Endangered and Threatened Wildlife and Plants; Proposed Rule To Revise the List of Endangered and Threatened Wildlife for the Gray Wolf (Canis lupus) in the Eastern United States, Initiation of Status Reviews for the Gray Wolf and for the Eastern Wolf (Canis lycaon)

DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

[Docket No. FWS-R3-ES-2011-0029; 92220-1113-000; ABC Code: C6]

RIN 1018-AX57

Endangered and Threatened Wildlife and Plants; Proposed Rule To Revise the List of Endangered and Threatened Wildlife for the Gray Wolf (

Canis lupus

) in the Eastern United States, Initiation of Status Reviews for the Gray Wolf and for the Eastern Wolf (

Canis lycaon

)

AGENCY:

Fish and Wildlife Service, Interior.

ACTION:

Proposed rule, initiation of status reviews.

SUMMARY:

We, the U.S. Fish and Wildlife Service (Service or USFWS) are re-evaluating the listing of the Minnesota population of gray wolves (

Canis lupus

) and propose to revise it to conform to current statutory and policy requirements. We propose to identify the Minnesota population as a Western Great Lakes (WGL) Distinct Population Segment (DPS) of the gray wolf and to remove this DPS from the List of Endangered and Threatened Wildlife. We propose these actions because the best available scientific and commercial information indicates that the WGL DPS does not meet the definitions of threatened or endangered under the Act.

This proposed rule, if made final, would remove the currently designated critical habitat for the gray wolf in Minnesota and Michigan and the current special regulations for gray wolves in Minnesota. We also propose to revise the range of the gray wolf (the species

C. lupus

) by removing all or parts of 29 eastern states that we now recognize were not part of the historical range of the gray wolf. New information indicates that these areas should not have been included in the original listing of the gray wolf.

In this proposed rule, we recognize recent taxonomic information indicating that the gray wolf subspecies

Canis lupus lycaon

should be elevated to the full species

C. lycaon.

Given that a complete status review of this newly recognized species has never been conducted, we are initiating a rangewide review of the conservation status of

C. lycaon

in the United States and Canada. This rule also constitutes the initiation of our five-year review of the status of gray wolves under section 4(c)(2) of the Act, as well as the initiation of status reviews specific to gray wolves in the Pacific Northwest and Mexican wolves in the Southwest United States and Mexico.

DATES:

Comment submission:

We will accept comments received or postmarked on or before July 5, 2011.

Public hearings:

We will hold two public hearings on this proposed rule scheduled on May 18, 2011 and on June 8, 2011. Informational meetings will be held from 6 p.m. to 7:15 p.m., followed by the public hearings from 7:30 p.m. to 9 p.m.

ADDRESSES:

Comment submission:

You may submit comments by one of the following methods:

Electronically:

Go to the Federal eRulemaking Portal:

http://www.regulations.gov.

In the Enter Keyword or ID box, enter FWS-R3-ES-2011-0029, which is the docket number for this rulemaking. Then, in the Search panel at the top 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 “Submit a Comment.”

By hard copy:

Submit by U.S. mail or hand-delivery to: Public Comments Processing, Attn: FWS-R3-ES-2011-0029; Division of Policy and Directives Management; U.S. Fish and Wildlife Service; 4401 N. Fairfax Drive, MS 2042-PDM; Arlington, VA 22203.

We will post all comments on

http://www.regulations.gov.

This generally means that we will post any personal information you provide us (see the Public Comments section below for more information).

Public hearings:

We have scheduled an informational meeting followed by a public hearing in Ashland, Wisconsin, on May 18, 2011, at the Northern Great Lakes Center, 29270 County Highway G. We have scheduled an informational meeting followed by a public hearing in Augusta, Maine, on June 8, 2011, at the Augusta Civic Center, 16 Cony Street. See the Public Hearings section below for more details.

FOR FURTHER INFORMATION CONTACT:

Laura Ragan, 612-713-5350. Direct all questions or requests for additional information to: GRAY WOLF QUESTIONS, U.S. Fish and Wildlife Service, Federal Building, 1 Federal Drive, Ft. Snelling, Minnesota 55111-4056. Additional information is also available on our Web site at

http://www.fws.gov/midwest/wolf.

Individuals who are hearing-impaired or speech-impaired may call the Federal Relay Service at 1-800-877-8337 for TTY assistance.

SUPPLEMENTARY INFORMATION:

Public Comments

We intend that any final action resulting from this proposal will be as accurate and as effective as possible. Therefore, comments, new information, or suggestions from the public, other concerned governmental agencies, the scientific community, industry, or any other interested party concerning this proposed rule are hereby solicited. In particular, we are seeking targeted information and comments on our national wolf strategy and our proposed revision of the Minnesota listing; see items (1)-(2) below. Also, as part of this proposed rule we are announcing initiation of a 5-year status review for

C. lupus

in the conterminous United States and Mexico; initiation of status reviews specific to, respectively, gray wolves in the Pacific Northwest and in the Southwest United States and Mexico; and initiation of a status review for

C. lycaon

throughout its range in the United States and Canada. For these status reviews to be complete and based on the best available scientific and commercial information, we request information on items (9)-(11) below from governmental agencies, Native American Tribes, the scientific community, industry, and any other interested parties.

(1) Biological, commercial trade, or other relevant information concerning our analysis of the current gray wolf listing and the adequacy of our national wolf strategy, with particular respect to our recommended gray wolf listing units (

i.e.,

taxonomic or population units);

(2) Information that forms the basis for revising the currently listed Minnesota group of gray wolves under section 4(c) of the Endangered Species Act of 1973, as amended (Act) (16 U.S.C. 1531

et seq.

), with particular respect to the factors in section 4(a) of the Act, which are:

(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.

(3) Biological, commercial trade, or other relevant data concerning any current or likely future threat, or lack thereof, to wolves in the WGL DPS;

(4) Additional information concerning the range, distribution, population size, population trends, and threats with respect to wolves in the WGL DPS;

(5) Current or planned activities in the WGL DPS and their possible impacts on the wolves and their habitat;

(6) Information concerning the adequacy of the recovery criteria described in the 1992 Recovery Plan for the Eastern Timber Wolf;

(7) The extent and adequacy of Federal, state, and Tribal protection and management that would be provided to wolves in the WGL DPS as delisted species; and

(8) The proposed geographic boundaries of the WGL DPS, and scientific and legal supporting information for alternative boundaries that might result in a larger or smaller DPS, including information on the discreteness and significance of the proposed DPS.

(9) New information concerning the biology and conservation of the gray wolf in the conterminous United States and Mexico that may be informative to the 5-year status review of

Canis lupus,

with particular attention to the listing units described under (1) above, including:

(a) Habitat requirements for feeding, breeding, and sheltering;

(b) Genetics and taxonomy;

(c) Historical and current range including distribution patterns;

(d) Historical and current population levels, and current and projected trends;

(e) Historical, current, and projected levels of suitable gray wolf habitat;

(f) Past, ongoing, and emerging threats to extant gray wolf populations, their habitat, or both; and

(g) Past and ongoing conservation measures for the gray wolf, its habitat, or both.

(10) Information concerning the status of the gray wolf in the Pacific Northwest United States and the gray wolf subspecies

baileyi

(Mexican wolf) in the Southwest United States and Mexico, including:

(a) Habitat requirements for feeding, breeding, and sheltering;

(b) Genetics and taxonomy;

(c) Historical and current range including distribution patterns;

(d) Historical and current population levels, and current and projected trends;

(e) Historical, current, and projected levels of suitable habitat;

(f) Past, ongoing, and emerging threats to these populations, their habitat, or both; and

(g) Past and ongoing conservation measures for these populations, their habitat, or both.

(11) Information concerning the biology, range, and population trends of

Canis lycaon,

including:

(a) Habitat requirements for feeding, breeding, and sheltering;

(b) Genetics and taxonomy;

(c) Historical and current range including distribution patterns;

(d) Historical and current population levels, and current and projected trends;

(e) Historical, current, and projected levels of suitable habitat;

(f) Past, ongoing, and emerging threats to extant populations, their habitat, or both;

(g) Past and ongoing conservation measures for the species, its habitat, or both; and

(h) The potential role that any portion of the historical range of the

C. lycaon

in the United States may play in the persistence and viability of the species.

You may submit your comments and materials by one of the methods listed in

ADDRESSES

. We will not accept comments sent by e-mail or fax or to an address not listed in

ADDRESSES

. Comments must be submitted to

http://www.regulations.gov

before midnight (Eastern Daylight Time) on the date specified in

DATES.

Finally, we will not consider hand-delivered comments that we do not receive, or mailed comments that are not postmarked, by the date specified in

DATES

.

We will post your entire comment—including your personal identifying information—on

http://www.regulations.gov.

If you provide personal identifying information, such as your street address, phone number, or e-mail address, 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.

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-R3-ES-2011-0029, or by appointment, during normal business hours at the following Ecological Services offices:

• Twin Cities, Minnesota Ecological Services Field Office, 4101 American Blvd. E., Bloomington, MN; 612-725-3548.

• Green Bay, Wisconsin Ecological Services Field Office, 2661 Scott Tower Dr., New Franken, WI; 920-866-1717.

• East Lansing, Michigan Ecological Services Field Office, 2651 Coolidge Road, Suite 101, East Lansing, MI; 517-351-2555.

• New England Ecological Services Field Office, U.S. Fish and Wildlife Service, 70 Commercial St., Suite 300, Concord, NH; 603-223-2541.

Public Hearings

We have scheduled an informational meeting followed by a public hearing in Ashland, Wisconsin, on May 18, 2011, at the Northern Great Lakes Center, 29270 County Highway G. The informational meeting will be held from 6 p.m. to 7:15 p.m., followed by a public hearing from 7:30 p.m. to 9 p.m.

A second informational meeting followed by a public hearing will be held in Augusta, Maine, on June 8, 2011, at the Augusta Civic Center, 16 Cony Street. The informational meeting will be held from 6 p.m. to 7:15 p.m., followed by a public hearing from 7:30 p.m. to 9 p.m.

Peer Review

In accordance with our policy, “Notice of Interagency Cooperative Policy for Peer Review in Endangered Species Act Activities,” which was published on July 1, 1994 (59 FR 34270), we will seek the expert opinion of at least three appropriate independent specialists regarding scientific data and interpretations contained in this proposed rule. The purpose of such review is to ensure that our decisions are based on scientifically sound data, assumptions, and analysis. We will send copies of this proposed rule to the peer reviewers immediately following publication in the

Federal Register

.

Background

National Overview

Below we provide an overview of our proposed national approach to recovery of wolves in the conterminous United States and Mexico. This overview provides the context for our proposed actions for wolves in the eastern United States. In this overview, we discuss the listing history for the gray wolf, evaluate the current gray wolf listing, present the structured decision-making process we have used to date to formulate our national wolf strategy, and describe the strategy itself.

Gray Wolf Listing History

Here we present a brief overview of previous Federal actions relating to the listing of gray wolves and the recovery plans that have been developed pursuant to these listing actions. Additional Federal actions for western Great Lakes wolves are discussed in

Previous Federal Actions for WGL Wolves

below.

