mexico, except for the Minnesota gray wolf population, which was classified as threatened (table 1)
deral Register citations for all the rulemaking actions described in the following paragraphs are provided in table 1, below.
In 1978, we published a rule reclassifying the gray wolf as an endangered population at the taxonomic species level ( C. lupus ) throughout the contiguous United States and Mexico, except for the Minnesota gray wolf population, which was classified as threatened (table 1). At that time, we considered the gray wolves in Minnesota to be a listable entity under the Act, and we considered gray wolves in Mexico and the 48 contiguous United States other than Minnesota to be another listable entity (43 FR 9607 and 9610, respectively, March 9, 1978). The earlier subspecies listings thus were subsumed into the listings for the gray wolf in Minnesota and the gray wolf in the rest of the contiguous United States and Mexico.
The 1978 reclassification was undertaken to “most conveniently” address changes in our understanding of gray wolf taxonomy and protect all gray wolves in the lower 48 United States. In addition, we sought to clarify that the gray wolf was only listed south of the Canadian border.
The 1978 reclassification rule stipulated that “biological subspecies would continue to be maintained and dealt with as separate entities” (43 FR 9609), and offered “the firmest assurance that [the Service] will continue to recognize valid biological subspecies for purposes of its research and conservation programs” (43 FR 9610). Accordingly, we implemented three gray wolf recovery programs in three regions of the country—the northern Rocky Mountains, the southwestern United States, and the eastern United States—to establish and prioritize recovery criteria and actions appropriate to the unique local circumstances of the gray wolf (table 1)
al subspecies for purposes of its research and conservation programs” (43 FR 9610). Accordingly, we implemented three gray wolf recovery programs in three regions of the country—the northern Rocky Mountains, the southwestern United States, and the eastern United States—to establish and prioritize recovery criteria and actions appropriate to the unique local circumstances of the gray wolf (table 1). Recovery in two of these regions (northern Rocky Mountains and southwestern United States) required reintroduction of gray wolves in experimental populations (table 1),
Between 2003 and 2015, we published several rules revising the 1978 contiguous United States and Mexico listings for C. lupus in an attempt to acknowledge taxonomy, comport with current policy and practices, and to recognize the biological recovery of gray wolves in the northern Rocky Mountains (NRM) and western Great Lakes (WGL) populations. Previous rules were challenged and subsequently invalidated or vacated by various courts based, in part, on their determinations that our distinct population segment (DPS) designations were legally flawed (table 1).
Of particular relevance to this proposed rule is our 2011 final rule, in which we recognized the expansion of the Minnesota wolf population by revising the entity to include all or portions of six surrounding States, identified the expanded population as the western Great Lakes DPS (WGL DPS), and revised the listings to remove the WGL DPS from the List due to recovery. Also in 2011, we published a final rule that implemented Section 1713 of Public Law 112-10, reinstating our 2009 delisting rule for the NRM DPS and, with the exception of Wyoming, removed gray wolves in that DPS from the List. In 2012, we finalized a rule removing gray wolves in Wyoming from the List. Subsequently, in 2013, we published a proposed rule to delist C
PS from the List due to recovery. Also in 2011, we published a final rule that implemented Section 1713 of Public Law 112-10, reinstating our 2009 delisting rule for the NRM DPS and, with the exception of Wyoming, removed gray wolves in that DPS from the List. In 2012, we finalized a rule removing gray wolves in Wyoming from the List. Subsequently, in 2013, we published a proposed rule to delist C. lupus in the remaining listed portions of the United States and Mexico outside of the delisted NRM and WGL DPSs, and keep Mexican wolf listed as an endangered subspecies, C. l. baileyi (table 1).
However, in 2014 the United States District Court for the District of Columbia vacated the final rule at 76 FR 81666 (December 28, 2011) that removed protections of the Act from the gray wolf in the western Great Lakes (table 1). The court's action was based, in part, on its conclusion that the Act does not allow the Service to use its authority to identify DPSs as “species” to remove the protections for part of an already listed species. The U.S. Court of Appeals disagreed, ruling in 2017 that the Service had the authority to designate a DPS from a larger listed entity and delist it in the same rule (table 1). That court nonetheless upheld the District Court's vacatur, concluding that the Service failed to reasonably analyze or consider two significant aspects of the rule: The impacts of partial delisting and historical range loss on the remainder of the listed entity.
Our 2012 decision to delist gray wolves in Wyoming was also vacated by the U.S. District Court for the District of Columbia. Because the 2013 proposal to delist the remaining listed portions of the gray wolf in the United States and Mexico relied in part on two subsequently vacated final rules, the 2011 WGL DPS rule as well as our 2012 rule delisting gray wolves in Wyoming, in 2015 we only finalized the portion of the rule listing the Mexican wolf as an endangered subspecies (table 1). In 2017, the D.C
rict of Columbia. Because the 2013 proposal to delist the remaining listed portions of the gray wolf in the United States and Mexico relied in part on two subsequently vacated final rules, the 2011 WGL DPS rule as well as our 2012 rule delisting gray wolves in Wyoming, in 2015 we only finalized the portion of the rule listing the Mexican wolf as an endangered subspecies (table 1). In 2017, the D.C. Circuit reversed the district court's decision and reinstated the delisting of gray wolves in Wyoming. Thus, wolves are currently delisted in the entire northern Rocky Mountains area (figure 1).
As a result of the above actions, the C. lupus listings in 50 CFR 17.11 currently include: (1) C. lupus in Minnesota listed as threatened, and (2) C. lupus in all or portions of 44 U.S. States and Mexico, listed as endangered (figure 1). In the United States, this includes: all of Alabama, Arkansas, California, Colorado, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Massachusetts, Maryland, Maine, Michigan, Missouri, Mississippi, North Carolina, North Dakota, Nebraska, New Hampshire, New Jersey, Nevada, New York, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, South Dakota, Tennessee, Texas, Virginia, Vermont, West Virginia, and Wisconsin; and portions of Arizona, New Mexico, Oregon, Utah, and Washington (figure 1).
For additional information on these Federal actions and their associated litigation history refer to the relevant associated rules or the Previous Federal Actions sections of our recent gray wolf actions (see table 1).
Table 1—Key Federal Regulatory Actions Under the Act and Predecessor Legislation 1 Pertaining to Gray Wolf and, Where Applicable, Outcomes of Court Challenges to These Actions [E = Endangered Species, T = Threatened Species, DPS = Distinct Population Segment, NRM = Northern Rocky Mountains, WGL = Western Great Lakes] Entity Year of action Type of action Federal Register citation Litigation history C. l
1—Key Federal Regulatory Actions Under the Act and Predecessor Legislation 1 Pertaining to Gray Wolf and, Where Applicable, Outcomes of Court Challenges to These Actions [E = Endangered Species, T = Threatened Species, DPS = Distinct Population Segment, NRM = Northern Rocky Mountains, WGL = Western Great Lakes] Entity Year of action Type of action Federal Register citation Litigation history C. l. lycaon 1967 1 List 32 FR 4001, March 11, 1967 C. l. irremotus 1973 1 List 38 FR 14678, June 4, 1973 C. l. lycaon 1974 List 39 FR 1171, January 4, 1974 C. l. irremotus 1974 List 39 FR 1171, January 4, 1974 C. l. baileyi 1976 List (E) 41 FR 17736, April 28, 1976 C. l. monstrabilis 2 1976 List (E) 41 FR 24064, June 14, 1976 C. lupus in lower 48 U.S. (except Minnesota) & Mexico 1978 Reclassify (E) 43 FR 9607, March 9, 1978 3 C. lupus in Minnesota 1978 Reclassify (T) 43 FR 9607, March 9, 1978 3 C. lupus 1978 (revised 1992) Recovery Plan for Eastern Timber Wolf (eastern gray wolf) n.a. C. lupus 1980 (revised 1987) Recovery Plan for NRM Gray Wolf n.a. C. lupus 1982 (revised 2017) Recovery Plan for Mexican Gray Wolf ( C. l. baileyi ) n.a. C. lupus 1994 Establish experimental population (southeastern Idaho, southern Montana, and Wyoming) 59 FR 60266, November 22, 1994 C. lupus 1994 Establish experimental population (central Idaho & southwest Montana) 59 FR 60252, November 22, 1994 C. lupus 1998 Establish experimental population (Arizona & New Mexico) 63 FR 1752, January 12, 1998 C. lupus DPSs: —Eastern DPS —Western DPS —Southwestern U.S. & Mexico DPS 2003 Designate DPS & classify/reclassify as: —Eastern DPS (T) —Western DPS (T) —Southwestern U.S. & Mexico DPS (E) Delist in unoccupied non-historical range 68 FR 15804, April 1, 2003 Rule vacated ( Defenders of Wildlife v. Norton, 354 F. Supp. 2d 1156 (D. Or. 2005); National Wildlife Federation v. Norton, 386 F. Supp. 2d 553 (D. Vt. 2005)) C. lupus WGL DPS 2007 Designate DPS & delist 72 FR 6052, February 8, 2007 Rule vacated ( Humane Society of the United States v
n DPS (T) —Southwestern U.S. & Mexico DPS (E) Delist in unoccupied non-historical range 68 FR 15804, April 1, 2003 Rule vacated ( Defenders of Wildlife v. Norton, 354 F. Supp. 2d 1156 (D. Or. 2005); National Wildlife Federation v. Norton, 386 F. Supp. 2d 553 (D. Vt. 2005)) C. lupus WGL DPS 2007 Designate DPS & delist 72 FR 6052, February 8, 2007 Rule vacated ( Humane Society of the United States v. Kempthorne, 579 F. Supp. 2d 7 (D.D.C. 2008)) C. lupus NRM DPS 2008 Designate DPS & delist 73 FR 10514, February 27, 2008 Rule vacated and remanded ( Defenders of Wildlife v. Hall, 565 F. Supp. 2d 1160 (D. Mont. 2008)) C. lupus DPSs: —WGL DPS —NRM DPS 2008 Reinstatement of protections—NRM & WGL DPSs 73 FR 75356, December 11, 2008 C. lupus WGL DPS 2009 Designate DPS & delist 74 FR 15070, April 2, 2009 Rule vacated ( Humane Society of the United States v. Salazar, 1:09-CV-1092-PLF (D.D.C. 2009)) C. lupus NRM DPS (except Wyoming) 2009 Designate DPS & delist (except in Wyoming) 74 FR 15123, April 2, 2009 Rule vacated ( Defenders of Wildlife v. Salazar, 729 F. Supp. 2d 1207 (D. Mont. 2010)) C. lupus WGL DPS 2009 Reinstatement of protections—WGL 74 FR 47483, September 16, 2009 C. lupus NRM DPS 2010 Reinstatement of protections—NRM DPS 75 FR 65574, October 26, 2010 C. lupus NRM DPS 2011 Reissuance of 2009 NRM DPS delisting rule (as required by Public Law 112-10-The Department of Defense and Full-Year Continuing Appropriations Act, 2011) 76 FR 25590, May 5, 2011 C. lupus WGL DPS 2011 Revise 1978 listing, designate DPS & delist 76 FR 81666, December 28, 2011 Rule vacated ( Humane Society of the U.S. v. Jewell, 76 F. Supp. 3d 69, 110 (D.D.C. 2014)) Vacatur upheld on appeal ( Humane Society of the U.S. v. Zinke, 865 F.3d 585 (D.C. Cir. 2017)) C. lupus in Wyoming 2012 Delist in Wyoming 77 FR 55530, September 10, 2012 Rule vacated ( Defenders of Wildlife v. Jewell, 68 F. Supp. 3d 193 (D.D.C. 2014) Vacatur reversed on appeal ( Defenders of Wildlife v. Zinke, 849 F.3d 1077 (D.C. Cir. 2017)) C. lupus in lower 48 U.S