Gray wolves were originally listed as subspecies or as regional populations of subspecies in the conterminous United States and Mexico. In 1967, we listed the eastern timber wolf (

Canis lupus lycaon

) in the Great Lakes region (32 FR 4001, March 11, 1967), and in 1973 we listed

C. l. irremotus

in the northern

Rocky Mountains (38 FR 14678, June 4, 1973). Both listings were promulgated under the Endangered Species Conservation Act of 1969; subsequently, on January 4, 1974, these subspecies were listed under the Endangered Species Act of 1973 (39 FR 1171). We listed a third gray wolf subspecies, the Mexican wolf (

C. l. baileyi

) as endangered on April 28, 1976 (41 FR 17740), in the southwestern United States and Mexico. On June 14, 1976 (41 FR 24064), we listed the Texas gray wolf subspecies (

C. l. monstrabilis

) as endangered in Texas and Mexico.

In 1978, we published a rule (43 FR 9607, March 9, 1978) reclassifying the gray wolf as an endangered population at the species level (

C. lupus

) throughout the conterminous 48 States and Mexico, except for the Minnesota gray wolf population, which was classified as threatened. At that time, we considered the Minnesota group of gray wolves to be a listable entity under the Act, and we considered the gray wolf group in Mexico and the 48 conterminous States other than Minnesota to be another listable entity (43 FR 9607, 9610, respectively, March 9, 1978). This reclassification was undertaken because of uncertainty about the taxonomic validity of some of the previously listed subspecies and because we recognized that wolf populations were historically connected, and that subspecies boundaries were thus malleable.

However, the 1978 rule also stated 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, March 9, 1978). Accordingly, recovery plans were developed for the wolf populations in the following regions of the United States: the northern Rocky Mountains in 1980, revised in 1987; the Great Lakes in 1978, revised in 1992; and the Southwest in 1982, the revision of which is now underway.

More detail on previous Federal actions for the Southwest and northern Rocky Mountains wolves is provided, respectively, within the 90-day finding for Mexican wolves (75 FR 46894) and in various notices and rulemakings for the management of northern Rocky Mountains wolves (59 FR 60252, November 22, 1994; 59 FR 60266, November 22, 1994; 68 FR 15804, April 1, 2003; 68 FR 15879, April 1, 2003; 70 FR 1286, January 6, 2005; 71 FR 6634, February 8, 2006; 71 FR 43410, August 1, 2006; 73 FR 4720, January 28, 2008; 73 FR 10514, February 27, 2008; 74 FR 15123, April 2, 2009) . Further detail on previous Federal actions related to the WGL DPS is provided in

Previous Federal Actions for WGL Wolves

below.

Evaluation of the 1978 Gray Wolf Listing

The Service now considers the 1978

Canis lupus

listing rule at 43 FR 9607 to be in need of revision. This need has been identified based on our review of the best available taxonomic information, which indicates that

C. lupus

historically did not occupy large portions of the eastern United States and on our reconsideration of the listing in light of current statutory and policy requirements under the Act. These considerations are discussed in turn below.

Taxonomy and Historical Ranges of Wolves in the United States

Our review of the best available taxonomic information indicates that

Canis lupus

did not occupy large portions of the eastern United States:

i.e.,

the northeastern United States was occupied by the eastern wolf (

C. lycaon

), now considered a separate species of

Canis

rather than a subspecies of

lupus,

and the southeastern United States was occupied by the red wolf (

Canis rufus

) rather than the gray wolf. Our review of North American wolf taxonomy also suggests that changes in listing classification are warranted in other portions of the country.

At the time the gray wolf was listed in 1978, and until the molecular genetics studies of the last few years, the range of the gray wolf prior to European settlement was generally believed to include most of North America. The only areas that were believed to have lacked gray wolf populations were the coastal and interior portions of California, the arid deserts and mountaintops of the western United States, and parts of the eastern and southeastern United States (Young and Goldman 1944, Hall 1981, Mech 1974, and Nowak 1995). We note, however, that some authorities have questioned the reported historical absence of gray wolves in parts of California (Carbyn

in litt.

2000, Mech

in litt.

2000).

Furthermore, we note long-held differences of opinion regarding the precise boundary of the gray wolf's historical range in the eastern and southeastern United States. Some researchers regarded Georgia's southeastern corner as the southern extent of gray wolf range (Young and Goldman 1944, Mech 1974); others believed gray wolves did not extend into the Southeast at all (Hall 1981) or did so to a limited extent, primarily at somewhat higher elevations (Nowak 1995). The southeastern and mid-Atlantic States were generally recognized as being within the historical range of the red wolf (

Canis rufus

), and it is not known how much range overlap historically occurred between the two

Canis

species. Morphological work by Nowak (2000, 2002, 2003) supported extending the historical range of the red wolf into southern New England or even farther northward, indicating either that the historical range of the gray wolf in the eastern United States was more limited than previously believed, or that the respective ranges of several wolf species expanded and contracted in the eastern and northeastern United States, intermingling in post-glacial times along contact zones.

The results of recent molecular genetic analyses (

e.g.,

Wilson

et al.

2000, Wilson

et al.

2003, Wheeldon and White 2009, Wilson

et al.

2009, Fain

et al.

2010, Wheeldon

et al.

2010) and morphometric studies (

e.g.,

Nowak 1995, 2000, 2002, 2003) explain some of the past difficulties in establishing the gray wolf's range in the eastern United States. These studies show that the mid-Atlantic and southeastern United States historically were occupied by the red wolf (

C. rufus

), and that New England and portions of the upper Midwest (eastern and western Great Lakes regions) historically were occupied by

C. lycaon;

they also indicate that the gray wolf (

C. lupus

) did not occur in the eastern United States.

Based on these recent studies, we view the historical range of the gray wolf as the central and western United States, including portions of the western Great Lakes region, the Great Plains, portions of the Rocky Mountains, the Intermountain West, the Pacific Northwest, and portions of the Southwest. All or parts of 29 southern and eastern States (Maine, Massachusetts, Connecticut, New Hampshire, Rhode Island, Vermont, New York, New Jersey, Pennsylvania, Delaware, Maryland, Virginia, North Carolina, South Carolina, Georgia, Florida, Ohio (the part outside WGL DPS), West Virginia, Kentucky, Tennessee, Alabama, Mississippi, Louisiana, Texas (east of Interstate Highway 35), Oklahoma (east of Interstate Highway 35 and southeast of Interstate Highway 44 north of Oklahoma City), Arkansas, Missouri (southeast of Interstate Highway 44 and southeast of Interstate Highway 70 east of St. Louis), Indiana (the part outside WGL DPS), and Illinois (the part outside WGL DPS)) were not within the gray wolf's historical range.

In sum, we now recognize three wolf species with ranges in the conterminous United States:

Canis lupus, Canis lycaon,

and

Canis rufus.

The ranges of

C. lupus

and

C. lycaon

overlap in the western Great Lakes region, as discussed in

Taxonomy of Wolves in the Western Great Lakes Region

below; however, in the eastern United States, the historical range of

C. lupus

is considered to fall outside the historical ranges of

C. lycaon

and

C. rufus.

Conformance With the Act's Definition of Species

Given the assurances we provided in the 1978

C. lupus

listing that we would continue to treat gray wolf subspecies as separate entities for conservation purposes (as noted in Gray Wolf Listing History, above), we identified a need to reconsider the listing in light of current statutory and policy standards regarding the Act's definition of species. The Act provides for listing at various taxonomic and subtaxonomic levels through its definition of “species” in section 3(16): The term species includes any

subspecies

of fish or wildlife or plants, and any

distinct population segment

of any species of vertebrate fish or wildlife which interbreeds when mature (16 U.S.C. 1532(16) (italics added). As a matter of procedure, then, the Service determines whether it is most appropriate to list an entity as a full species, a subspecies, or a DPS of either a species or subspecies. The gray wolf has a Holarctic range; the current listing encompasses the United States-Mexico segment of the population and consists, in turn, of multiple entities.

The specific provision for listing distinct population segments of vertebrates was enacted through the 1978 Amendments to the Act (Pub. L. 95-362, November 10, 1978); these amendments replaced the ability to list “populations” with the ability to list “distinct population segments” and treat them as species under the Act. To interpret and implement the 1978 DPS amendment, the Service and the National Marine Fisheries Service jointly published the Policy Regarding the Recognition of Distinct Vertebrate Population Segments Under the Endangered Species Act (DPS policy) (61 FR 4722, February 7, 1996), setting policy standards for designating populations as “distinct.”

The March 1978 gray wolf listing predated the November 1978 amendments to the Act. Although the 1978 rule lists two

C. lupus

entities,

i.e.,

the endangered and threatened entities described above, these listings were not predicated upon a formal DPS analysis and do not comport with current policy standards. Nonetheless, subsequent recovery plans and all gray wolf rulemakings since 1996 have focused on units reflective of the evident intent of the 1978 rule to manage and recover gray wolves as “separate entities” (43 FR 9609),

i.e.,

subspecies or populations. This proposed rule and our proposed National Wolf Strategy, below, constitute an effort to bring the 1978 listing in line, insofar as possible, with the Act's requirements and current policy standards.

Structured Decision-Making for Wolves

In 2008, the Service embarked on a structured decision-making process as a means of developing a more integrated and comprehensive strategy for gray wolf conservation in the lower 48 States and Mexico. The overall intent of the process was to identify appropriate wolf entities (

i.e.,

listing units) for full status review, anticipating that such review would lead to either confirmation or revision of the existing gray wolf listing. We aimed to identify a coherent set of listing units based on best available scientific and commercial information, conformance with existing regulatory and policy requirements, and fundamental wolf management objectives.

We first conducted several iterations of the process in an internal Service effort to develop a viable framework for considering the scientific and policy questions that drive decision-making for wolves. The resulting framework incorporated decision analysis principles and techniques for crafting alternative listing units and then assessing the relative performance of each alternative in terms of achieving management objectives.

Management of wolves is shared among the Service, States, and Tribes. Thus, following our development of a satisfactory decision-making framework, representatives from several States involved with gray wolf conservation joined us to further explore alternative units that could qualify for future status review (Tribal representatives declined to participate). After acquainting state participants with the decision-making framework, we convened a State-Federal workshop in August 2010 to generate and assess alternative taxonomic and population units at various scales and in various configurations, including the 1978 listing as the

status quo

alternative.

Workshop participants also explored the different values that drive wolf decision-making; these values were expressed as the following fundamental management objectives: (1) Promote and sustain wolf recovery; (2) comply with the requirements of the Act; (3) minimize the regulatory burden on States, Tribes, and the general public; (4) facilitate State and Tribal management of wolves; (5) minimize wolf-human conflicts; and (6) promote public acceptance of wolf listing and recovery actions.

Workshop outcomes provided important input to our continuing effort to formulate a comprehensive vision of wolf conservation. Based on further Service deliberations, this comprehensive vision has evolved into the proposed national wolf strategy discussed below. It is important to note that this strategy is a broad outline, the components of which are in various stages of execution.

National Wolf Strategy

The Service's national wolf strategy is 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.

The strategy is based on three precepts. First, 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. Second, the strategy promotes the continued representation in this country of all substantially unique genetic lineages found historically in the lower 48 States. Third, wolf conservation under the Act is concerned with reducing extinction risks to imperiled entities; the strategy thus focuses on conservation of the four extant gray wolf entities identified through the structured decision-making process and being considered for section 4 actions: (1) The western Great Lakes population, (2) the northern Rocky Mountains (NRM) population, (3) gray wolves in the Pacific Northwest, and (4) the Southwestern population of Mexican wolves.

Various reviews and listing actions are underway for these gray wolf populations. The WGL DPS is proposed for delisting in the proposed rule being published in today's

Federal Register

. With regard to the NRM gray wolf population, Congress is considering legislation that would direct us to reissue our 2009 final rule (74 FR 15123, April 2, 2009), that delisted the NRM DPS in the States of Idaho and Montana,

and in portions of Oregon, Washington, and Utah. This rule retained ESA protections of wolves in Wyoming as non-essential experimental. If passed, we would publish a separate notice in the

Federal Register

. Negotiations regarding potential future post-delisting wolf management in Wyoming are ongoing.

The biological and conservation status of wolves in the Pacific Northwest (we are considering this to be the area west of the NRM gray wolf population, including portions of Oregon, Washington, northern California, and western Nevada) is being assessed to determine their appropriate listing classification. When this review is completed, we will evaluate a potential Pacific Northwest DPS in accordance with our DPS policy and will reclassify this population as appropriate through an additional rulemaking process. The status of the Southwestern population (

i.e.,

Mexican wolves within their historical range) is being reviewed pursuant to our 90-day finding on two listing petitions (75 FR 46894, August 4, 2010). We anticipate that the Southwestern population will be proposed for listing as either the subspecies

C. l. baileyi

or as a DPS of

C. lupus;

in the meantime, recovery planning will continue to proceed for these wolves.

As separate actions move forward for the NRM, Pacific Northwest, and Southwest, wolves in these regions will retain their current classification as endangered, except where delisted and where currently listed as non-essential experimental populations (see 50 CFR 17.84(k)). We plan to move forward with a rulemaking to replace the remainder of the 1978 listing with more targeted regional units, as appropriate, concurrently with publication of the final rule for the WGL DPS.

It is likely that revision of the 1978 gray wolf listing into finer-scale taxonomic or population units will result in removal of the Act's protections in areas of the historical

C. lupus

range, such as the Great Plains States and areas of the western States, that do not support extant wolf populations

and

do not play a role in the recovery of any of the four gray wolf entities. Although some of these areas are within the species' historical range, these areas lack sufficient suitable habitat for wolf pack persistence. Thus, we believe recovery in these areas is both unrealistic and unnecessary. We note, however, that such areas would not necessarily be precluded from wolf conservation efforts under other authorities,

e.g.,

Tribes, States, and Federal land management agencies.

Our national wolf strategy also addresses the two other wolf taxa that fall within the range described for

Canis lupus

in the 1978 listing,

C. lycaon

and

C. rufus.

With regard to

Canis lycaon,

we are announcing a rangewide status review of this species, which occurs in Canada and the western Great Lakes region of the United States. The historical range of

C. lycaon

also extends into the northeastern United States, which the 1978 listing inaccurately treated as part of the range of

C. lupus.

The role of the Northeast region in conservation of

C. lycaon

will be considered in the rangewide review, which will look at the status of extant populations in terms of uniqueness, demography, and extinction risks. A determination as to whether to proceed with any

C. lycaon

listing action—and, if listing is warranted, whether or not to include the northeastern United States in the listed range—will depend on the results of the status review. Notification of our intentions with regard to

C. lycaon

will be provided in conjunction with publication of the final rule for the WGL DPS. Meanwhile, we propose to revise the range of the gray wolf (the species

C. lupus

) by removing all or parts of 29 eastern states that we now recognize were not part of the historical range of the gray wolf. New information indicates that these areas should not have been included in the original listing of the gray wolf. These States are specified under

Taxonomy and Historical Ranges of Wolves in the United States,

above.

Finally, with regard to

Canis rufus,

we propose to remove the southeastern states included in the 1978 gray wolf listing from the List due to error, because we now recognize were not part of the historical range of the gray wolf. These states instead constitute the range of

Canis rufus;

see

Taxonomy and Historical Ranges of Wolves in the United States,

above. Red wolves currently are listed as endangered where found (32 FR 4001, March 11, 1967); this listing will be retained and recovery efforts for red wolves will continue as warranted (Red Wolf Recovery and Species Survival Plan; Service 1990).

Five-Year Review of Gray Wolves

Under section 4(c)(2) of the Act, we have a duty to review listed species' status every 5 years and determine whether a change in listing status is appropriate. We announce initiation of the 5-year review for the gray wolf in this rule and seek new information as requested in Public Comments above.

Western Great Lakes Wolves

Previous Federal Actions for WGL Wolves

The eastern timber wolf (

Canis lupus lycaon

) was listed as endangered in Minnesota and Michigan in the first list of species that were protected under the 1973 Act, published in May 1974 (USDI 1974). On March 9, 1978, we published a rule (43 FR 9607) reclassifying the gray wolf at the species level (

Canis lupus

) as endangered throughout the conterminous 48 States and Mexico, except for the Minnesota population, which we classified to threatened. The separate subspecies listings, including

C. l. lycaon,

thus were subsumed into the listings for the gray wolf in Minnesota and the gray wolf in the rest of the conterminous United States and Mexico. In that 1978 rule, we also identified Isle Royale National Park, Michigan, and Minnesota wolf management zones 1, 2, and 3, as critical habitat. We also promulgated special regulations under section 4(d) of the Act for operating a wolf management program in Minnesota at that time. The depredation control portion of the special regulation was later modified (50 FR 50793; December 12, 1985); these special regulations are found in 50 CFR 17.40(d)(2).

On April 1, 2003, we published a final rule revising the listing status of the gray wolf across most of the conterminous United States (68 FR 15804). Within that rule, we identified three DPSs for the gray wolf (see Gray Wolf Listing History, above), including an Eastern DPS, which was reclassified from endangered to threatened, except where already classified as threatened. In addition, we established a second section 4(d) rule that applied provisions similar to those previously in effect in Minnesota to most of the Eastern DPS. The special rule was codified in 50 CFR 17.40(o).

U.S. District Court rulings in Oregon and Vermont on January 31, 2005, and August 19, 2005, respectively, invalidated the April 1, 2003, final rule. Consequently, the status of gray wolves outside of Minnesota reverted back to endangered status, as had been the case prior to the 2003 reclassification. The courts also invalidated the three DPSs identified in the April 1, 2003, rule, as well as the associated special regulations.

On March 27, 2006, we published a proposal (71 FR 15266-15305) to identify a WGL DPS of the gray wolf, to remove the WGL DPS from the protections of the Act, to remove designated critical habitat for the gray wolf in Minnesota and Michigan, and to

remove special regulations for the gray wolf in Minnesota. The proposal was followed by a 90-day comment period, during which we held four public hearings on the proposal.

On February 8, 2007, the Service issued a rule that identified and delisted the WGL DPS of the gray wolf (

Canis lupus

) (72 FR 6052). Three parties challenged this rule (

Humane Society of the United States

v.

Kempthorne,

579 F. Supp. 2d 7 (D.D.C. 2008)), and on September 29, 2008, the court ruled in favor of the plaintiffs and vacated the rule and remanded it to the Service. On remand, the Service was directed to provide an explanation as to how simultaneously identifying and delisting a DPS is consistent with the Act's text, structure, policy objectives, legislative history, and any relevant judicial interpretations.

The court's primary question was whether the Service has the authority to identify a DPS within a larger already-listed entity and, in the same decision, determine the DPS does not warrant the Act's protections even though the other populations of the species retain the old listing status. Our authority to make these determinations and to revise the list accordingly is a reasonable interpretation of the language of the Act, and our ability to do so is an important component of the Service's program for the conservation of threatened and endangered species.

Our authority to revise the existing listing of a species (the gray wolf in Minnesota and the gray wolf in the lower 48 States and Mexico, excluding Minnesota) to identify a Western Great Lakes DPS and determine that it is healthy enough that it no longer needs the Act's protections is found in the precise language of the Act. Moreover, even if that authority were not clear, our interpretation of this authority to make determinations under section 4(a)(1) and to revise the endangered and threatened species list to reflect those determinations under section 4(c)(1) is reasonable and fully consistent with the Act's text, structure, legislative history, relevant judicial interpretations, and policy objectives.

We consulted with the Solicitor of the Department of the Interior to address the issue in the court's opinion. On December 12, 2008, a formal opinion was issued by the Solicitor, “U.S. Fish and Wildlife Service Authority Under Section 4(c)(1) of the Endangered Species Act to Revise Lists of Endangered and Threatened Species to `Reflect Recent Determinations'” (U.S. DOI 2008). The Service fully agrees with the analysis and conclusions set out in the Solicitor's opinion. This proposed action is consistent with the opinion. The complete text of the Solicitor's opinion can be found at

http://www.fws.gov/midwest/wolf/.

On December 11, 2008, we published a notice reinstating protections for the gray wolf in the western Great Lakes (and northern Rocky Mountains) pursuant to court orders (73 FR 75356).

On April 2, 2009, we published a final rule identifying the western Great Lakes populations of gray wolves as a DPS and revising the list of Endangered and Threatened Wildlife by removing the DPS from that list (74 FR 15070). We did not seek additional public comment on the 2009 final rule. On June 15, 2009, five parties filed a complaint against the Department and the Service alleging that we violated the Act, the Administrative Procedure Act (APA), and the court's remand order by publishing the 2009 final rule (74 FR 15070). On July 2, 2009, pursuant to a settlement agreement between the parties, the court issued an order remanding and vacating the 2009 final rule.

On March 1, 2000, we received a petition from Mr. Lawrence Krak of Gilman, Wisconsin, and on June 28, 2000, we received a petition from the Minnesota Conservation Federation. Mr. Krak's petition requested the delisting of gray wolves in Minnesota, Wisconsin, and Michigan. The Minnesota Conservation Federation requested the delisting of gray wolves in a Western Great Lakes DPS. Because the data reviews resulting from the processing of these petitions would be a subset of the review begun by our July 13, 2000, proposal (65 FR 43450) to revise the current listing of the wolf across most of the conterminous United States, we did not initiate separate reviews in response to those two petitions. While we addressed these petitions in our February 8, 2007, final rule (72 FR 6052), this rule was vacated by the subsequent District Court ruling. While we view our actions on these petitions as final upon publication of the

Federal Register

determinations, we nevertheless restate our 90-day findings that the action requested by each of the petitions may be warranted, as well as our 12-month finding that the action requested by each petition is warranted.