Supp. 3d 69, 110 (D.D.C. 2014)) Vacatur upheld on appeal ( Humane Society of the U.S. v. Zinke, 865 F.3d 585 (D.C. Cir. 2017)) C. lupus in Wyoming 2012 Delist in Wyoming 77 FR 55530, September 10, 2012 Rule vacated ( Defenders of Wildlife v. Jewell, 68 F. Supp. 3d 193 (D.D.C. 2014) Vacatur reversed on appeal ( Defenders of Wildlife v. Zinke, 849 F.3d 1077 (D.C. Cir. 2017)) C. lupus in lower 48 U.S. (except NRM & WGL DPSs) and Mexico 2013 Propose delist in lower 48 U.S. & list C. l. baileyi (E); status review of wolves in Pacific Northwest 78 FR 35664, June 13, 2013 C. l. baileyi 2015 List E 80 FR 2488, January 16, 2015 C. l. baileyi 2015 Revised 1998 C. lupus experimental population and associated it with C. l. baileyi listing 80 FR 2512, January 16, 2015 C. lupus WGL DPS and C. lupus in Wyoming 2015 Reinstatement of protections—WGL DPS & Wyoming 80 FR 9218, February 20, 2015 C. lupus in Wyoming 2017 Reinstatement of 2012 delisting—Wyoming 82 FR 20284, May 1, 2017 1 Action taken under the Endangered Species Preservation predecessor legislation (Endangered Species Act of 1966, Endangered Species Conservation Act of 1969). 2 Later subsumed into C. l. baileyi due to taxonomic changes. 3 In this rule we also identified critical habitat in Michigan and Minnesota and promulgated special regulations under section 4(d) of the Act for operating a wolf- management program in Minnesota. The special regulation was later modified (50 FR 50793, December 12, 1985). BILLING CODE 4333-15-P EP15MR19.005
BILLING CODE 4333-15-C General Background
The 1978 Reclassification
When the gray wolf ( C. lupus ) was reclassified in March 1978 (replacing multiple subspecies listings with two C. lupus population listings as described further in Previous Federal Actions), it had been extirpated from much of its historical range in the contiguous United States
2, 1985). BILLING CODE 4333-15-P EP15MR19.005
BILLING CODE 4333-15-C General Background
The 1978 Reclassification
When the gray wolf ( C. lupus ) was reclassified in March 1978 (replacing multiple subspecies listings with two C. lupus population listings as described further in Previous Federal Actions), it had been extirpated from much of its historical range in the contiguous United States. Although the 1978 reclassification listed two gray wolf entities (a threatened population in Minnesota and an endangered population throughout the rest of the contiguous United States and Mexico), these listings were not predicated upon a formal DPS analysis, because the reclassification predated the November 1978 amendments to the Act, which revised the definition of “species” to include distinct population segments of vertebrate fish or wildlife, and our 1996 DPS Policy.
As indicated in Previous Federal Actions, the 1978 reclassification was employed as an approach of convenience to ensure the gray wolf was protected wherever it was found (as described in 47 FR 9607, March 9, 1978) in the lower 48 States and Mexico, rather than an indication of where gray wolves actually existed or where gray wolf recovery would occur. Thus, the 1978 reclassification resulted in inclusion of large areas of the contiguous United States where gray wolves were extirpated, as well as the mid-Atlantic and southeastern United States—west to central Texas and Oklahoma—an area that is generally accepted not to be within the historical range of C. lupus (Young and Goldman 1944, pp. 413-416, 478; Nowak 1995, p. 395, fig. 20). While this generalized approach to the listing appropriately protected dispersing wolves throughout the historical range of C
olves were extirpated, as well as the mid-Atlantic and southeastern United States—west to central Texas and Oklahoma—an area that is generally accepted not to be within the historical range of C. lupus (Young and Goldman 1944, pp. 413-416, 478; Nowak 1995, p. 395, fig. 20). While this generalized approach to the listing appropriately protected dispersing wolves throughout the historical range of C. lupus in the United States and Mexico and facilitated recovery of the northern Rocky Mountains and western Great Lakes populations, it also erroneously included areas outside the species' historical range and was misread by some members of the public as an expression of a larger gray wolf recovery effort not required by the Act and never intended by the Service. In fact, as discussed below (see National Wolf Strategy ), our recovery efforts have consistently focused on reestablishing wolf populations in specific areas of the country.
National Wolf Strategy
We first described our national wolf strategy in our May 5, 2011, proposed rule to revise the List for the gray wolf in the eastern United States (76 FR 26086). This strategy was intended to: (1) Lay out a cohesive and coherent approach to addressing wolf conservation needs, including protection and management, in accordance with the Act's statutory framework; (2) ensure that actions taken for one wolf population do not cause unintended consequences for other populations; and (3) be explicit about the role of historical range in the conservation of extant wolf populations. Included in this strategy is the precept that, in order to qualify for any type of listing or delisting action, wolf entities must conform to the Act's definition of “species,” whether as taxonomic species or subspecies or as distinct population segments.
Our May 5, 2011, proposed rule states that our strategy focuses on conservation of four extant gray wolf entities being considered for C. l. baileyi ) separately as endangered in 2015
that, in order to qualify for any type of listing or delisting action, wolf entities must conform to the Act's definition of “species,” whether as taxonomic species or subspecies or as distinct population segments.
Our May 5, 2011, proposed rule states that our strategy focuses on conservation of four extant gray wolf entities being considered for C. l. baileyi ) separately as endangered in 2015. However, as indicated in Previous Federal Actions, our 2011 final rule designating and delisting the WGL DPS was subsequently vacated.
In addition to the rules described above, we completed a status review for gray wolves in the Pacific Northwest (western Washington and western Oregon) in 2013 (table 1). We determined that these wolves are not discrete, under our DPS policy, from wolves in the NRM DPS (see 78 FR 35707-35713) and, therefore, are not a valid listable entity under the Act. Wolves in the Pacific Northwest are a mix of individuals derived from wolves in the northern Rocky Mountains and Canada (or both) and represent the expanding fronts of these populations (78 FR 35707-35713, USFWS 2018, pp. 4, 14-15, 23). Since publication of our 2013 status review, wolves have also expanded into northern California. Wolves in northern California are not discrete from those in the Pacific Northwest based on documented movement of wolves between Oregon and California (USFWS 2018, pp. 14-15). Therefore, wolves in western Washington, western Oregon, and northern California are not a valid DPS because they are not discrete from the NRM DPS.
Approach for This Proposed Rule
The Entities Addressed in This Rule
In this proposed rule, we consider the status of the gray wolf within the geographic boundaries of the two currently listed C. lupus entities to determine whether these wolves should remain on the List in their current status, be reclassified, or be removed from the List. These two currently listed entities are: (1) C. lupus in Minnesota, and (2) C
osed Rule
The Entities Addressed in This Rule
In this proposed rule, we consider the status of the gray wolf within the geographic boundaries of the two currently listed C. lupus entities to determine whether these wolves should remain on the List in their current status, be reclassified, or be removed from the List. These two currently listed entities are: (1) C. lupus in Minnesota, and (2) C. lupus in the lower 48 United States and Mexico outside of Minnesota, the NRM DPS (Montana, Idaho, Wyoming, eastern third of Washington and Oregon, and north-central Utah), and the area covered by the experimental population area for C. l. baileyi (the designated area in which the subspecies is being re-introduced; see 63 FR 1752, January 12, 1998). These two entities are currently listed as threatened and endangered, respectively.
While our past status reviews have focused on C. lupus DPSs and taxonomic units that align with our national wolf strategy (see table 1), this status review considers the current C. lupus listed entities described above. We do this:
(1) To address the Court of Appeals concerns with our 2011 final rule delisting the WGL DPS, specifically, concern pertaining to the impacts of partial delisting on the remainder of the already-listed species (see Previous Federal Actions);
(2) To avoid a rulemaking that conflicts with multiple court opinions regarding our prior attempts to designate and delist wolf DPSs (see table 1); and
s:
(1) To address the Court of Appeals concerns with our 2011 final rule delisting the WGL DPS, specifically, concern pertaining to the impacts of partial delisting on the remainder of the already-listed species (see Previous Federal Actions);
(2) To avoid a rulemaking that conflicts with multiple court opinions regarding our prior attempts to designate and delist wolf DPSs (see table 1); and
(3) Because, with the exception of C. l. baileyi, which is listed separately as endangered wherever found (see Previous Federal Actions), the taxonomy of C. lupus is complex, controversial, and unresolved (USFWS 2018, pp. 1-4; also see How We Address Taxonomic Uncertainties in this Rule, below).
How We Address the C. lupus Entities in This Rule
The two currently listed gray wolf entities are vestiges of a 40-year-old action (the 1978 reclassification (see Background)). Our knowledge of wolf biology and taxonomy has vastly changed since then. Additionally, our previous efforts to revise the listed entities have not withstood judicial scrutiny (see Previous Federal Actions). Our policies and practices pertaining to listable entities have also changed since the 1978 reclassification. As a result, these entities do not conform with our current policies and standard practice. Specifically: (1) These two entities are not discrete from one another under our current policy on vertebrate distinct population segments (DPSs) (61 FR 4722, February 7, 1996); (2) the listing for the larger entity includes areas known to overlap with the range of the separately listed gray wolf subspecies C. l. baileyi; and (3) wolves currently listed in the western United States are not discrete from the recovered Northern Rocky Mountains population, which we removed from the List in 2009 (table 1).
e distinct population segments (DPSs) (61 FR 4722, February 7, 1996); (2) the listing for the larger entity includes areas known to overlap with the range of the separately listed gray wolf subspecies C. l. baileyi; and (3) wolves currently listed in the western United States are not discrete from the recovered Northern Rocky Mountains population, which we removed from the List in 2009 (table 1).