On March 15, 2010, we received a petition from the Minnesota Department of Natural Resources requesting that the gray wolf in Minnesota be removed from the List of Endangered or Threatened Wildlife under the Act. Likewise, on April 26, 2010, we received a petition from the Wisconsin Department of Natural Resources requesting that the gray wolf in Minnesota and Wisconsin be delisted. On April 26, 2010, we received a petition from the Sportsmen's Alliance, representing five other organizations, requesting that gray wolves in the Great Lakes area be delisted. On June 17, 2010, we received a petition from Safari Club International, Safari Club International Foundation and the National Rifle Association of America requesting that wolves of the western Great Lakes be delisted. In response to those four petitions, on September 14, 2010, we published a 90-day finding determining that the petitions presented substantial information that delisting may be warranted and reinitiated a full status review. Therefore, this delisting proposal constitutes our 12-month finding that the action requested by each petition is warranted.

In response to a separate petition, on June 10, 2010, we made a 90-day finding that there was no evidence of any breeding population of wolves to support the requested listing of a DPS of the gray wolf in New England (75 FR 32869).

Species Concepts

As noted in Conformance with the Act's Definition of Species above, the Act defines “species” as including any species or subspecies of fish or wildlife or plants, and any distinct vertebrate population segment of fish or wildlife that interbreeds when mature (16 U.S.C. 1532(16)). It has not been uncommon in the years since the Act was passed for significant controversy to arise over the propriety of recognizing various groups of organisms as eligible for protection under the Act. Our implementing regulations (50 CFR 424.11) require us to use standard taxonomic distinctions (such as species and subspecies) when they are available, clearly defined, and generally accepted. In determining that a taxonomic entity qualifies as a species or subspecies we carefully evaluate the best available taxonomic data to determine whether we have sufficient information to conclude that a taxonomic entity qualifies as a species under the Act.

In identifying species, there is not a single set of criteria, and, therefore, no single species concept that is accepted by all taxonomists. In 1942, Ernst Mayr identified five different species concepts (Mayr 1942), and many more have been recognized since then (Wilkins 2006; 2003; Mayden 1997, pp. 381-384). Many of these species concepts can be associated with one of two major classes of concepts or approaches. The first is the biological species concept (BSC). This concept is based on reproductive relationships among populations. The

ability to interbreed and realize gene flow between two populations is the indication that they belong to the same species. The concept is most commonly associated with Mayr (1963), but has antecedents during the development of evolutionary biology in the 20th century. The second major class of concepts is the phylogenetic species concept (PSC). Under this group of concepts, species are identified by their genealogical (lineages) or phylogenetic (evolutionary) relationships and diagnosability. The many variations of these concepts and others are reviewed by Wiley (1981), Avise (2004), and Coyne and Orr (2004).

There is, likewise, no scientific consensus on what constitutes a subspecies, and some authorities (Wilson and Brown 1953) have questioned the utility of the subspecies level of classification. Following is a description of various subspecies criteria that have been proposed and applied in the taxonomic literature. Because some criteria are more stringent than others, a putative, or generally accepted, subspecies may meet the criteria and be recognized following one concept, but found to be invalid under a more stringent concept. Nowak (1995, p. 394) discussed the standards he used when he revised the subspecies of

Canis lupus:

“My investigation largely disregarded such questions [concerning use of very localized characters] and concentrated on general trends in measurable size and proportion that could be evaluated on a continent-wide or worldwide basis. Substantive statistical breaks in such trends, as discussed above, were taken as evidence of taxonomic division.” In The Mammals of North America, Hall (1981, p. viii) included the following in his “Criteria for Species versus Subspecies.”

If crossbreeding occurs in nature at a place or places where the geographic ranges of two kinds of mammals meet, the two kinds are to be treated as subspecies of one species. If no crossbreeding occurs, the two kinds are to be regarded as two distinct, full species.

Mayr (1963, glossary) defined subspecies as, “an aggregate of local populations of a species inhabiting a geographic subdivision of the range of the species, and differing taxonomically from other populations of the species.” He further explains “differing taxonomically” as differing “by diagnostic morphological characters” (Mayr 1963, p. 348). Mayr (1969, p. 190) also describes a quantitative method for determining whether populations differ taxonomically: “A so-called 75-percent rule is widely adopted. According to this, a population is recognized as a valid subspecies if 75 percent of the individuals differ from “all” (97 percent) of the individuals of a previously recognized subspecies. At the point of intersection between the two curves where this is true, about 90 percent of population A will be different from about 90 percent of the individuals of population B (to supply a symmetrical solution)”.

Patten and Unitt (2002, p. 27) provide another definition of subspecies as “diagnosable clusters of populations of biological species occupying distinct geographic ranges.” They do not require that diagnosability be absolute, but advocate 90 percent separation as a more stringent criterion than the 75-percent rule.

Avise (2004, p. 362) attempted to incorporate phylogenetic information within a biological species concept in providing the following guidance on recognizing subspecies: “Within such units [=species], “subspecies” warranting formal recognition could then be conceptualized as groups of actually or potentially interbreeding populations (normally mostly allopatric) that are genealogically highly distinctive from, but reproductively compatible with, other such groups. Importantly, the empirical evidence for genealogical distinction must come, in principle, from concordant genetic partitions across multiple, independent, genetically based molecular (or phenotypic; Wilson and Brown 1953) traits.”

A common feature of all of the above definitions is that they recognize that subspecies are groups of populations, and most recognize that subspecies can be variable and overlap, to some degree, in distinguishing characters. Taxonomists do not assign an individual to one subspecies or another; instead individuals are assigned a specific taxonomic classification based on the population in which they exist.

The existence of multiple concepts of species and subspecies is not the only complicating factor in the debate surrounding the classification of organisms; it is further complicated by the way organisms occur in the natural world. Taxonomists are determined to categorize natural organisms into specific groups and identify and name those groups, while also striving to understand the evolutionary processes that give rise to these specific groups (Hey 2001, pp. 328-329). When viewed on the ground, a particular organism may appear to clearly fit into one group or another, but when their evolutionary history is viewed, these groups are revealed as changeable and without clear boundaries. In the reverse, individuals may appear different (that is be morphometrically distinct) but in fact be of the same taxon (that is, genetically similar). In many situations, it is difficult to determine where one species ends and another begins. This is especially true in wide-ranging species and in the zones where multiple forms (for example, where either two species or two subspecies) contact each other or meet, which is the situation with wolves in the WGL region. Ultimately, species are evolving, dynamic populations, and at times are difficult to categorize. Nevertheless, Congress directs that the Service classify populations as species, subspecies, and DPSs, despite the difficulty and complexity of various taxonomic concepts.

Taxonomy of Wolves in the Western Great Lakes Region

The taxonomic status of the wolves in the western Great Lakes region has long been debated. They have been considered a subspecies of gray wolf,

Canis lupus lycaon

(Goldman 1944), Nowak 1995, 2002, 2003); a

Canis lupus

population that has been influenced by interbreeding with coyotes (Lehman

et al.

1991); members of a full species,

Canis lycaon

(or eastern wolf) that is separate from

Canis lupus

(Wilson

et al.

2000, Baker

et al.

2003); possibly the same species as the red wolf,

C. rufus

(Wilson

et al.

2000); the result of hybridization between

C. rufus

and

C. lupus

(Nowak 2002, 2003, 2009); and, most recently, as a mixed population of

C. lupus, C. lycaon,

and their intercrosses (for example, Wheeldon and White 2009, Fain

et al.

2010, Wheeldon

et al.

2010). These varying interpretations of the taxonomic status of western Great Lakes wolves are summarized, respectively, below.

Wolves in Michigan, Wisconsin, and eastern Minnesota were considered by Goldman (1944, p. 437 and Figure 14) to be within the range of the subspecies

Canis lupus lycaon.

Goldman based his classification on variation in body size and proportions, and in pelage (coat) color. According to Goldman, this was the subspecies of gray wolf historically found across a wide range east of the Mississippi River in the United States and in southeastern Canada. Wolves immediately to the west of the Mississippi River were considered to be part of the subspecies

Canis lupus nubilus.

This taxonomic interpretation was followed by Hall and Kelson (1959, p. 849) and Hall (1981, p. 932).

Nowak's (1995, p. 396; 2003, p. 243) revision of the subspecies taxonomy reduced the range of

C. l. lycaon

to southern Ontario and Quebec and northern portions of New York,

Pennsylvania, and Ohio. Nowak's classification was primarily based on statistical analysis of measurements of skull features. He considered gray wolves that historically occupied Michigan, Wisconsin, and Minnesota to be within the range of

C. l. nubilus.

Based on analysis of additional specimens, Nowak (2002, p. 119; 2003; 2009, p. 238) continued to recognize western Great Lakes wolves as

C. l. nubilus,

but noted that historical specimens from the Upper Peninsula (UP) of Michigan were somewhat transitional between the two subspecies.

Based on a study of DNA variation in North American wolves, Wilson

et al.

(2000, p. 2165) proposed that the taxonomic standing of eastern wolves be restored to full species as

Canis lycaon.

They found that eastern wolves were divergent from

Canis lupus

in both mitochondrial DNA (mtDNA) and autosomal microsatellite DNA composition. They considered the geographic range of

C. lycaon

as extending west across the Great Lakes region to Minnesota and Manitoba.

Leonard and Wayne (2008, pp. 2-3) have reported on maternally inherited mtDNA sequence haplotypes (DNA sequences or groups of alleles of different genes on a single chromosome that are inherited together as a single unit) from historical (“prerecovery”) wolves from Ontario, Quebec, Michigan, and Wisconsin compared with the recent population of the area. Their interpretation of these results is that the 6 unique haplotypes) identified in 15 historical individuals indicate that the pre-recovery population was “an endemic American wolf,” which they call “the Great Lakes wolf” (p. 1). However, only the two haplotypes most common in the historical sample still occur in the modern wolf population of the western Great Lakes area. Leonard and Wayne (2007) conclude that the modern population does not contain the diversity of Great Lakes wolf haplotypes found in the prerecovery population and that the current population is primarily a mixture of

Canis lupus

and coyote hybrids, with minor influence from the endemic Great Lakes wolf (p. 3).

Koblmüller

et al.

(2009) examined wolves from the western Great Lakes region using three types of genetic markers: mtDNA; Y-chromosome haplotypes based on microsatellite DNA loci on the Y-chromosome, which is a paternally-inherited marker; and autosomal microsatellite DNA, which provides information on recent and ongoing interactions among populations rather than evolutionary lineage information. The historical sample from Minnesota was found to exhibit a third Great Lakes wolf mtDNA haplotype that is common in the modern population. However, the Y-chromosome haplotypes identified in the historical sample were more similar to those of western gray wolves, suggesting that interbreeding between Great Lakes wolves and western gray wolves had taken place before 1910, the year of collection.

Koblmüller

et al.

(2009) conclude that, despite what they consider both ancient and recent incidences of interbreeding with coyotes and western gray wolves, Great Lakes wolves remain morphologically distinct and represent a “distinct taxon” of gray wolf (

Canis lupus

) that is adapted to the region. They do not, however, conclude that this taxon is differentiated enough to be recognized as a species separate from gray wolves, as proposed by Wilson

et al.

(2000).

Several recent studies conclude that the eastern wolf is a unique species and should be recognized as

C. lycaon

(Wheeldon and White 2009; Wilson

et al.

2009; Fain

et al.

2010, p. 15; Wheeldon

et al.