(1) Lack of Discreteness of the Two C. lupus Listed Entities
Under the Act we can list a species, subspecies, or vertebrate DPS. Neither of the two entities currently on the List represents an entire species or subspecies, thus to comply with the statute, these listings must be DPSs. Our 1996 DPS policy specifies that a vertebrate population must be both discrete and significant to qualify as a DPS (61 FR 4722-4725; February 7, 1996). To qualify as “discrete,” a population must be “markedly separated from other populations of the same taxon as a consequence of physical, physiological, ecological, or behavioral factors” (61 FR 4725). However, as indicated, the populations in these two entities are no longer discrete (U.S. Fish and Wildlife Service (USFWS) 2018, pp. 22-23). Therefore, because it is clear that neither entity would qualify as a DPS under our 1996 DPS policy (61 FR 4725), we consider the conservation status of the two listed wolf entities as one combined entity in this proposed rule. We refer to the combined entity simply as “the gray wolf entity” throughout this proposed rule.
o longer discrete (U.S. Fish and Wildlife Service (USFWS) 2018, pp. 22-23). Therefore, because it is clear that neither entity would qualify as a DPS under our 1996 DPS policy (61 FR 4725), we consider the conservation status of the two listed wolf entities as one combined entity in this proposed rule. We refer to the combined entity simply as “the gray wolf entity” throughout this proposed rule.
(2) C. l. baileyi listing
As indicated above (see Previous Federal Actions), in 2015 we revised the listing for gray wolf by reclassifying the subspecies C. l. baileyi as a separately listed entity with the status of endangered, wherever found. Although the rulemaking does not include language expressly excluding C. l. baileyi from the previously listed C. lupus entity, we indicated in our 2015 final rule listing the subspecies that the effect of the regulation was to revise the List by making a separate entry for the Mexican wolf (80 FR 2488, 2511, January 16, 2015). Therefore, because we already assessed the status of, and listed, the Mexican wolf separately, we do not consider individuals or populations of C. l. baileyi in this proposed rule. In geographical terms, we do not consider wolves occurring in Mexico and within the experimental population area in this proposed rule. Canis lupus baileyi is the only subspecies known to occur in these areas, and we have no information suggesting that other gray wolves occur in these areas.
can wolf separately, we do not consider individuals or populations of C. l. baileyi in this proposed rule. In geographical terms, we do not consider wolves occurring in Mexico and within the experimental population area in this proposed rule. Canis lupus baileyi is the only subspecies known to occur in these areas, and we have no information suggesting that other gray wolves occur in these areas.
(3) Lack of Discreteness of Western Wolves Within and Outside the Gray Wolf Entity
In the coastal States of the western United States, wolves within the gray wolf entity occur in an area comprising western Oregon, western Washington, and northern California. These wolves are part of the expanding fronts (or edges) of the recovered and delisted wolf population in the NRM DPS and wolves crossing into the United States
How We Address Taxonomic Uncertainties in This Rule
The taxonomy and evolutionary history of wolves in North America are complex and controversial, particularly with respect to the taxonomic assignment of wolves in the northeastern United States and portions of the Great Lakes region (eastern wolves) (see Taxonomy of Gray Wolves in North America ). Available information indicates ongoing scientific debate and a lack of resolution on the taxonomy of eastern wolves. Some scientists consider eastern wolves to be a distinct species, C. lycaon; some consider them gray wolves ( C. lupus ); and some consider them the product of hybridization between gray wolves and coyotes (USFWS 2018, p. 1). Further, none of these viewpoints is more widely accepted by the scientific community.
For the purposes of this proposed rule, we consider eastern wolves to be members of the species C. lupus because there is not clear support for a recognizable and independent evolved eastern wolf species. Therefore, in our assessment of the status of the gray wolf entity, we include eastern wolves and eastern wolf range that occurs within the geographical boundaries of the gray wolf entity
nity.
For the purposes of this proposed rule, we consider eastern wolves to be members of the species C. lupus because there is not clear support for a recognizable and independent evolved eastern wolf species. Therefore, in our assessment of the status of the gray wolf entity, we include eastern wolves and eastern wolf range that occurs within the geographical boundaries of the gray wolf entity.
We note that in our 2013 proposed rule to delist wolves in the lower 48 United States and Mexico (table 1), we accepted the conclusions of Chambers et al. (2012, entire) on the taxonomy of eastern wolves and recognized eastern wolves as the distinct species C. lycaon. However, peer reviewers of our 2013 proposed rule indicated that Chambers et al. was not universally accepted and our rule did not represent the best available science (National Center for Ecological Analysis and Synthesis 2014, entire). Also, new information published on the topic since publication of our 2013 rule indicates the taxonomy of eastern wolves continues to be controversial and unresolved (USFWS 2018, pp. 1-2). Finally, the uncertainty of the existence of a separate species is reflected in the fact that C. lycaon is not recognized by authoritative taxonomic organizations such as the American Society of Mammalogists or the International Commission on Zoological Nomenclature.
Scientists also disagree on the taxonomic assignment of wolves in the southeastern United States generally recognized as “red wolves.” However, we recognize the red wolf as the species C. rufus, and note that it is listed as endangered where found (32 FR 4001, March 11, 1967). We do not consider red wolves further in this rule, and the red wolf listing is not affected by this proposal.
Summary of Our Approach
In this proposed rule, we assess the status of gray wolves occurring within the geographic area outlined by the two currently listed gray wolf ( C
d wolf as the species C. rufus, and note that it is listed as endangered where found (32 FR 4001, March 11, 1967). We do not consider red wolves further in this rule, and the red wolf listing is not affected by this proposal.
Summary of Our Approach
In this proposed rule, we assess the status of gray wolves occurring within the geographic area outlined by the two currently listed gray wolf ( C. lupus ) entities combined (figure 1), but we do not include in our assessment individuals or populations of the Mexican gray wolf ( C. l. baileyi ) (wolves that occur in Mexico and the nonessential experimental population area in the southwestern United States) as these wolves are separately listed as an endangered subspecies (80 FR 2488, January 16, 2015). Further, for the purposes of this proposed rule, we consider any eastern wolves within the geographic boundaries of the two currently listed gray wolf entities to be members of the species C. lupus. As stated previously, this proposed rule supersedes the June 13, 2013, proposed rule to delist C. lupus in the remaining listed portions of the United States and Mexico outside of the delisted NRM and WGL (78 FR 35663).
Species Information
We provide detailed background information on gray wolves in the United States in a separate Gray Wolf Biological Report (see USFWS 2018, entire). This document can be found along with this proposed rule at http://regulations.gov in Docket No. FWS-HW-ES-2018-0097 (see Supplemental Documents ). We summarize relevant information from this report below. For additional information, including sources of the information presented below, see USFWS (2018, entire) and references therein.
Biology and Ecology
Gray wolves are the largest wild members of the dog family and have a broad circumpolar range. They are highly territorial, social animals that live and hunt in packs. They are well adapted to traveling fast and far in search of food, and catching and eating large mammals
including sources of the information presented below, see USFWS (2018, entire) and references therein.
Biology and Ecology
Gray wolves are the largest wild members of the dog family and have a broad circumpolar range. They are highly territorial, social animals that live and hunt in packs. They are well adapted to traveling fast and far in search of food, and catching and eating large mammals. In North America they are primarily predators of medium to large mammals, including deer, elk, and other species.
Gray wolves are habitat generalists. They can successfully occupy a wide range of habitats and are not dependent on wilderness for their survival. An inadequate prey density and a high level of human persecution appear to be the only factors that limit habitat suitability and gray wolf distribution. Thus, virtually any area that has sufficient prey and adequate protection from persecution can be suitable habitat for gray wolves.
Wolf populations are remarkably resilient as long as food supply and regulation of human-caused mortality are adequate. In the absence of high levels of anthropogenic influences, wolf populations are generally believed to be regulated by the distribution and abundance of prey on the landscape, though density-dependent, intrinsic mechanisms ( e.g., social strife, territoriality, disease) may limit populations when ungulate densities are high. Where harvest occurs, high levels of reproduction and immigration can compensate for high mortality rates. Pack social structure is very adaptable—breeding members can be quickly replaced from within or outside the pack, and pups can be reared by another pack member should their parents die. Consequently, wolf populations can rapidly overcome severe disruptions, such as pervasive human-caused mortality or disease. Wolf populations can increase rapidly after severe declines if the source of mortality is reduced
ial structure is very adaptable—breeding members can be quickly replaced from within or outside the pack, and pups can be reared by another pack member should their parents die. Consequently, wolf populations can rapidly overcome severe disruptions, such as pervasive human-caused mortality or disease. Wolf populations can increase rapidly after severe declines if the source of mortality is reduced. Also, the species' dispersal capabilities allow a wolf population to quickly expand and colonize nearby areas, even areas separated by broad expanses of unsuitable habitat.
Taxonomy of Gray Wolves in North America
The taxonomy of the genus Canis in North America has a complex and contentious history, particularly with respect to two generally recognized phenotypes (morphological forms) that occur in eastern North America: The “red wolf” and “eastern wolf.” As indicated above (see How We Address Taxonomic Uncertainties in this Rule ), we continue to recognize the red wolf as the species C. rufus and do not discuss the taxonomy of the species further in this rule (for more information, see our 2018 Red Wolf Species Status Assessment). We discuss the eastern wolf further below.
The “eastern wolf” has been the source of perhaps the most significant disagreement on North American canid taxonomy among scientists. The “eastern wolf” has been variously described as a species, a subspecies of gray wolf, an ecotype of gray wolf, or the product of hybridization between gray wolves and coyotes. Hybridization is widely recognized to have played, and to continue to play, an important role among “eastern wolves,” with varying views on the role of hybridization between “eastern wolves” and coyotes, “eastern wolves” and gray wolves, and gray wolves and coyotes
bed as a species, a subspecies of gray wolf, an ecotype of gray wolf, or the product of hybridization between gray wolves and coyotes. Hybridization is widely recognized to have played, and to continue to play, an important role among “eastern wolves,” with varying views on the role of hybridization between “eastern wolves” and coyotes, “eastern wolves” and gray wolves, and gray wolves and coyotes. Minnesota appears to be the western edge of a hybrid zone between western gray wolves and eastern wolves—wolves in western Minnesota appear to be gray wolves both morphologically and genetically while wolves in eastern Minnesota and much of the Great Lakes area appear to be “eastern wolf,” introgressed with western gray wolf to varying degrees.
No controversy exists regarding the number of wolf species in western North America—all are widely recognized as gray wolves ( C. lupus ). However, the science pertaining to gray wolf subspecies designations, unique evolutionary lineages, ecotypes, and admixture of formerly isolated populations continues to develop and remains unresolved. Even so, genetic studies indicate that wolves in Washington include individuals from the northern Rocky Mountains, individuals from British Columbia, and individuals of mixed ancestry. Wolves currently occupying Oregon and California are derived from dispersers from the northern Rocky Mountains.
Range and Population Trends Prior to 1978 Reclassification
Historical Range of the Gray Wolf Entity
We view the historical range to be the range of gray wolves within the gray wolf entity at the time of European settlement. We determined that this timeframe is appropriate because it precedes the major changes in range in response to excessive human-caused mortality (USFWS 2018, pp. 7-11).
At the time of the 1978 reclassification, the historical range of the gray wolf was generally believed to include most of North America and, consequently, most of the gray wolf entity
olf entity at the time of European settlement. We determined that this timeframe is appropriate because it precedes the major changes in range in response to excessive human-caused mortality (USFWS 2018, pp. 7-11).
At the time of the 1978 reclassification, the historical range of the gray wolf was generally believed to include most of North America and, consequently, most of the gray wolf entity. In the lower 48 United States, they were reportedly absent from parts of California, the arid deserts and mountaintops of the western United States, and parts of the eastern United States. However, some authorities question the species' historical absence in parts of California. In addition, long-held differences of opinion exist among scientists regarding the precise boundary of the gray wolf's historical range in the eastern United States. Some believe the range of gray wolves extended as far south as southern Georgia while others believe it did not extend into the southeast at all. The southeastern and mid-Atlantic States are generally recognized as being within the historical range of the red wolf, but it is not known how much range overlap historically occurred between these two species. Because of the various scientific positions on gray wolf species and range, the historical extent of gray wolf range for much of the gray wolf entity in the eastern United States remains uncertain.
Based on our review of the best available information, we view the historical range of the gray wolf within the gray wolf entity to follow that presented in Nowak (1995) and depicted in figure 2. This includes all areas within the gray wolf entity except western California, a small portion of southwestern Arizona, and the southeastern United States (see figure 2 and USFWS 2018, pp. 7-11).
While some authorities question the absence of gray wolves in parts of California, limited preserved physical evidence of wolves in California exists
presented in Nowak (1995) and depicted in figure 2. This includes all areas within the gray wolf entity except western California, a small portion of southwestern Arizona, and the southeastern United States (see figure 2 and USFWS 2018, pp. 7-11).
While some authorities question the absence of gray wolves in parts of California, limited preserved physical evidence of wolves in California exists. Therefore, we rely on early reports of wolves in the State that describe the species as occurring in the northern and Sierra Mountain regions of California. Further, while recognizing that the extent of overlap of C. rufus and C. lupus ranges is unknown, because the southeastern United States are generally recognized as within the range of C. rufus, we consider it to be generally outside the range of C. lupus. However, we acknowledge that the historical range of C. lupus is uncertain and the topic of continued debate among scientists.
Historical Abundance of the Gray Wolf Entity
Historical abundance of gray wolves within the gray wolf entity is largely unknown. Based on the reports of European settlers, gray wolves were common in much of the West. While historical (at the time of European settlement) estimates are notoriously difficult to verify, one study estimates that hundreds of thousands of wolves occurred in the western United States and Mexico. In the Great Lakes area, there were an estimated 4,000 to 8,000 in Minnesota, 3,000 to 5,000 in Wisconsin, and fewer than 6,000 in Michigan. No estimates are available for historical abundance in the Northeast.
Historical Trends in Range and Abundance for the Gray Wolf Entity
Gray wolf range and numbers throughout the gray wolf entity declined significantly during the 19th and 20th centuries as a result of killing of wolves by humans through poisoning, unregulated trapping and shooting, and government-funded wolf-extermination efforts
estimates are available for historical abundance in the Northeast.
Historical Trends in Range and Abundance for the Gray Wolf Entity
Gray wolf range and numbers throughout the gray wolf entity declined significantly during the 19th and 20th centuries as a result of killing of wolves by humans through poisoning, unregulated trapping and shooting, and government-funded wolf-extermination efforts. By the time subspecies were first listed under the Act in 1974 (table 1), the gray wolf had been eliminated from most of its historical range within the lower 48 United States, including within most of the gray wolf entity.
Distribution, and Abundance of the Gray Wolf Entity at the Time of the 1978 Reclassification
By the time gray wolf subspecies were listed under the Act in 1974 (table 1), the species occurred in only a small fraction of its historical range. Aside from a few scattered individuals, wolves occurred in only two places within the gray wolf entity (and the entire lower 48 United States). A population persisted in northeastern Minnesota, and a small, isolated group of about 40 wolves occurred on Isle Royale, Michigan. The Minnesota wolf population was the only major U.S. population in existence outside Alaska at this time and numbered about 1,000 individuals. While the Minnesota population was small compared to historical numbers and range within the lower 48 United States, it had not undergone a significant decline since about 1900. By 1978, when several gray wolf subspecies were consolidated into a single lower 48 United States/Mexico listing and a separate Minnesota listing under the Act, the gray wolf population in Minnesota had increased to an estimated 1,235 wolves in 138 packs (in the winter of 1978-79) and had an estimated range of 14,038 square miles (mi 2 ) (36,500 square kilometers (km 2 )) (figure 2). Although it was suspected that wolves inhabited Wisconsin at this time, it was not until 1979 that wolf presence was confirmed in the State
e Minnesota listing under the Act, the gray wolf population in Minnesota had increased to an estimated 1,235 wolves in 138 packs (in the winter of 1978-79) and had an estimated range of 14,038 square miles (mi 2 ) (36,500 square kilometers (km 2 )) (figure 2). Although it was suspected that wolves inhabited Wisconsin at this time, it was not until 1979 that wolf presence was confirmed in the State.
Current Distribution and Abundance of the Gray Wolf Entity
The vast majority of wolves within the gray wolf entity now exist as a large, stable or growing metapopulation (partially isolated set of subpopulations) of more than 4,400 individuals that is broadly distributed across the northern portions of three States in the Great Lakes area. This metapopulation is also connected, via documented dispersals, to the large and expansive population of about 12,000-14,000 wolves in eastern Canada. As a result, gray wolves in the
In addition to the metapopulation in the Great Lakes area, as of 2017, three breeding pairs and four packs with no documented reproduction occur within the gray wolf entity in Oregon, Washington, and California. These wolves originated from large populations of approximately 15,000 wolves in western Canada and about 1,700 wolves in the northern Rocky Mountains. Effective dispersal has been documented among California, Oregon, and Washington as well as between these States and other northern Rocky Mountains States and Canada. Thus, wolves in the Pacific coast States are an extension of the metapopulation of wolves in western Canada and the northern Rocky Mountains.
Finally, a number of lone long-distance dispersing wolves have been documented outside core populations of the Great Lakes area and western United States since the early 2000s. Confirmed records of individual wolves have been reported from North Dakota, South Dakota, Utah, Colorado, Nevada, Missouri, Indiana, Illinois, Nebraska, and Kansas
lves in western Canada and the northern Rocky Mountains.
Finally, a number of lone long-distance dispersing wolves have been documented outside core populations of the Great Lakes area and western United States since the early 2000s. Confirmed records of individual wolves have been reported from North Dakota, South Dakota, Utah, Colorado, Nevada, Missouri, Indiana, Illinois, Nebraska, and Kansas. The total number of confirmed records in each of these States, since the early 2000s, ranges from one in Nevada to at least 27 in North Dakota, with the latter also having an additional 45 probable but unverified reports.
BILLING CODE 4333-15-P EP15MR19.006
BILLING CODE 4333-15-C Gray Wolf Recovery Plans and Recovery Implementation
Section 4(f) of the Act directs us to develop and implement recovery plans for the conservation and survival of endangered and threatened species unless we determine that such a plan will not promote the conservation of the species. Recovery plans are non-regulatory documents that identify site-specific management actions that may be necessary to achieve conservation and survival of the species. They also identify objective, measurable criteria (recovery criteria) which, when met, would result in a determination that the species should be removed from the List. Methods for monitoring recovery progress may also be included in recovery plans.
The Act does not describe recovery in terms of the proportion of historical range that must be occupied by a species, nor does it ever allude to restoration throughout the entire historical range as a conservation purpose. In fact, the Act itself does not contain the phrase “historical range.” Thus, the Act does not require us to restore the gray wolf (or any other species) to all of its historical range or any specific percentage of currently suitable habitat
historical range that must be occupied by a species, nor does it ever allude to restoration throughout the entire historical range as a conservation purpose. In fact, the Act itself does not contain the phrase “historical range.” Thus, the Act does not require us to restore the gray wolf (or any other species) to all of its historical range or any specific percentage of currently suitable habitat. For some species, expansion of their distribution or abundance may be necessary to achieve recovery, but the amount of expansion is driven by a species' biological needs affecting viability (ability to sustain
As indicated in Previous Federal Actions, following our 1978 reclassification, we drafted recovery plans and implemented recovery programs for gray wolves in three regions of the contiguous United States (table 1). Wolves in one of these regions— C. l. baileyi, in the southwestern United States and Mexico—were recently listed separately as an endangered subspecies and are not considered in this rule (see Approach for this Proposed Rule). Wolves in another of these regions—the northern Rocky Mountains—have recovered and were delisted (table 1). We discuss recovery of wolves in the third region—the eastern United States—as it relates to the status of the gray wolf entity, below. We did not develop a recovery plan for wolves in the U.S. west coast States because we did not identify this area as necessary to the recovery of the species following our 1978 reclassification. We have not since developed a recovery plan for these wolves because we determined in our 2013 status review that they are biologically part of (although outside the legal boundary of) an already recovered and delisted population (see National Wolf Strategy ).
Recovery Criteria
There are many paths to accomplish recovery of a species, and recovery may be achieved without all recovery criteria being fully met
ce developed a recovery plan for these wolves because we determined in our 2013 status review that they are biologically part of (although outside the legal boundary of) an already recovered and delisted population (see National Wolf Strategy ).
Recovery Criteria
There are many paths to accomplish recovery of a species, and recovery may be achieved without all recovery criteria being fully met. We use recovery criteria in concert with evidence that threats have been minimized sufficiently and populations have achieved long-term viability to determine when a species can be reclassified from endangered to threatened or delisted. Recovery of a species is a dynamic process requiring adaptive management that may, or may not, fully follow the guidance provided in a recovery plan. Recovery plans, including recovery criteria, are subject to change based upon new information and are revised accordingly and when practicable. In a similar sense, implementation of planned actions is subject to changing information and availability of resources. We have taken these considerations into account in the following discussion.
The 1978 Recovery Plan (hereafter Recovery Plan) and the 1992 Revised Recovery Plan for the Eastern Timber Wolf (hereafter Revised Recovery Plan) were developed to guide recovery of the eastern timber wolf subspecies. Those recovery plans contain the same two recovery criteria, which are meant to indicate when recovery of the eastern timber wolf throughout its historical range in the eastern United States has been achieved. The first recovery criterion states that the survival of the wolf in Minnesota must be assured. We, and the Eastern Timber Wolf Recovery Team (Peterson in litt. 1997, 1998, 1999a, 1999b), have concluded that this recovery criterion remains valid. It addresses a need for reasonable assurances that future State, tribal, and Federal wolf management and protection will maintain a viable recovered population of wolves within the borders of Minnesota for the foreseeable future
ota must be assured. We, and the Eastern Timber Wolf Recovery Team (Peterson in litt. 1997, 1998, 1999a, 1999b), have concluded that this recovery criterion remains valid. It addresses a need for reasonable assurances that future State, tribal, and Federal wolf management and protection will maintain a viable recovered population of wolves within the borders of Minnesota for the foreseeable future.