2010). Wheeldon and White (2009, pp. 3-4) state that both the present-day and pre-recovery wolf populations in the western Great Lakes region are genetically similar and that both were derived from hybridization between

C. lupus

and the eastern wolf,

C. lycaon.

Fain

et al.

(2010, p. 10) recognize

C. lycaon

as a unique species of North American wolf, and based on mtDNA and Y-chromosome haplotypes and autosomal microsatellite markers, they establish that the population of wolves in the western Great Lakes region comprise

C. lupus, C. lycaon,

and their hybrids. Contrary to Koblmüller

et al.

(2009), Fain

et al.

(2010, p. 14) found no evidence of interbreeding with coyotes. Furthermore, they conclude that the western Great Lakes States were included in the historical range of

C. lycaon

and that hybridization between the two species “predates significant human intervention” (Fain

et al.

2010, pp. 13-14).

Wheeldon

et al.

(2010, p. 2) used multiple genetic markers to clarify the taxonomic status of

Canis

species in the western Great Lakes region of Minnesota, Wisconsin, Michigan, and western Ontario. They conclude that the current western Great Lakes wolf population is “composed of gray-eastern wolf hybrids that probably resulted from historic hybridization between the parental species” (Wheeldon

et al.

2010, p. 10), and that the appropriate taxonomic designation for the western Great Lakes hybrid wolves is

C. lupus × lycaon,

replacing Nowak's (2009) wolf subspecies designation of

C. lupus lycaon.

We note, however, that a name in the form of

C. lupus × lycaon

has no standing as an available species name under the rules of zoological nomenclature (ICZN 1999).

It is clear from the studies discussed above that the taxonomic classification of wolves in the western Great Lakes region is one that has been, and will continue to be, of great debate in the scientific community. Most researchers, however, appear to agree that there is a unique and genetically identifiable form of wolf that occupies the western Great Lakes region, and that this form has hybridized with

Canis lupus,

whose origins were from elsewhere in North America. Researchers differ in whether this unique form of wolf should be recognized as a species (Wilson

et al.

2000; Fain

et al.

2010, p. 15; Wheeldon

et al.

2010), a subspecies (Nowak 1995), or a distinct taxon or ecotype but without applying a formal scientific name to that form (Koblmüller

et al.

2009). In choosing among these three alternatives, we find that the large divergence of both mtDNA and Y-chromosome haplotypes between Great Lakes wolves and

C. lupus

is greater than that found between subspecies of

Canis lupus

and favors recognition of the eastern wolf as a species. Currently, the best available scientific information supports recognition of the eastern wolf,

C. lycaon,

as a species (rather than, as previous believed, as a subspecies of gray wolf), and establishes that this species has intercrossed with

C. lupus

in the western Great Lakes region to constitute a population composed of

C. lupus, C. lycaon,

and their hybrids (Wheeldon and White 2009, p. 1; Fain

et al.

2010, p. 14; Mech

et al.

2010; Wheeldon

et al.

2010).

The existence of two wolf species in the western Great Lakes region was not known or suspected in 1978, when the Service replaced the listings of four subspecies of gray wolf, including

C. lupus lycaon,

with the listing of all

Canis lupus

and

Canis lupus

subspecies in the conterminous United States and Mexico as endangered, except for the Minnesota population, which was listed as threatened (USFWS 1978). Since that time, increasingly powerful genetic techniques for the characterization of populations have been developed and applied to wild populations, including wolves. These advances have shown that hybridization between species is much more prevalent than was appreciated in 1978 (Schwenk

et al.

2011); thus the detection of hybridization in western Great Lakes wolves is not unique among mammalian species.

Nowak's (1995, 2002, 2003) exclusion of the western Great Lakes region from

C. l. lycaon

was likely influenced by his inclusion of both

C. lupus

and

C. lycaon

in his western Great Lakes sample. In any event, the various genetic investigations of western Great Lakes wolves clearly show a distribution of eastern wolf (

C. lycaon

) genetic markers throughout the region.

We do not accept the proposal of Wilson

et al.

(2000) that

C. lycaon

and

C. rufus

(red wolf) are the same species. Their conclusion was based on red wolf and

C. lycaon

occurring on the same branch of a phylogenetic network representing mtDNA differences (Wilson

et al.

2000, Figure 5A). This relationship has not been found in subsequent studies (Wilson

et al.

2003; Leonard and Wayne 2008, p. 2; Fain

et al.

2010, p. 9), which placed the red wolf and

C. lycaon

on different branches separated by intervening coyote lineages. This suggests that the red wolf and

C. lycaon

may have evolved independently from common ancestors with modern coyotes, but does not support uniting them as a single species.

Genetic Composition of Wolves in the Western Great Lakes Region

Estimates of the genetic composition of the wolves of the western Great Lakes region with respect to the two species (

C. lupus

and

C. lycaon

) are based on the frequencies of different paternal (Y-chromosome) and maternal (mtDNA) markers specific to the each species in samples of wolves from the region. For mtDNA, 66 percent of sampled wolves had

C. lycaon

haplotypes (Fain

et al.

2010, p. 13; Wheeldon

et al.

2010). For Y-chromosome haplotypes, 54 percent (Wheeldon

et al.

2010) or 50 percent (Fain

et al.

2010, p. 7) of sampled wolves had haplotypes of

C. lycaon.

Male wolves carry both paternal and maternal markers. Of male wolves sampled by Fain

et al.

(2010, p. 12), 41 percent had both maternal and paternal haplotypes of

C. lycaon,

and 13 percent had both maternal and paternal haplotypes of

C. lupus.

Based on a larger sample that also included some wolves from western Ontario, Wheeldon

et al.

(2010) reported 42 percent of the sampled male wolves had both maternal and paternal haplotypes of

C. lycaon

and 21 percent had both maternal and paternal haplotypes of

C. lupus.

Maternal and paternal haplotypes were mixed with respect to the two species for the remaining wolves in both studies.

Although it is clear that

C. lycaon

and

C. lupus

have hybridized in the western Great Lakes region, same-species combinations of paternal and maternal markers in male wolves are more common than expected by random mating (Wheeldon

et al.

2010). This suggests that there is some constraint on complete hybridization between the two species and that complete blending of the two components of the population is not inevitable. The limited number of historical specimens from the western Great Lakes region that have been genetically characterized all have mtDNA indicative of

C. lycaon

(Leonard and Wayne 2008, pp. 2-3; Wheeldon and White 2009, p. 1), but four of these from the early 20th century also had

C. lupus

Y-chromosome haplotypes, which indicates that hybridization had occurred by that time. The opportunity for hybridization between

C. lycaon,

which belongs to a North American lineage, and

C. lupus,

which evolved in Eurasia, has existed since

C. lupus

entered North America about 500,000 years ago (Kurtén and Anderson 1980), yet a predominantly

C. lycaon

population of wolves still persists in the western Great Lakes region.

Wolf-Coyote Relationships

For a discussion on interpretations of wolf-coyote relationships in the western Great Lakes, see the discussion under Factor E. Other Natural or Manmade Factors Affecting Its Continued Existence in this proposed rule.

Procedural Aspects of Proposal Applying to the Gray Wolf (C. lupus)

When the Service revised the endangered species list in 1978 to include the species

Canis lupus

in the lower 48 States and Mexico, regulatory protections were applied to all gray wolves in the lower 48 States, including all subspecies of gray wolves, which were subsumed at that time into

C. lupus.

That rule classified the Minnesota gray wolf population as a threatened “species” and gray wolves elsewhere in the lower 48 States and Mexico as another “species” with endangered status. The best scientific information available supports the existence of distinct taxa and populations within the

C. lupus

listing and changes our understanding of North American wolf taxonomy. With regard to the WGL wolf population, current scientific data indicate that

Canis lycaon,

which was understood in 1978 to be a subspecies of

C. lupus,

should be recognized as a full species, and that

C. lycaon

and

C. lupus

both occur, and to some extent, interbreed in the western Great Lakes area (see

Taxonomy of Wolves in the Western Great Lakes Region

).

The existence of this new information does not by itself change the regulatory status of the gray wolf (

C. lupus

) under the Act—such changes must be made through rulemaking. This proposed rule recognizes the taxonomic changes and the improved status of the WGL gray wolf populations and proposes those appropriate and necessary administrative changes for the gray wolf in the WGL and portions of the eastern United States.

Based on our current understanding of wolf systematics, we recognize that not all individual wolves in the WGL region are in fact, gray wolves,

Canis lupus.

Within this rule we are proposing changes to the listing for

C. lupus

and are initiating a status review for

C. lycaon.

These two actions combined will address all wolves in the WGL region.

The procedural aspects of this proposed rule (

e.g.,

the revision of the 1978 listing of the group of gray wolves in Minnesota as a “species” to a DPS and the delisting of that DPS) refer to the gray wolf (

C. lupus

), because that is the named entity currently on the List of Endangered and Threatened Wildlife. Our proposed action here is to establish the existence of a WGL distinct population segment of

C. lupus

and to determine that the DPS is neither endangered nor threatened, despite its proximity to a closely related species,

C. lycaon

—a species whose status we will evaluate for possible protection under the Act in the near future.

Biology and Ecology of Wolves in the Western Great Lakes

Gray wolves are the largest wild members of the Canidae, or dog family, with adults ranging from 18 to 80 kilograms (kg) (40 to 175 pounds (lb)) depending upon sex and subspecies (Mech 1974). The average weight of male wolves in Wisconsin is 35 kg (77 lb) and ranges from 26 to 46 kg (57 to 102 lb), while females average 28 kg (62 lb) and range from 21 to 34 kg (46 to 75 lb) (Wisconsin Department of Natural Resources (WI DNR) 1999). Wolves' fur color is frequently a grizzled gray, but it can vary from pure white to coal black. Wolves may appear similar to coyotes (

Canis latrans

) and some domestic dog breeds (such as the German shepherd or Siberian husky) (

C. lupus familiaris

). Wolves' longer legs, larger feet, wider head and snout, and straight tail distinguish them from both coyotes and dogs.

Wolves primarily are predators of medium and large mammals. Wild prey species in North America include white-tailed deer (

Odocoileus virginianus

) and mule deer (

O. hemionus

), moose (

Alces alces

), elk (

Cervus elaphus

), woodland caribou (

Rangifer caribou

) and barren

ground caribou (

R. arcticus

), bison (

Bison bison

), muskox (

Ovibos moschatus

), bighorn sheep (

Ovis canadensis

) and Dall sheep (

O. dalli

), mountain goat (

Oreamnos americanus

), beaver (

Castor canadensis

), snowshoe hare (

Lepus americanus

), and muskrat (

Ondatra zibethicus

), with small mammals, birds, and large invertebrates sometimes being taken (Chavez and Gese 2005, Mech 1974, Stebler 1944, WI DNR 1999, Huntzinger

et al.

2005). In the WGL DPS, during the last 25 years, wolves have also killed domestic animals including horses (

Equus caballus

), cattle (

Bos taurus

), sheep (

Ovis aries

), goats (

Capra hircus

), llamas (

Lama glama

), pigs (

Sus scrofa

), geese (

Anser sp.

), ducks (

Anas sp.