Although the recovery criteria identified in the Recovery Plan predate identification of the conservation biology principles of representation (conserving the adaptive genetic diversity of a taxon), resiliency (ability to withstand demographic and environmental variation), and redundancy (sufficient populations to provide a margin of safety), those principles were incorporated into the recovery criteria. The Recovery Team insisted that the remnant Minnesota wolf population be maintained and protected to achieve wolf recovery in the eastern United States. Maintenance of the Minnesota wolf population is vital in terms of representation because these wolves include both western gray wolves and wolves that are admixtures of western gray wolves and eastern wolves. In other words, they contain the genetic components of both western gray wolves and eastern wolves. The successful growth of the remnant Minnesota population has maintained and maximized the representation of that genetic diversity among wolves in the Great Lakes area.
Maintenance of the Minnesota wolf population is also vital in terms of resiliency. Although the Revised Recovery Plan did not establish a specific numerical criterion for the Minnesota wolf population, it did identify, for planning purposes only, a population goal of 1,251-1,400 animals for that Minnesota population (USFWS 1992, p. 28). A population of this size not only increases the likelihood of maintaining its genetic diversity over the long term, but also reduces the adverse impacts of unpredictable demographic and environmental events
numerical criterion for the Minnesota wolf population, it did identify, for planning purposes only, a population goal of 1,251-1,400 animals for that Minnesota population (USFWS 1992, p. 28). A population of this size not only increases the likelihood of maintaining its genetic diversity over the long term, but also reduces the adverse impacts of unpredictable demographic and environmental events. Furthermore, the Revised Recovery Plan recommends a wolf population that is spread across about 40 percent of Minnesota (Zones 1 through 4) (USFWS 1992, p. 28), adding a geographic component to the resiliency of the Minnesota wolf population.
The second recovery criterion in the Recovery Plan states that at least one viable wolf population should be reestablished within the historical range of the eastern timber wolf outside of Minnesota and Isle Royale, Michigan (USFWS 1992, pp. 24-26). The reestablished population enhances both the resiliency and redundancy of the Great Lakes metapopulation.
The Recovery Plan provides two options for reestablishing this second population. If it is an isolated population, that is, located more than 100 miles (mi) (160 kilometers (km)) from the Minnesota wolf population, the second population should consist of at least 200 wolves for at least 5 years, based upon late-winter population estimates, to be considered viable. Late-winter estimates are made at a time when most winter mortality has already occurred and before the birth of pups, thus, the count is made at the annual low point of the population. Alternatively, if the second population is located within 100 mi (160 km) of a self-sustaining wolf population (for example, the Minnesota wolf population), it should be maintained at a minimum of 100 wolves for at least 5 years, based on late-winter population estimates, to be considered viable
curred and before the birth of pups, thus, the count is made at the annual low point of the population. Alternatively, if the second population is located within 100 mi (160 km) of a self-sustaining wolf population (for example, the Minnesota wolf population), it should be maintained at a minimum of 100 wolves for at least 5 years, based on late-winter population estimates, to be considered viable. A nearby second population would be considered viable at a smaller size because it would be geographically close enough to exchange wolves with the Minnesota population (that is, they would function as a metapopulation), thereby bolstering the smaller second population both genetically and numerically.
The original Recovery Plan did not specify where in the eastern United States the second population should be reestablished. Therefore, the second population could have been established anywhere within the triangular Minnesota-Maine-Florida area covered by the Recovery Plan and the Revised Recovery Plan, except on Isle Royale (Michigan) or within Minnesota. The Revised Recovery Plan identified potential gray wolf reestablishment areas in northern Wisconsin, the Upper Peninsula of Michigan, the Adirondack Forest Preserve of New York, a small area in eastern Maine, and a larger area of northwestern Maine and adjacent northern New Hampshire (USFWS
In 1998, the Eastern Timber Wolf Recovery Team clarified the application of the recovery criterion for the second population to the wolf population that had developed in northern Wisconsin and the adjacent Upper Peninsula of Michigan. This second population is less than 100 mi (160 km) from the Minnesota wolf population. The Recovery Team recommended that the numerical recovery criterion for the Wisconsin-Michigan population be considered met when consecutive late-winter wolf surveys document that the population equals or exceeds 100 wolves (excluding Isle Royale wolves) for the 5 consecutive years between the first and last surveys (Peterson in litt. 1998)
than 100 mi (160 km) from the Minnesota wolf population. The Recovery Team recommended that the numerical recovery criterion for the Wisconsin-Michigan population be considered met when consecutive late-winter wolf surveys document that the population equals or exceeds 100 wolves (excluding Isle Royale wolves) for the 5 consecutive years between the first and last surveys (Peterson in litt. 1998).
Recovery Progress
Wolves in the Great Lakes area greatly exceed the recovery criteria (USFWS 1992, pp. 24-26) for (1) a secure wolf population in Minnesota, and (2) a second population outside Minnesota and Isle Royale consisting of 100 wolves for 5 successive years. Based on the eight surveys conducted since 1998, the wolf population in Minnesota has exceeded 2,000 individuals over the past 20 years, and populations in Michigan and Wisconsin have exceeded 100 individuals every year since 1996 (USFWS 2018, appendix 1). Based on the criteria set by the Eastern Wolf Recovery Team in 1992 and reaffirmed in 1997 and 1998 (Peterson in litt. 1997, in litt. 1998), this region contains sufficient wolf numbers and distribution to ensure the long-term survival of the gray wolf entity.
The maintenance and expansion of the Minnesota wolf population has allowed for the preservation of the genetic diversity that remained in the Great Lakes area when its wolves were first protected in 1974. Furthermore, the Wisconsin-Michigan wolf population far exceeds the numerical recovery criterion even for a completely isolated second population. Therefore, even in the unlikely event that this two-State population were to become totally isolated and wolf immigration from Minnesota and Ontario completely ceased, it would still remain a viable wolf population for the foreseeable future, as defined by the Revised Recovery Plan (USFWS 1992, pp. 25-26)
ar exceeds the numerical recovery criterion even for a completely isolated second population. Therefore, even in the unlikely event that this two-State population were to become totally isolated and wolf immigration from Minnesota and Ontario completely ceased, it would still remain a viable wolf population for the foreseeable future, as defined by the Revised Recovery Plan (USFWS 1992, pp. 25-26). Finally, each of the wolf populations in Wisconsin and Michigan has exceeded 200 animals for about 20 years, so if either were somehow to become isolated, they would remain viable, and each State has committed to manage its wolf population at or above viable population levels. The wolf's numeric and distributional recovery criteria in the Great Lakes area have been met.
Historical Context of Our Analysis
When reviewing the current status of a species, it is important to understand and evaluate the effects of lost historical range on the viability of the species in its current range. In fact, when we consider the status of a species in its current range, we are considering whether, without the species' lost historical range, the species is endangered or threatened. Range reduction may result in: Reduced numbers of individuals and populations; changes in available resources (such as food) and, consequently, range carrying capacity; changes in demographic characteristics (survival, reproductive rate, metapopulation structure, etc.); and changes in genetic diversity and gene flow. These in turn can increase a species' vulnerability to a wide variety of threats, such as habitat loss, restricted gene flow, or having all or most of its populations affected by a catastrophic event such as a hurricane, fire, or disease outbreak. In other words, past range reduction can reduce the redundancy, resiliency, and representation of a species in its remaining range, such that a species may meet the definition of an “endangered species” or “threatened species” under the Act
s habitat loss, restricted gene flow, or having all or most of its populations affected by a catastrophic event such as a hurricane, fire, or disease outbreak. In other words, past range reduction can reduce the redundancy, resiliency, and representation of a species in its remaining range, such that a species may meet the definition of an “endangered species” or “threatened species” under the Act. Thus, loss of historical range is not necessarily determinative of a species' status, but must be considered in the context of all factors affecting a species. In addition to considering the effects that loss of historical range has had on the current and future viability of the species, we must also consider the causes of that loss of historical range. If the causes of the loss are still continuing, then that loss is also relevant as evidence of the effects of an ongoing threat.
As indicated above, gray wolves historically occupied most of the range of the gray wolf entity (see Historical Range). The gray wolf range of the gray wolf entity began receding after the arrival of Europeans as a result of deliberate killing of wolves by humans and government funded bounty programs aimed at eradication (USFWS 2018, pp. 7-11). Further, many historical habitats were converted into agricultural land (Paquet and Carbyn 2003, p. 483), and natural food sources such as deer and elk were reduced, eliminated, or replaced with domestic livestock, which can become anthropogenic food sources for gray wolves (Young 1944 in Fritts et al. 1997, p. 8). The resulting reduction in range and population were dramatic—by the 1970s gray wolves occupied only a small fraction of their historical range (figure 2). Although the range of the gray wolf in the gray wolf entity has significantly expanded since 1978, its size and distribution remain below historical levels
ome anthropogenic food sources for gray wolves (Young 1944 in Fritts et al. 1997, p. 8). The resulting reduction in range and population were dramatic—by the 1970s gray wolves occupied only a small fraction of their historical range (figure 2). Although the range of the gray wolf in the gray wolf entity has significantly expanded since 1978, its size and distribution remain below historical levels. Today, gray wolves within the gray wolf entity exist as a metapopulation spread across northern Minnesota, Michigan, and Wisconsin, and a small number of colonizing wolves in the west coast United States (USFWS 2018, pp. 22-23) (figure 2).
The alterations to gray wolf historical numbers and populations within the gray wolf entity increased the vulnerability of the gray wolf entity to a wide variety of threats that would not be at issue without such massive range reduction. Some of these threats were identified in the 1978 reclassification (43 FR 9607, March 9, 1978), including reduction in available food (prey) resources, and direct killing by humans. In addition to these considerations, in this proposed rule we also consider availability of suitable habitat, disease and parasites, and climate change. We analyze these potential threats to the gray wolf entity below under Summary of Factors Affecting the Species.
While range reduction may also result in changes in genetic diversity and gene flow, or cause changes in population demographics, we do not address genetic diversity or demographics of the gray wolf entity below because we are not aware of any information indicating that these are potential threats to wolves in the gray wolf entity. Wolves in the entity appear to be genetically and demographically healthy
nge reduction may also result in changes in genetic diversity and gene flow, or cause changes in population demographics, we do not address genetic diversity or demographics of the gray wolf entity below because we are not aware of any information indicating that these are potential threats to wolves in the gray wolf entity. Wolves in the entity appear to be genetically and demographically healthy. Not only do they include wolves of differing and mixed genetic origin, but they exist as part of larger metapopulations—adverse effects resulting from genetic drift, demographic shifts, and local environmental fluctuations can be countered by influxes of individuals and their genetic diversity from other subpopulations of the metapopulation.
Summary of Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533) and its implementing regulations (50 CFR part 424) set forth the procedures for adding species to, reclassifying species on, or removing species from the Federal List of Endangered and i.e., reclassification from endangered to threatened).
For the purposes of this proposed rule, we define the “foreseeable future” to be the extent to which, given the amount and substance of available data, we can anticipate events or effects, or reliably extrapolate threat trends that relate to the status of the gray wolf entity. It took a considerable length of time for public attitudes and regulations to result in a social climate that promoted and allowed for wolf recovery within the gray wolf entity. The length of time over which this shift occurred, and the ensuing stability in those attitudes, gives us confidence that this social climate will persist. Also, the Great Lakes States, which contain the vast majority of wolves within the gray wolf entity, have had a solid history of cooperating with and assisting in wolf recovery and have made a commitment, through legislative actions, to continue these activities
over which this shift occurred, and the ensuing stability in those attitudes, gives us confidence that this social climate will persist. Also, the Great Lakes States, which contain the vast majority of wolves within the gray wolf entity, have had a solid history of cooperating with and assisting in wolf recovery and have made a commitment, through legislative actions, to continue these activities. Washington, Oregon, and California are also committed to conserving wolves as demonstrated by development of management plans and laws and regulations that protect wolves. We are not aware of any information indicating that the commitment of the Great Lakes States and west coast States to gray wolf conservation will change and conclude that this commitment will continue. When evaluating the available information, with respect to foreseeable future, we take into account reduced confidence as we forecast further into the future. Finally, we note that there is a proposed revision to 50 CFR part 424 that creates a regulatory framework for the phrase “foreseeable future.” This proposal is not a departure from how we have implemented the phrase, but rather is meant to codify the framework we have been implementing. Thus, while we are not bound to the proposed revised regulations because they are not final, our interpretation of “foreseeable future” in this rule is consistent with them.
In considering what factors might constitute threats, we must look beyond the exposure of the species to a particular factor to evaluate whether the species may respond to the factor in a way that causes actual impacts to the species. If there is exposure to a factor and the species responds negatively, the factor may be a threat, and during the status review, we attempt to determine how significant a threat it is. The threat is significant if it drives or contributes to the risk of extinction of the species, such that the species warrants listing as endangered or threatened as those terms are defined by the Act
to the species. If there is exposure to a factor and the species responds negatively, the factor may be a threat, and during the status review, we attempt to determine how significant a threat it is. The threat is significant if it drives or contributes to the risk of extinction of the species, such that the species warrants listing as endangered or threatened as those terms are defined by the Act. However, the mere identification of factors that could affect a species negatively may not be sufficient to compel a finding that the species warrants listing. The information must include evidence sufficient to suggest that the potential threat is likely to materialize and that it has the capacity ( i.e., it should be of sufficient magnitude and extent) to affect the species' status such that it meets the definition of an endangered species or threatened species under the Act.
Gray wolves that occur in the gray wolf entity are currently listed as endangered under the Act, except those wolves in Minnesota, which are listed as threatened. In this analysis we evaluate threat factors currently facing the gray wolf entity and those that are reasonably likely to have a negative effect on the viability of wolf populations in the gray wolf entity if the protections of the Act were not in place. Our analysis of threat factors below does not consider the potential for effects to C. lupus in areas where the species has been extirpated—rather, effects are considered in the context of the present population. As explained in our significant portion of the range (SPR) final policy (79 FR 37578; July 1, 2014), we take into account the effect lost historical range may have on the current and future viability of a species in the range it currently occupies, and also whether the causes of that loss are evidence of ongoing or future threats to the species. We do this through our analysis of factors affecting the species
in our significant portion of the range (SPR) final policy (79 FR 37578; July 1, 2014), we take into account the effect lost historical range may have on the current and future viability of a species in the range it currently occupies, and also whether the causes of that loss are evidence of ongoing or future threats to the species. We do this through our analysis of factors affecting the species. A species' current condition reflects the effects of historical range loss and, because threat factors are evaluated in the context of the species' current condition, historical range contraction may affect the outcome of our analysis.
Based on our review of the best available scientific and commercial information, we have identified several factors that could potentially be significant threats to the gray wolf entity. We summarize our analysis of these factors, and factors identified at the time of listing, below. We considered and evaluated the best available scientific and commercial data for our analyses.
Human-Caused Mortality
Human-caused mortality was identified as the main factor causing the decline of gray wolves at the time of listing (43 FR 9611, March 9, 1978), and an active eradication program is the sole reason that wolves were extirpated from their historical range in the United States (Weaver 1978, p. i). European settlers attempted to eliminate the wolf entirely, primarily due to the threat or reality of attacks on livestock, and the U.S. Congress passed a wolf bounty that covered the Northwest Territories in 1817. Bounties on wolves subsequently became the norm for States across the species' range. For example, in Michigan, an 1838 wolf bounty became the ninth law passed by the First Michigan Legislature; this bounty remained in place until 1960. A Wisconsin bounty was instituted in 1865 and was repealed about the time wolves were extirpated from the State in 1957. Minnesota maintained a wolf bounty until 1965
nties on wolves subsequently became the norm for States across the species' range. For example, in Michigan, an 1838 wolf bounty became the ninth law passed by the First Michigan Legislature; this bounty remained in place until 1960. A Wisconsin bounty was instituted in 1865 and was repealed about the time wolves were extirpated from the State in 1957. Minnesota maintained a wolf bounty until 1965. As the first provisional governments in the Pacific Northwest region were formed, they too enacted wolf bounties (Hampton 1997, pp. 107-108).
Protection of the gray wolf under the Act and State endangered-species statutes prohibited the intentional killing of wolves except under very limited circumstances, such as in defense of human life, for scientific or conservation purposes, or under special regulations intended to reduce wolf depredations of livestock or other domestic animals. Aside from the reintroduction of wolves into portions of the northern Rocky Mountains, the regulation of human-caused wolf mortality is the primary reason wolf numbers have significantly increased and their range has expanded since the mid-to-late 1970s.
Two Minnesota studies provide some limited insight into the extent of human-caused wolf mortality before and after the species' listing. On the basis of bounty data from a period that predated wolf protection under the Act by 20 years, Stenlund (1955, p. 33) found an annual human-caused mortality rate of 41 percent. Fuller (1989, pp. 23-24) provided 1980-86 data from a north-central Minnesota study area and found an annual human-caused mortality rate of 29 percent, a figure that includes 2-percent mortality from legal depredation-control actions. Drawing conclusions from comparisons of these two studies, however, is difficult due to the confounding effects of habitat quality, exposure to humans, prey density, differing time periods, and vast differences in study design
entral Minnesota study area and found an annual human-caused mortality rate of 29 percent, a figure that includes 2-percent mortality from legal depredation-control actions. Drawing conclusions from comparisons of these two studies, however, is difficult due to the confounding effects of habitat quality, exposure to humans, prey density, differing time periods, and vast differences in study design. Nonetheless, these figures provide clear support for the contention that human-caused mortality decreased significantly once the wolf became protected under the Act.
Humans kill wolves for a number of reasons. In locations where people, livestock, and wolves coexist, some wolves are killed to resolve conflicts with livestock and pets (Fritts et al. 2003, p. 310; Woodroffe et al. 2005, pp. 86-107, 345-347). Occasionally, wolves are killed accidentally ( e.g., wolves are hit by vehicles, mistaken for coyotes and shot, caught in traps set for other animals, or subject to accidental capture-related mortality during conservation or research efforts) (Bangs et al. 2005, p. 346). A few wolves have been killed by people who stated that they believed their physical safety was being threatened. Many wolf killings, however, are intentional, illegal, and never reported to authorities.
The number of illegal killings is difficult to estimate and impossible to accurately determine because they generally occur with few witnesses. Illegal killing was estimated to make up 70 percent of the total mortality rate in a north-central Minnesota wolf population and 24 percent in the northern Rocky Mountains population (Liberg et al. 2011, pp. 3-5). Liberg et al. (2011, pp. 3-5) suggest more than two-thirds of total poaching may go undetected, and that illegal killing may pose a threat to wolves; however, poaching has not prevented population resurgence in either the Great Lakes area or the northern Rocky Mountains, as evidenced by population growth in those areas
rcent in the northern Rocky Mountains population (Liberg et al. 2011, pp. 3-5). Liberg et al. (2011, pp. 3-5) suggest more than two-thirds of total poaching may go undetected, and that illegal killing may pose a threat to wolves; however, poaching has not prevented population resurgence in either the Great Lakes area or the northern Rocky Mountains, as evidenced by population growth in those areas.
Vehicle collisions contribute to wolf mortality rates throughout their range in the lower 48 United States. This type of mortality is expected to rise with increasing wolf populations and as wolves colonize areas with more human development and a denser network of roads and vehicle traffic; however, mortalities due to vehicle collisions will likely constitute a small proportion of total mortalities.
Each of the States in the current range of gray wolves in the contiguous United States conduct scientific research and monitoring of wolf populations. Even the most intensive and disruptive of these activities (anesthetizing for the purpose of radio-collaring) involves a very low rate of mortality for wolves (73 FR 10542, February 27, 2008). We expect that capture-related mortality during wolf monitoring, nonlethal control, and research activities will remain below three percent of the wolves captured, and will have an insignificant impact on population dynamics.
We are unaware of any wolves that have been removed from the wild solely for educational purposes in recent years. Wolves that are used for such purposes are typically privately held captive-reared offspring of wolves that were already in captivity for other reasons. However, States may get requests to place wolves that would otherwise be euthanized in captivity for research or educational purposes. Such requests have been and will continue to be rare, would be closely regulated by the State wildlife-management agencies through the requirement for State permits for protected species, and would not substantially increase human-caused wolf mortality rates
asons. However, States may get requests to place wolves that would otherwise be euthanized in captivity for research or educational purposes. Such requests have been and will continue to be rare, would be closely regulated by the State wildlife-management agencies through the requirement for State permits for protected species, and would not substantially increase human-caused wolf mortality rates.
Other sources of human-caused mortality include intentional and legal actions, such as lethal depredation control and killing wolves in defense of human life or property. Although most wolf-human conflicts are solved using nonlethal methods, in a few instances lethal control is warranted to control a wolf to protect human life and property. The number of wolves killed for this purpose is small. For example, from 2004 to 2014, State or Federal agents killed 26 wolves for these purposes in the State of Michigan (an average of around 0.5 percent of the population each year) (Roell et al. 2010, p. 9; Beyer in litt. 2018). In the western States, since the first pack was confirmed in Washington in 2008, one wolf has been killed by a private individual who claimed self-defense. Although the number of wolves killed in defense of human life and property may be slightly higher in areas with greater human density and may increase after delisting as authority for this action expands (see Post-delisting Management), overall this type of mortality is rare and is not expected to have a significant impact on wolf populations.