), turkeys (

Meleagris gallopavo

), chickens (

Gallus sp.

), guinea fowl (

Numida meleagris

), pheasants (

Phasianus colchicus

), dogs, cats (

Felis catus

), and captive white-tailed deer (Paul 2004, 2005; Wydeven 1998; Wydeven

et al.

2001; Wydeven and Wiedenhoeft 1999, 2000, 2001, 2005).

Wolves are social animals, normally living in packs of 2 to 12 wolves. Winter pack size in Michigan's Upper Peninsula (UP) averaged from 2.7 to 4.6 wolves during the 1995 through 2005 period and ranged from 2 to 14 wolves per pack (Huntzinger

et al.

2005). Pack size in Wisconsin is similar, averaging 3.8 to 4.1 wolves per pack, and ranging from 2 to 11 wolves in winter 2004-05 (Wydeven and Wiedenhoeft 2005). In Minnesota the average pack size found in the 1988-89, 1997-98, and 2003-04 winter surveys was higher—5.55, 5.4, and 5.3 wolves per pack, respectively (Erb and Benson 2004).

Packs are primarily family groups consisting of a breeding pair, their pups from the current year, offspring from one or two previous years, and occasionally an unrelated wolf. Packs typically occupy, and defend from other packs and individual wolves, a territory of 20 to 214 square (sq) miles (mi) (50 to 550 sq kilometers (km)). Midwest wolf packs tend to occupy territories on the lower end of this size range. Michigan Upper Peninsula territories averaged 103 sq mi (267 sq km in 2000-01 (Drummer

et al.

2002), Wisconsin territories 37 sq mi (96 sq km) in 2004-05 (Wydeven and Wiedenhoeft 2005), and Minnesota territory size averaged 39 sq mi (102 sq km) in 2003-04 (Erb and Benson 2004). Normally, only the top-ranking (“alpha”) male and female in each pack breed and produce pups. Litters are born from early April into May; they range from 1 to 11 pups, but generally include 4 to 6 pups (Michigan Department of Natural Resources (MI DNR) 1997; USFWS 1992; USFWS

et al.

2001). Normally a pack has a single litter annually, but the production of 2 or 3 litters in one year has been routinely documented in Yellowstone National Park (USFWS

et al.

2002; Smith

et al.

2005).

Yearling wolves frequently disperse from their natal packs, although some remain with their natal pack. Adult wolves and pups older than 5 months also may disperse but at much lower frequencies (Fuller 1989). Dispersers may range over large areas as lone animals after leaving their natal pack or they may locate suitable unoccupied habitat and a member of the opposite sex and begin their own pack. These dispersal movements allow a wolf population to quickly expand and colonize areas of suitable habitat that are nearby or even those that are isolated by a broad area of unsuitable habitat. Additional details on extraterritorial movements are found in

Delineating the Boundaries of the Proposed WGL Gray Wolf DPS,

below.

Recovery of Western Great Lakes Wolves

Recovery Criteria

Recovery plans are not regulatory documents and are instead intended to provide guidance to the Service, States, and other partners on methods of minimizing threats to listed species and achieving recovery. These documents include, among other elements required under section 4(f) of the Act, criteria for determining when a species can be delisted. There are many paths to accomplishing recovery of a species; in fact, recovery of a species is a dynamic process requiring adaptive management that may, or may not, strictly adhere to the guidance provided in a recovery plan.

We use recovery criteria in concert with evidence that threats have been minimized sufficiently and populations have achieved long-term viability to judge when a species can be reclassified from endangered to threatened or delisted. 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) contain the same two recovery criteria. 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.

The Recovery Plan for the Eastern Timber Wolf was based on the best available information on wolf taxonomy at the time of its original publication and subsequent revision. As discussed above in

Taxonomy of Wolves in the Western Great Lakes Region,

since the publication of those plans, several studies have produced conflicting results regarding the taxonomic identity of the wolf that historically occupied the eastern States. Currently, the Service subscribes to the view that what was formerly recognized as the subspecies

C. lupus lycaon

should be recognized as a unique species,

C. lycaon.

Regardless of its taxonomic identity, however, this recovery program has always focused on recovering the wolf population that survived in, and has expanded outward from, northeastern Minnesota. Thus, the Plans guide our analysis of recovery of the wolves in the western Great Lakes area.

Although the recovery criteria identified in the Recovery Plan predate the scientific field of conservation biology, the conservation principles of representation (conserving the genetic diversity of a taxon), resilience (the ability to withstand demographic and environmental variation), and redundancy (sufficient populations to provide a margin of safety) were incorporated into these criteria. Maintenance of the Minnesota wolf population is vital in terms of representation and resilience, because the remaining genetic diversity of wolves in the eastern United States (other than red wolves) was carried by the several hundred wolves that survived in Minnesota into the early 1970s. The Recovery Team insisted that the remnant Minnesota wolf population be maintained and protected to achieve wolf recovery in the eastern United States. The successful growth of the remnant Minnesota population has maintained and maximized the representation of that genetic diversity among wolves in the WGL. 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 would increase the likelihood of maintaining its genetic diversity over the long term. This large Minnesota wolf population also provides resiliency to reduce the adverse impacts of unpredictable demographic and environmental events. Furthermore, the Revised Recovery Plan specifies 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 WGL metapopulation.

The Recovery Plan provides two options for reestablishing this second population. If it is an isolated population, that is, located more than 100 mi (160 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. 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 re-established. 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 UP 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 1992, pp. 56-58). Neither the 1978 nor the 1992 recovery criteria suggest that the restoration of the gray wolf throughout all or most of what was thought to be its historical range in the eastern United States, or to all of these potential re-establishment areas, is necessary to achieve recovery under the Act.

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 UP 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).

Recovery Trends for Wolves in the Western Great Lakes Region

Minnesota Recovery

During the pre-1965 period of wolf bounties and legal public trapping, wolves persisted in the remote northeastern portion of Minnesota but were eliminated from the rest of the State. Estimated numbers of Minnesota wolves before their listing under the Act in 1974 include 450 to 700 wolves in 1950-53 (Fuller

et al.

1992, p. 43, based on data in Stenlund 1955, p. 19), 350 to 700 wolves in 1963 (Cahalane 1964, p. 10), 750 wolves in 1970 (Leirfallom 1970, p. 11), 736 to 950 wolves in 1971-72 (Fuller

et al.

1992, p. 44), and 500 to 1,000 wolves in 1973 (Mech and Rausch 1975, p. 85). Although these estimates were based on different methodologies and are not directly comparable, each puts the pre-listing abundance of wolves in Minnesota at 1,000 or less. This was the only significant wolf population in the United States outside Alaska during those time periods.

After the gray wolf was listed as endangered under the Act in 1974, the Minnesota population estimates increased (see table 1 below). Mech estimated the population to be 1,000 to 1,200 wolves in 1976 (USFWS 1978, pp. 4, 50-52), and Berg and Kuehn (1982, p. 11) estimated that there were 1,235 wolves in 138 packs in the winter of 1978-79. In 1988-89, the Minnesota Department of Natural Resources (MN DNR) repeated the 1978-79 survey and also used a second method to estimate wolf numbers in Minnesota. The resulting independent estimates were 1,500 and 1,750 wolves in at least 233 packs; the lower number was derived by a method comparable to the 1978-79 survey (Fuller

et al.

1992, pp. 50-51).

During the winter of 1997-98, the MN DNR repeated a statewide wolf population and distribution survey, using methods similar to those of the two previous surveys. Field staff of Federal, State, Tribal, and county land management agencies and wood products companies were queried to identify occupied wolf range in Minnesota. Data from 5 concurrent radio telemetry studies tracking 36 packs, representative of the entire Minnesota wolf range, were used to determine average pack size and territory area. Those figures were then used to calculate a statewide estimate of wolf and pack numbers in the occupied range, with single (non-pack) wolves factored into the estimate (Berg and Benson 1999, pp. 1-2).

Table 1—Minimum Winter Wolf Populations in Minnesota, Wisconsin, and Michigan (Excluding Isle Royale) From 1976 Through 2010. (Note That There are Several Years Between the First Three estimates. Minnesota Does Not Conduct Annual Surveys.)

Year

Number of wolves

Minnesota

Wisconsin

Michigan

Wisconsin and Michigan total

1976

1,000-1,200

1978-79

1,235

1988-89

1,500-1,750

31

3

34

1989-90

34

10

44

1990-91

40

17

57

1991-92

45

21

66

1992-93

40

30

70

1993-94

57

57

114

1994-95

83

80

163

1995-96

99

116

215

1996-97

148

113

261

1997-98

2,445

180

139

319

1998-99

205

169

374

1999-2000

248

216

464

2000-01

257

249

506

2001-02

327

278

604

2002-03

335

321

656

2003-04

3,020

373

360

733

2004-05

435

405

840

2005-06

467

434

899

2006-07

546

509

1,055

2007-08

2,921

549

520

1,069

2008-09

637

577

1,214

2009-10

690

557

1,247

The 1997-98 survey concluded that approximately 2,445 wolves existed in about 385 packs in Minnesota during that winter period (90 percent confidence interval from 1,995 to 2,905 wolves) (Berg and Benson 1999, p. 4). This figure indicated the continued growth of the Minnesota wolf population at an average rate of about 3.7 percent annually from 1970 through 1997-98. Between 1979 and 1989 the annual growth rate was approximately 3 percent, and it increased to between 4 and 5 percent in the next decade (Berg and Benson 1999, p. 5; Fuller

et al.

1992, p. 51). As of the 1998 survey, the number of Minnesota wolves had reached approximately twice the number specified in the recovery planning goal for Minnesota (USFWS 1992, p. 28).

Minnesota DNR conducted another survey of the State's wolf population and range during the winter of 2003-04, again using methodology similar to the previous surveys. That survey concluded that an estimated 3,020 wolves in 485 packs occurred in Minnesota (90 percent confidence interval for this estimate is 2,301 to 3,708 wolves) (Erb and Benson 2004, pp. 7, 9). The MN DNR conducted its most recent survey of wolf population and range during the winter of 2007-08. That survey concluded that an estimated 2,921 wolves in 503 packs occurred in Minnesota (90 percent confidence interval for this estimate is 2,192 to 3,525 wolves). The results of the past three surveys suggest that the wolf population has been numerically stable over the past 10 or more years (Erb 2008, p. 6).

As wolves increased in abundance in Minnesota, they also expanded their distribution. During 1948-53, the primary wolf range was estimated at 11,954 sq mi (31,080 sq km) (Stenlund 1955, p. 19). A 1970 questionnaire survey in Minnesota resulted in an estimated wolf range of 14,769 sq mi (38,400 sq km) (calculated by Fuller

et al.

1992, p. 43, from Leirfallom 1970). Fuller

et al.

(1992, p. 44), using data from Berg and Kuehn (1982), estimated that Minnesota primary wolf range encompassed 14,038 sq mi (36,500 sq km) during the winter of 1978-79. By 1982-83, pairs or breeding packs of wolves were estimated to occupy an area of 22,000 sq mi (57,050 sq km) in northern Minnesota (Mech

et al.