Lethal control of depredating wolves was authorized in Minnesota while wolves have been listed (under the authority of a regulation (50 CFR 17.40(d)) under section 4(d) of the Act), but such control was not authorized in Michigan or Wisconsin, except for the several years when such control was authorized under a permit from the USFWS or while wolves were delisted under previous actions. Lethal control of depredating wolves is not authorized in the listed portion of Oregon, Washington, or in California
authority of a regulation (50 CFR 17.40(d)) under section 4(d) of the Act), but such control was not authorized in Michigan or Wisconsin, except for the several years when such control was authorized under a permit from the USFWS or while wolves were delisted under previous actions. Lethal control of depredating wolves is not authorized in the listed portion of Oregon, Washington, or in California. The Minnesota wolf-depredation-control program euthanized from 20 (in 1982) to 262 (in 2015) wolves annually, and averaged between 2.2 to 7.6 percent of the wolf population annually. During the times wolves were listed and depredation control was the primary means of management in the State, the Minnesota wolf population continued to grow or remain stable while experiencing these levels of lethal control. During the times that lethal control of depredating wolves was conducted in Wisconsin and Michigan, there was no evidence of resulting adverse impacts to the maintenance of a viable wolf population in those States. In Wisconsin, a total of 256 wolves were killed for depredation control in the State, including 46 legally shot by private landowners, during the 59 months that wolves were delisted in the State. A total of 50 wolves were killed by the Michigan Department of Natural Resources (MI DNR) and the U.S. Department of Agriculture, Animal and Plant Health Inspection Service (USDA-APHIS), Wildlife Services in response to depredation events during that time period. Following delisting, wolf depredation control in Wisconsin and Michigan would again occur, and be carried out according to their State management plans. We anticipate the level of mortality due to depredation control that would take place would be similar to what was observed during those times. See the Post-delisting Management section for a more detailed discussion of legal control of problem wolves (primarily for depredation control)
was previously believed to occur in wolf populations. This means that human-caused mortality is not simply added to “natural” mortality, but rather replaces a portion of it. Creel and Rotella (2010) reexamined this concept with regard to wolves and found that, contrary to the previously held belief, wolf population growth declined as human-caused mortality increased (Creel and Rotella 2010, p. 3). Their study concludes that wolves can be harvested within limits, but that human-caused mortality was strongly additive in total mortality (Creel and Rotella 2010, p. 6).
The wolf population in the northern Rocky Mountains States of Idaho, Montana, and Wyoming provides a good example of the effects of increased human-caused mortality on population growth rates. From 1995 to 2008, wolf populations increased an average of 23 percent annually (range: 9 percent to 50 percent; USFWS et al. 2016, table 6b), while from 1999 to 2008, human-caused mortality removed an average of approximately 12 percent of the minimum estimated population each year (range: 7 percent to 16 percent; see USFWS et al. 2000-2009). Between 2009 and 2015, some or all of the northern Rocky Mountains States (dependent upon the Federal status of wolves) instituted fair-chase wolf hunting seasons with the objective of slowing or reversing population growth while continuing to maintain wolf populations well above federal recovery requirements in their respective States. During those years when legal harvest occurred, human-caused mortality increased to an average of 29 percent of the minimum estimated population (range: 23 percent to 36 percent; see USFWS et al. 2010, 2012-2016), while the annual growth rate declined to an average of approximately 1 percent annually (range: -7 percent to 4 percent; see USFWS et al. 2010, 2012-2016). Where harvest occurs, the species' high levels of reproduction and immigration can compensate for mortality rates of 17 percent to 48 percent (USFWS 2018, p. 6)
ted population (range: 23 percent to 36 percent; see USFWS et al. 2010, 2012-2016), while the annual growth rate declined to an average of approximately 1 percent annually (range: -7 percent to 4 percent; see USFWS et al. 2010, 2012-2016). Where harvest occurs, the species' high levels of reproduction and immigration can compensate for mortality rates of 17 percent to 48 percent (USFWS 2018, p. 6). Thus, although 2009 to 2015 is a relatively short time period from which to draw inferences, the population trends observed in the Northern Rocky Mountains suggest that the northern Rocky Mountains wolf population may be able to sustain an approximate 30 percent annual human-caused mortality rate while continuing to maintain a stable to slightly increasing population.
The States of Minnesota, Michigan, and Wisconsin have committed to continue to regulate human-caused mortality so that it does not reduce the wolf population below recovery level and have adequate laws and regulations to fulfill those commitments and ensure that the wolf population in the Great Lakes area remains above recovery levels (See Post-delisting Management). Washington, Oregon, and California are also committed to conserving wolves as demonstrated by development of management plans and laws and regulations that protect wolves. Furthermore, each post-delisting management entity (State, Tribal, and Federal) has experienced and professional wildlife staff to ensure those commitments can be accomplished.
Effects on Wolf Social Structure
Human-caused mortality of reproductive gray wolves could negatively affect gray wolf populations because wolves have a complex social system in which usually only the dominant male and female in a pack breed. Consequently, the death of one or both of the breeders may negatively affect the pack (by leading to pack dissolution) and the population as a whole (by slowing or reducing population growth)
Human-caused mortality of reproductive gray wolves could negatively affect gray wolf populations because wolves have a complex social system in which usually only the dominant male and female in a pack breed. Consequently, the death of one or both of the breeders may negatively affect the pack (by leading to pack dissolution) and the population as a whole (by slowing or reducing population growth). However, studies indicate these effects are context-dependent and that the availability of replacement breeders and timing of mortality can moderate the consequences of breeder loss (Borg et al. 2014, entire; Brainerd et al. 2008, entire). In populations that are at or near carrying capacity, where breeder replacement and subsequent reproduction occurs relatively quickly, population growth rate is largely unaffected by breeder loss (Borg et al. 2014, pp. 6-7). Large colonizing populations (> 75 wolves) have similar times to breeder replacement and subsequent reproduction as populations at or near carrying capacity, while small recolonizing populations (≤75 wolves) take about twice as long to replace breeders and subsequently reproduce (Brainerd et al. 2008, pp. 89, 93). Therefore, the effects of breeder loss may be greatest on small recolonizing gray wolf populations. Studies also indicate that mortality of breeding gray wolves is more likely to lead to pack dissolution and reduced reproduction when mortality occurs during the breeding season (Borg et al. 2014, p. 8) and when pack sizes are small (Borg et al. 2014, pp. 5-6; Brainerd et al. 2008, p. 94).
Gray wolf pack social structure is very adaptable and resilient. Breeding members can be quickly replaced from either within or outside the pack, and pups can be reared by another pack member should their parents die (USFWS 2018, p. 6). Consequently, wolf populations can rapidly overcome severe disruptions, such as pervasive human-caused mortality or disease
p. 5-6; Brainerd et al. 2008, p. 94).
Gray wolf pack social structure is very adaptable and resilient. Breeding members can be quickly replaced from either within or outside the pack, and pups can be reared by another pack member should their parents die (USFWS 2018, p. 6). Consequently, wolf populations can rapidly overcome severe disruptions, such as pervasive human-caused mortality or disease. Although we acknowledge that breeder loss can and will occur in the future regardless of Federal status, we conclude that the effects of breeder loss on wolf populations (or the gray wolf entity) as a whole are likely to be minimal as long as adequate regulatory mechanisms are in place to ensure sufficient population size is maintained.
The Role of Public Attitudes
In our 1978 rule reclassifying wolves, we indicated that regulations prohibiting the killing of wolves, even wolves that may be attacking livestock and pets, such as the Federal regulations in place at that time in Minnesota, may work against gray wolves by creating an adverse public attitude toward the species. We acknowledge that public attitudes towards wolves vary with demographics, change over time, and can affect human behavior toward wolves, including poaching (illegal killing) of wolves (see the following studies and reviews: Kellert 1985, 1990, 1999; Nelson and Franson 1988; Kellert et al. 1996; Wilson 1999; Browne-Nuñez and Taylor 2002; Williams et al. 2002; Manfredo et al. 2003; Naughton-Treves et al. 2003; Schanning 2009; Mertig 2004; Chavez et al. 2005; Schanning and Vazquez 2005; Beyer et al. 2006; Hammill 2007; Treves et al. 2009; Wilson and Bruskotter 2009; Treves and Martin 2011; Treves et al. 2013; Madden and McQuinn 2014). However, the factors that affect people's attitudes and et al. 2013, p. 316 and references therein; also see Olson et al. 2014, entire and Chapron and Treves 2016, entire)
Schanning 2009; Mertig 2004; Chavez et al. 2005; Schanning and Vazquez 2005; Beyer et al. 2006; Hammill 2007; Treves et al. 2009; Wilson and Bruskotter 2009; Treves and Martin 2011; Treves et al. 2013; Madden and McQuinn 2014). However, the factors that affect people's attitudes and et al. 2013, p. 316 and references therein; also see Olson et al. 2014, entire and Chapron and Treves 2016, entire). Thus, it is unclear how delisting and the changes in wolf management subsequent to delisting, such as implementation of wolf harvests, may affect attitudes, human behavior and, ultimately, wolf mortality.
We expect that some segments of the public will be more tolerant of wolf management at the State level because it may be perceived by some as more flexible than Federal regulation, whereas other segments may continue to prefer Federal management due to a perception that it is more protective. State wildlife agencies have professional staff dedicated to disseminating accurate, science-based information about wolves and wolf management within their respective States. In addition, several States have convened advisory committees to engage stakeholders in discussing and addressing conflicts related to wolves (for example, Washington ( https://wdfw.wa.gov/about/advisory/wag/ ) and Wisconsin ( https://dnr.wi.gov/topic/WildlifeHabitat/wolf/committee.html )). As the status and management of the gray wolf evolves, continued collaboration between managers and researchers to monitor public attitudes toward wolves and their management will be necessary.
Human-Caused Mortality Summary
Despite human-caused mortalities of wolves, wolf populations have continued to increase in both numbers and range. Wolf population growth will likely slow as densities increase in suitable habitat
ent of the gray wolf evolves, continued collaboration between managers and researchers to monitor public attitudes toward wolves and their management will be necessary.
Human-Caused Mortality Summary
Despite human-caused mortalities of wolves, wolf populations have continued to increase in both numbers and range. Wolf population growth will likely slow as densities increase in suitable habitat. Wolves are less likely to persist in more unfavorable habitats due to depredation management, illegal killing, incidental mortality (for example, vehicle collision), natural mortality (disease, starvation, and intraspecific aggression), and other means. Once wolf populations become established, we should expect to see populations fluctuate around an equilibrium resulting from fluctuations in birth and mortality rates.
Minnesota, Wisconsin, and Michigan will utilize adaptive management to respond to wolf population increases or decreases to maintain populations at sustainable levels well above management objectives. State management plans in these three states that would be implemented following delisting manage for a minimum wolf population of 1,600 in Minnesota, 250 in Wisconsin (with a management goal of 350), and 200 in Michigan. These minimum population numbers are well above Federal recovery requirements defined in the Eastern Timber Wolf Recovery Plan. As wolf population numbers are currently much higher in each of these three States, we can expect to see some reduction in wolf populations in the Great Lakes areas if they are delisted as States implement lethal depredation control and begin to institute wolf hunting seasons with the objective of slowing or reversing population growth. However, the ultimate goal of these three States is to maintain wolf populations well above Federal recovery requirements in their respective States.
The 2010 State management plan for Oregon and the 2016 plan for California do not include population-management goals (Oregon Department of Fish and Wildlife (ODFW) 2010, p
nting seasons with the objective of slowing or reversing population growth. However, the ultimate goal of these three States is to maintain wolf populations well above Federal recovery requirements in their respective States.