1988, p. 86). That study also identified an additional 15,577 sq mi (40,500 sq km) of peripheral range, where habitat appeared suitable but no wolves or only lone wolves existed. The 1988-89 study produced an estimate of 23,165 sq mi (60,200 sq km) as the contiguous wolf range at that time in Minnesota (Fuller

et al.

1992, pp. 48-49; Berg and Benson 1999, p. 3, 5), an increase of 65 percent over the primary range calculated for 1978-79.

The 1997-98 study concluded that the contiguous wolf range had expanded to 33,971 sq mi (88,325 sq km), a 47 percent increase in 9 years (Berg and Benson 1999, p. 5). By that time the Minnesota wolf population was using most of the available primary and peripheral range identified by Mech

et al.

(1988, p. 86). The wolf population in Minnesota had increased in abundance and distribution to the point that its contiguous range covered approximately 40 percent of the State during 1997-98. In contrast, the 2003-04 survey failed to show a continuing expansion of wolf range in Minnesota, and any actual increase in wolf numbers since 1997-98 was attributed to increased wolf density within a stabilized range (Erb and Benson 2004, p. 7). The results of the 2007-08 survey also indicated that wolf range in Minnesota remained “essentially unchanged” since 2004 (Erb 2008, not paginated).

Although the Minnesota DNR does not conduct a formal wolf population survey annually, it includes the species in its annual carnivore track survey. This survey, standardized and operational since 1994, provides an annual index of abundance for several species of large carnivores by counting their tracks along 20-mile (32-km) long standardized survey routes in northern Minnesota. In 2009, wolves were detected on 71 percent of the 58 routes surveyed, and the resulting indices of abundance and distribution were not appreciably different from recent years (Erb 2009, not paginated).

Summary for Minnesota

The Minnesota wolf population has increased from an estimated 1,000 individuals in 1976 to nearly 3,000 today and the estimated wolf range in the State has expanded by approximately 225 percent (from approximately 15,000 sq mi (24,100 sq km) to approximately 34,000 sq mi (54,700 sq km)) since 1970. Over the past 10-12 years, the population size and range have remained stable, as most of the primary and peripheral habitat has been occupied. Based on the current abundance and distribution of the Minnesota wolf population, we believe its continued survival is ensured, and it achieves the first recovery criterion of the Revised Recovery Plan.

Wisconsin Recovery

Wolves were considered to have been extirpated from Wisconsin by 1960. No formal attempts were made to monitor the State's wolf population from 1960 through 1978. Although individual wolves and an occasional wolf pair were reported from 1960 through 1975, (Thiel 1978, Thiel 1993), there was no documentation of wolf reproduction occurring in Wisconsin, and the wolves that were reported may have been dispersing animals from Minnesota.

Wolves are believed to have reestablished breeding packs in Wisconsin in the winter of 1975-76. The Wisconsin Department of Natural Resources (WI DNR) began wolf population monitoring in 1979-80, estimating a statewide population of 25 wolves at that time (Wydeven and Wiedenhoeft 2000, pp. 151, 159; Wydeven

et al.

2009c, pp. 93-97). This population remained relatively stable for several years, and then declined to approximately 15 to 19 wolves in the mid-1980s. In the late 1980s, the Wisconsin wolf population began an increase that has continued into 2010, when 690 wolves were counted (Wydeven

et al.

2010, Figure 3).

Since 1979, WI DNR has intensively surveyed its wolf population on an annual basis using a combination of aerial, ground, and satellite radio telemetry complemented by snow tracking and wolf sign surveys (Wydeven

et al.

2006a, pp. 4-5; Wydeven

et al.

2009c, pp. 90-91). Wolves are trapped from May through September and fitted with radio collars, with a goal of having at least one radio collared wolf in approximately half of the wolf packs in Wisconsin. Aerial locations are obtained from each functioning radio collar about once per week, and pack territories are estimated and mapped from the movements of the individuals who exhibit localized patterns. From December through March, the pilots make special efforts to visually locate and count the individual wolves in each radio-tracked pack.

Snow tracking is used to supplement the information gained from aerial sightings and to provide pack size estimates for packs lacking a radio-collared wolf. Tracking is done by assigning survey blocks to trained trackers, who then drive snow-covered roads in their blocks and follow all wolf tracks they encounter. Snowmobiles are used to locate wolf tracks in more remote areas with few roads. The results of the aerial and ground surveys are carefully compared to properly separate packs and to avoid over-counting (Wydeven

et al.

2006a, pp. 4-5). The estimated number of wolves in each pack is based on the aerial and ground observations made of the individual wolves in each pack over the winter.

Because the monitoring methods focus on wolf packs, lone wolves are likely undercounted in Wisconsin. As a result, the annual population estimates are probably slight underestimates of the actual wolf population within the State during the late-winter period. Fuller (1989, p. 19) noted that lone wolves are estimated to compose from 2 to 29 percent of the total population in the area. Wisconsin DNR surveys have estimated 2-15 percent of the winter population as loners (Wydeven

et al.

2009c, p. 96). These surveys, however, are focused on heavily forested portions of northern and central Wisconsin; therefore, dispersing wolves traveling other portions of the State are less likely to be detected, and often such wolves are only documented after vehicle collisions or accidental shootings. Broader use of trail cameras by members of the public is improving the WI DNR's ability to detect lone wolves across the State.

As previously stated, population estimates are made at the low point of the annual wolf population cycle. Thus, Wisconsin wolf population estimates are conservative in two respects. They undercount lone wolves, and the count is made at the annual low point of the population. This methodology is consistent with the recovery criteria established in the Revised Recovery Plan, which established numerical criteria to be measured with data obtained by late-winter surveys. Based on these considerations, an estimated 690 to 733 wolves in 181 packs, including 35 wolves on Native American reservations, were in Wisconsin in early 2010, representing an 8 percent increase from 2009 (Wydeven

et al.

2010, pp. 12-13).

In the winter of 1994-95, wolves were first documented in Jackson County, Wisconsin, well to the south of the area occupied by other Wisconsin wolf packs in the northern part of the State (Thiel

et al.

2009, pp. 109-110). The number of wolves in this central Wisconsin area has dramatically increased since that time. During the winter of 2009-10, there were 100-106 wolves in 25 packs in the central forest wolf range (Zone 2 in the Wisconsin Wolf Management Plan; Wydeven

et al.

2010, p. 5) and an additional 46 to 48 wolves in 12 or 13 packs in the marginal habitat in Zone 3, located between Zone 1 (northern forest wolf range) and Zones 2 and 4 (Wydeven

et al.

2010, p. 5).

During the winter of 2004-05, 11 to 13 wolves were believed to be primarily occupying Native American reservation lands in Wisconsin (Wydeven in litt. 2005); this increased to 16 to 17 in 2005-06, 17 to 19 in 2007-08 (Wydeven and Wiedenhoeft 2008, Summary), approximately 27 in 2008-2009 (Wydeven and Wiedenhoeft 2008, p. 1), and approximately 35 in 2009-10 (Wydeven

et al.

2010, p. 1). The 2009-10 survey consisted of 3 packs totaling 10-11 wolves on the Bad River Chippewa Reservation and a pack of 2 wolves on the Lac Courtes Oreilles Chippewa Reservation, both in northwestern Wisconsin. There also were two packs of five wolves each on the Lac du Flambeau Reservation in north-central Wisconsin. A pack of four wolves and three pairs occurred on the Menominee Reservation and a three-wolf pack occurred on the Stockbridge Reservation, both in northeastern Wisconsin (Wydeven

et al.

2010, Table 6). A pack of four to five wolves spent time on portions of the Red Cliff Chippewa Reservation along the Lake Superior shoreline. Wolf packs also used scattered lands of the St. Croix Chippewa in northwest Wisconsin, the Ho Chunk Nation in central Wisconsin, and Potawatomi in northeast Wisconsin. The Tribal land of the Ho-Chunk, St. Croix Chippewa, and Potawatomi are composed mostly of scattered parcels of land, and are not likely to provide significant amounts of wolf habitat. About 90 percent of packs in northern Wisconsin Zone 1, and northern portions of Zone 3 are located in ceded territory where Chippewa Bands have retained hunting and gathering rights.

In 2002, wolf numbers in Wisconsin alone surpassed the 1992 Revised Recovery Plan criterion for a second population within 100 miles of the Minnesota population (100 wolves for a minimum of 5 consecutive years (USFWS 1992, p. 4)). Furthermore, in 2004, Wisconsin wolf numbers

exceeded the 1992 recovery criterion of 200 animals for 6 successive late-winter surveys for an isolated wolf population (USFWS 1992, p. 4). Wisconsin population estimates for 1985 to 2010 increased from 15 to 690 wolves (see table 1 above) and from 4 to 181 packs (Wydeven

et al.

2010, figure 3). This represents an annual population increase of 21 percent through 2000, and an average annual increase of 6 percent for the most recent 6 years. The slower rates of increase since 2000 are an indication that the State's wolf population growth and geographic expansion are beginning to level off.

Michigan Recovery

Except for Isle Royale, wolves were extirpated from Michigan as a reproducing species long before they were listed as endangered under the Act in 1974. Prior to 1989, the last known breeding population of wild Michigan wolves outside Isle Royale occurred in the mid-1950s. However, as wolves began to reoccupy northern Wisconsin, the Michigan Department of Natural Resources (MI DNR) began noting single wolves at various locations in the UP of Michigan. Wolf recovery in Michigan began with the documentation of three wolves traveling together and making territorial marks in the central UP during the fall of 1988; and the subsequent birth of pups in this territory during spring 1989 (Beyer

et al.

2009, p. 73). Since that time, wolf packs have spread throughout the UP, with immigration occurring from Wisconsin on the west and possibly from Ontario on the east. Wolves now are found in every county of the UP, with the possible exception of Keweenaw County (Huntzinger

et al.

2005, p. 6; Roell 2009, pers. comm.).

The MI DNR annually monitors the wolf population in the UP by conducting a winter survey. Roads and trails are searched intensively and extensively for wolf tracks and other wolf sign using trucks and snowmobiles (Potvin

et al.

2005). Complete surveys conducted from 1999 to 2006 provided an opportunity to evaluate multiple sampling approaches (MI DNR 2008). Based on these evaluations, it was determined that a geographically stratified sampling protocol produced unbiased, precise estimates of wolf abundance (Potvin

et al.

2005; Drummer, unpublished data). The sampling protocol implemented in 2007 allows trackers to spend more time in smaller areas (MI DNR 2008).

The UP is divided into 21 survey units from which a stratified random sample is drawn, covering roughly 50 percent of the UP every year (MI DNR 2008). Pack locations are derived from previous surveys, citizen reports, and extensive ground and aerial tracking of radio-collared wolves. During the winter of 2009-10, 557 wolves in 109 packs were resident in the UP (MI DNR in litt. 2010, Table 1). Surveys along the border of adjacent survey units are coordinated to avoid double counting of wolves and packs occupying those border areas. In areas with a high density of wolves, ground surveys by four to six surveyors with concurrent aerial tracking are used to accurately delineate territories of adjacent packs and count their members (Beyer

et al.

2004, pp. 2-3; Huntzinger

et al.

2005, pp. 3-6; Potvin

et al.