The 2010 State management plan for Oregon and the 2016 plan for California do not include population-management goals (Oregon Department of Fish and Wildlife (ODFW) 2010, p. 27; California Department of Fish and Wildlife (CDFW) 2016a, p. 12); however, this is likely to be addressed in the forthcoming Oregon plan revision as the draft plan revision currently suggests that 300 wolves are the “minimum population management threshold” for the State (ODFW 2017, p. 17). While the 2011 Washington State management plan does not include population-management goals, it includes recovery objectives intended to ensure the reestablishment of a self-sustaining population of wolves in Washington (Wiles et al. 2011, p. 9; also see Post-delisting Management in the West ). In these States, wolf populations will likely be managed to ensure progress towards recovery objectives while also minimizing livestock losses caused by wolves.
Habitat and Prey Availability
Gray wolves are habitat generalists (Mech and Boitani 2003, p. 163) and once occupied or transited most of the United States, except the southeast. However, much of the historical range of gray wolves (Chambers et al. 2012, pp. 34-42) in the contiguous United States has been modified due to human use. While lone wolves can travel through, or temporarily live, almost anywhere (Jimenez et al. 2017, p. 1), large portions of gray wolf historical range is no longer suitable habitat to support wolf packs (Oakleaf et al. 2006, p. 559; Carroll et al. 2006, p. 32, Mladenoff et al. 1995, p. 287). Much of the area that wolves currently occupy corresponds to what is considered “suitable” wolf habitat in the lower 48 States as modeled by Oakleaf et al. (2006, entire), Carroll et al. (2006, entire), Mladenoff (1995, entire), and Mladenoff et al
cal range is no longer suitable habitat to support wolf packs (Oakleaf et al. 2006, p. 559; Carroll et al. 2006, p. 32, Mladenoff et al. 1995, p. 287). Much of the area that wolves currently occupy corresponds to what is considered “suitable” wolf habitat in the lower 48 States as modeled by Oakleaf et al. (2006, entire), Carroll et al. (2006, entire), Mladenoff (1995, entire), and Mladenoff et al. (1999, entire). It is also expected that wolves will continue to recolonize areas of the Pacific Northwest where suitable habitat has been identified (Maletzke et al. 2015, entire; ODFW 2015, entire). We consider suitable habitat as forested terrain containing adequate wild ungulate populations (elk, white-tailed deer, and mule deer) to support a wolf population. Suitable habitat has minimal roads and human development, as human access to areas inhabited by wolves can result in wolf mortality.
Great Lakes Area: Suitable Habitat
Various researchers have investigated habitat suitability for wolves in the central and eastern portions of the United States. Most of these efforts have focused on using a combination of human density, density of agricultural lands, deer density or deer biomass, and road density, or have used road density alone to identify areas where wolf populations are likely to persist or become established (Mladenoff et al. 1995, pp. 284-285; 1997, pp. 23-27; 1998, pp. 1-8, 1999; pp. 39-43; Harrison and Chapin 1997, p. 3; 1998, pp. 769-770; Wydeven et al. 2001, pp. 110-113; Erb and Benson 2004, p. 2; Potvin et al. 2005, pp. 1661-1668; Mladenoff et al. 2009, pp. 132-135).
To a large extent, road density has been adopted as the best predictor of habitat suitability in the Midwest due to the connection between roads and human-caused wolf mortality. Several studies demonstrated that wolves generally did not maintain breeding packs in areas with a road density greater than about 0.9 to 1.1 linear mi per mi 2 (0.6 to 0.7 km per km 2 ) (Thiel 1985, pp. 404-406; Jensen et al. 1986, pp. 364-366; Mech et al
s been adopted as the best predictor of habitat suitability in the Midwest due to the connection between roads and human-caused wolf mortality. Several studies demonstrated that wolves generally did not maintain breeding packs in areas with a road density greater than about 0.9 to 1.1 linear mi per mi 2 (0.6 to 0.7 km per km 2 ) (Thiel 1985, pp. 404-406; Jensen et al. 1986, pp. 364-366; Mech et al. 1988, pp. 85-87; Fuller et al. 1992, pp. 48-51). Work by Mladenoff and associates indicated that colonizing wolves in Wisconsin preferred areas where road densities were less than 0.7 mi per mi 2 (0.45 km per km 2 ) (Mladenoff et al. 1995, p. 289). However, research in the Upper Peninsula of Michigan indicates that, in some areas with low road densities, low deer density appears to limit wolf occupancy (Potvin et al. 2005, pp. 1667-1668) and may prevent recolonization of portions of the Upper Peninsula. In Minnesota, a combination of road density and human density is used by Minnesota Department of Resources (MN DNR) to model suitable habitat. Areas with a human density up to 20 people per mi 2 (8 people per km 2 ) are suitable if they also have a road density less than 0.8 mi per mi 2 (0.5 km per km 2 ). Areas with a human density of less than 10 people per mi 2 (4 people per km 2 ) are suitable if they have road 2 (0.7 km per km 2 ) (Erb and Benson 2004, table 1).
Road density is a useful parameter because it is easily measured and mapped, and because it correlates directly and indirectly with various forms of other human-caused wolf mortality factors. A rural area with more roads generally has a greater human density, more vehicular traffic, greater access by hunters and trappers, more farms and residences, and more domestic animals. As a result, there is a greater likelihood that wolves in such an area will encounter humans, domestic animals, and various human activities
directly with various forms of other human-caused wolf mortality factors. A rural area with more roads generally has a greater human density, more vehicular traffic, greater access by hunters and trappers, more farms and residences, and more domestic animals. As a result, there is a greater likelihood that wolves in such an area will encounter humans, domestic animals, and various human activities. These encounters may result in wolves being hit by motor vehicles, being controlled by government agents after becoming involved in depredations on domestic animals, being shot intentionally by unauthorized individuals, being trapped or shot accidentally, or contracting diseases from domestic dogs (Mech et al. 1988, pp. 86-87; Mech and Goyal 1993, p. 332; Mladenoff et al. 1995, pp. 282, 291). Based on mortality data from radio-collared Wisconsin wolves from 1979 to 1999, natural causes of death predominate (57 percent of mortalities) in areas with road densities below 1.35 mi per mi 2 (0.84 km per km 2 ), but human-related factors produced 71 percent of the wolf deaths in areas with higher road densities (Wydeven et al. 2001, pp. 112-113).
Some researchers have used a road density of 1 mi per mi 2 (0.6 km per km 2 ) of land area as an upper threshold for suitable wolf habitat. However, the common practice in more recent studies is to use road density to predict probabilities of persistent wolf pack presence in an area. Areas with road densities less than 0.7 mi per mi 2 (0.45 km per km 2 ) are estimated to have a greater than 50 percent probability of wolf pack colonization and persistent presence, and areas where road density exceeded 1 mi per mi 2 (0.6 km per km 2 ) have less than a 10 percent probability of occupancy (Mladenoff et al. 1995. pp. 288-289; Mladenoff and Sickley 1998, p. 5; Mladenoff et al. 1999, pp. 40-41)
road densities less than 0.7 mi per mi 2 (0.45 km per km 2 ) are estimated to have a greater than 50 percent probability of wolf pack colonization and persistent presence, and areas where road density exceeded 1 mi per mi 2 (0.6 km per km 2 ) have less than a 10 percent probability of occupancy (Mladenoff et al. 1995. pp. 288-289; Mladenoff and Sickley 1998, p. 5; Mladenoff et al. 1999, pp. 40-41). Wisconsin researchers view areas with greater than 50 percent probability as “primary wolf habitat,” areas with 10 to 50 percent probability as “secondary wolf habitat,” and areas with less than 10 percent probability as unsuitable habitat (Wisconsin Department of Natural Resources (WI DNR) 1999, pp. 47-48).
The territories of packs that do occur in areas of high road density, and hence with low expected probabilities of occupancy, are generally near broad areas of more suitable habitat that are likely serving as a source of wolves, thereby assisting in maintaining wolf presence in the higher road density areas and, therefore, less-suitable areas (Mech 1989, pp. 387-388; Wydeven et al. 2001, p. 112). The predictive ability of this model was questioned (Mech 2006a, 2006b) and responded to (Mladenoff et al. 2006), and an updated analysis of Wisconsin pack locations and habitat was completed (Mladenoff et al. 2009). This model maintains that road density is still an important indicator of suitable wolf habitat; however, lack of agricultural land is also a strong predictor of habitat that wolves occupy.
It appears that essentially all suitable habitat in Minnesota is now occupied, range expansion has slowed or possibly ceased, and the wolf population within the State has stabilized (Erb and Benson 2004, p. 7; Erb and Don Carlos 2009, pp. 57, 60). This suitable habitat closely matches the areas designated as Wolf Management Zones 1 through 4 in the Revised Recovery Plan (USFWS 1992, p. 72), which are identical in area to Minnesota Wolf Management Zone A (MN DNR 2001, appendix III)
expansion has slowed or possibly ceased, and the wolf population within the State has stabilized (Erb and Benson 2004, p. 7; Erb and Don Carlos 2009, pp. 57, 60). This suitable habitat closely matches the areas designated as Wolf Management Zones 1 through 4 in the Revised Recovery Plan (USFWS 1992, p. 72), which are identical in area to Minnesota Wolf Management Zone A (MN DNR 2001, appendix III).
Recent surveys for Wisconsin wolves and wolf packs show that wolves have now recolonized the areas predicted by habitat models to have low, moderate, and high probability of occupancy (primary and secondary wolf habitat). The late-winter 2017-18 Wisconsin wolf survey identified packs occurring throughout the central Wisconsin forest area (Wolf Management Zone 2) and across the northern forest zone (Zone 1), with highest pack densities in the northwest and north-central forest (WI DNR 2018, entire).
Michigan wolf surveys in winter 2017-18 continue to show wolf pairs or packs (defined by Michigan DNR as two or more wolves traveling together) in every Upper Peninsula county (Huntzinger et al. 2005, p. 6; MI DNR 2018, entire).
Habitat suitability studies in the Upper Midwest indicate that the only large areas of suitable or potentially suitable habitat areas that are currently unoccupied by wolves are located in the northern Lower Peninsula of Michigan (Mladenoff et al. 1997, p. 23; Mladenoff et al. 1999, p. 39; Potvin 2003, pp. 44-45; Gehring and Potter 2005, p. 1239). One published Michigan study (Gehring and Potter 2005, p. 1239) estimates that these areas could host 46 to 89 wolves; a graduate thesis estimates that 110-480 wolves could exist in the northern Lower Peninsula (Potvin 2003, p. 39). The northern Lower Peninsula is separated from the Upper Peninsula by the Straits of Mackinac, whose 4-mile (6.4-km) width freezes during mid- and late-winter in some years
ublished Michigan study (Gehring and Potter 2005, p. 1239) estimates that these areas could host 46 to 89 wolves; a graduate thesis estimates that 110-480 wolves could

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