2005, p. 1661). As with Wisconsin, the Michigan surveys likely miss lone wolves, thus underestimating the actual population.

Based on annual surveys in late winter, estimates of wolves in the UP increased from 57 wolves in 1994 to 557 in late winter 2009-10 (see table 1 above). Over the last 10 years, the annualized rate of increase has been about 12 percent (MI DNR in litt. 2010, table 1). This rate has varied from year to year, but there appear to be two distinct phases of population growth, with relatively rapid growth (25.8 percent average) from 1995 through 2000 and slower growth (10.1 percent average) from 2001 through 2010. In 2005, the number of wolves in the Michigan population alone surpassed the recovery criterion for an isolated wolf population of 200 animals for 6 successive late-winter surveys, as specified in the Revised Recovery Plan (USFWS 1992, pp. 24-26).

To date, no wolf packs are known to be primarily using Tribal-owned lands in Michigan (Roell 2011, pers. comm.). Native American Tribes in the UP of Michigan own small, scattered parcels of land relative to the size of wolf pack territories. Thus, no one Tribal property would likely support a wolf pack. However, as wolves occur in all counties in the UP and are wide-ranging, Tribal land is likely used periodically by wolves.

In October 2004, a coyote trapper mistakenly captured and killed a wolf in Presque Isle County in the northern Lower Peninsula (LP) of Michigan. This was the first verification of a wolf in the northern LP in at least 65 years (Roell

et al.

2010, p. 4). This wolf had been trapped and radio-collared by the MI DNR the previous year (2003) while it was a member of an eastern UP pack. Since 2004, Michigan has surveyed the northern LP to determine whether wolves had successfully colonized the area. From 2005 through 2007, the survey had two components: A prioritized area search and a targeted area search based on citizen reports of wolves or wolf sign. USDA-Wildlife Services, Little Traverse Bay Band of Odawa Indians, and Central Michigan University worked cooperatively on the surveys. Nine units ranging in size from 200-400 sq mi (322-644 sq km) were surveyed; however, no wolf sign was found (Roell

et al.

2010, p. 4). Beginning in 2008, a targeted search approach was used. The MI DNR issued a press release asking citizens to report any wolves or wolf sign; again, no wolves were detected in winters of 2008-10 (Roell

et al.

2009, p. 5; Roell 2010, pers. comm.).

In the summer of 2009, video images of single wolves were recorded in two of the three northern LP counties nearest to the UP (Roell

et al.

2010, p. 4). The videos, taken in Emmet County in May 19, 2009, and Presque Isle County in July 27, 2009, may have been of the same animal (Roell 2009, pers. comm.). In 2010, USDA Wildlife Services and MI DNR staff confirmed a single breeding pair with three pups in Cheboygan County in the northern LP (MI DNR 2010). This is the first time a wolf pack has been verified in the LP since the early 1900s. In 2008, the DNR recognized the likelihood that small numbers of wolves would eventually move into the northern LP and form persistent packs (Potvin 2003, pp. 29-30; Gehring and Potter 2005, p. 1242; Beyer

et al.

2006, p. 35), and revised its Wolf Management Plan in part to incorporate provisions for wolf management in the northern LP (MI DNR 2008a, p. 46).

The wolf population of Isle Royale National Park, Michigan, is not considered to be an important factor in the recovery of wolves in the WGL. The Park population is small and isolated and lacks genetic uniqueness (Wayne

et al.

1991, pp. 47-49). In addition, this island population probably has not had any contact with mainland wolf populations since its founding pair crossed the Lake Superior ice in the late 1940s (Peterson

et al.

1998, p. 828). For genetic reasons and constraints on expansion due to the island's small size, this wolf population does not contribute significantly towards meeting numerical recovery criteria; however, long-term research on this wolf population has added a great deal to our knowledge of the species. The wolf population on Isle Royale has ranged from 12 to 50 wolves since 1959, and was 19 wolves in the winter of 2009-2010 (Vucetich and Peterson 2010, p. 5).

Summary for Wisconsin and Michigan

The two-State wolf population, excluding Isle Royale wolves, has

exceeded 100 wolves since late-winter 1993-94 and has exceeded 200 wolves since late-winter 1995-96. Therefore, the combined wolf population for Wisconsin and Michigan has exceeded the second recovery criterion of the 1992 Revised Recovery Plan for a nonisolated wolf population, since 1999. Furthermore, the two-State population has exceeded the recovery criterion for an isolated second population since 2001.

Other Areas In and Near the Proposed Western Great Lakes DPS

No surveys have been conducted to document the number of wolves present in North Dakota or South Dakota, but an increasing number of wolves has apparently been detected in the eastern portions of these States. The eastern boundaries of North Dakota and South Dakota are approximately 19 and 81 mi (30 and 130 km), respectively, from occupied habitat in Minnesota. Biologists who are familiar with wolves in these States, however, generally agree that the wolves found there are primarily lone dispersers, although there were reports of pups being seen in the Turtle Mountains of North Dakota, in 1994 (Collins in litt. 1998).

Other records include an adult male shot near Devil's Lake, North Dakota in 2002, another adult male shot in Richland County in extreme southeastern North Dakota in 2003 (Fain in litt. 2006), and a vehicle-killed adult male found near Sturgis, South Dakota, in 2006 (Larson in litt. 2006). In contrast to the other South Dakota wolves of the last 25 years, the animal found near Sturgis was genetically identified as having come from the Greater Yellowstone area (Fain in litt. 2006). Most recently, a wolf was shot in Roberts County, South Dakota in January 2009 (reportedly running with two or three other wolves) (Prieksat in litt. 2009), and another wolf was found dead in a foothold trap that was set as part of an ongoing USDA Wildlife Service's coyote control operation in southeastern Eddy County, North Dakota (Bicknell in litt. 2009). See

Delineating the Boundaries of the Proposed WGL Gray Wolf DPS

in this proposed rule for a detailed discussion of movement of wolves.

Wolf dispersal is expected to continue as wolves travel away from the more saturated habitats in the primary range into peripheral areas where wolves are extremely sparse or absent. Unless they return to the primary range and join or start a pack there, they are unlikely to contribute to long-term maintenance of WGL wolf populations.

Although it is possible for these dispersers to encounter and mate with a mature wolf outside the primary range, the lack of large expanses of unfragmented habitat make it unlikely that wolf packs will persist in these peripheral areas; lack of contiguous habitat is expected to seriously impede further expansion. The only exception is the northern LP of Michigan, where several studies indicate that a persistent wolf population may develop (Gehring and Potter 2005, p. 1242; Potvin 2003, pp. 29-30), albeit dependent on occasional to frequent immigration of UP wolves. Despite the constraints on further expansion described here, however, current wolf populations in Minnesota, Wisconsin, and the UP of Michigan have already greatly exceeded the recovery levels defined in the 1992 Revised Recovery Plan, and maintenance of these numbers is not contingent on recruitment of wolves from areas outside the primary range that has been established for the WGL.

Summary of Wolf Recovery in the Western Great Lakes Region

Wolves in the proposed WGL DPS 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 criteria set by the Eastern Wolf Recovery Team in 1992 and reaffirmed in 1997 and 1998 (Peterson in litt. 1997, in litt. 1998), the proposed DPS contains sufficient wolf numbers and distribution to ensure their long-term survival within the DPS.

The maintenance and expansion of the Minnesota wolf population has maximized the preservation of the genetic diversity that remained in the proposed WGL DPS when its wolves were first protected in 1974. Furthermore, the Wisconsin-Michigan wolf population has exceeded the numerical recovery criterion even for a completely isolated second population. Therefore, even in the unlikely event that this two-State population was 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 11 and 10 years, respectively, 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 WGL have been met.

Have the Wolves of the Western Great Lakes Region Been Restored?

Leonard and Wayne (2008, p. 3) have stated that Great Lakes wolves have not been restored based on absence of certain historical mtDNA haplotypes from the current population, an estimated historical population size far greater than the current population size, and the admixture of coyote and western wolf haplotypes in the current population.

The spatial representativeness of both the historical and recent samples reported by Leonard and Wayne (2008) has been questioned by Mech (2009). For example, 16 recent but no historical samples from Minnesota were included in the study. Leonard and Wayne (2009) responded that they did not believe that genetic differences were likely to be pronounced at the geographic scale discussed by Mech and Paul (2008) and Mech (2009).

The current population of wolves in Minnesota, Wisconsin, and Michigan is derived from expansion of the remnant population in northeastern Minnesota (Fain

et al.

2010, p. 12), which was likely to have included both

C. lupus

and

C. lycaon

(Mech and Frenzel 1971; Mech 2010, p. 135), and in the case of UP Michigan, with possible contributions from

C. lycaon

from southern Ontario (Fain

et al.

2010, p. 12).

Subsequent studies with larger samples of the current wolf population find, despite acknowledged influence of western wolves, the current population is generally representative of the historical population (Fain

et al.

2010, p. 14; Wheeldon

et al.

2010). Koblmüller

et al.

(2009, pp. 10-11) found “comparatively slight” differentiation at autosomal microsatellite DNA loci between historical and current Great Lakes wolves. Wheeldon and White (2009, p. 4) present microsatellite DNA evidence that the hybridization processes noted by Leonard and Wayne (2008) were taking place over a century ago, so that the current population is comparable to the historical population with respect to admixture. Hybridization between eastern wolves and western wolves in the western Great Lakes region occurred prior to significant human effects on population size or habitat (Fain

et al.

2010, p. 14). According to Fain

et al.

(2010, p. 14), the current population of wolves in the western Great Lakes “represents an ancient component of the northeast ecosystem and have been

established throughout the region for thousands of years.”

The loss of mtDNA haplotypes found in historical but not the current western Great Lakes wolf population reported by Leonard and Wayne (2008, pp. 2-3) and the loss of allelic diversity (Fain

et al.

2010, p. 11), indicate that a genetic bottleneck occurred when wolves were nearly extirpated from the western Great Lakes region and the period of slow recovery that immediately followed. Despite these “founder effects” on the genetic composition of the western Great Lakes population, various measures of genetic diversity remain comparable to other wolf populations (Koblmüller

et al.

2009; Fain

et al.

2010, p. 12; Wheeldon

et al.

2010), at least partially owing to contributions from western wolves (

C. lupus

).

Wolves in the WGL region display a healthy level of heterozygosity (Fain

et al.

2010, p. 12), and show no evidence of genetic bottlenecks (Koblmuller

et al.

2009, p. 1). Schwartz and Vucetich (2009, p. 2) have stated that “By all accounts, the return of wolves to the Great Lakes region has been successful * * * they are doing superbly—both in terms of population viability and ecological function.” Cronin and Mech (2009, p. 2) state, “We suggest that wolves in the [W]GL region can simply be called a wolf population with mixed ancestry.” They further state that, “It is generally acknowledged that the Great Lakes wolf population is fit, with abundant genetic variation” (Cronin and Mech 2009, p. 2).

Distinct Vertebrate Population Segment Policy Overview

Pursuant to the Act, we consider whether information is sufficient to indicate that listing, reclassifying, or delisting any species, subspecies, or, for vertebrates, any DPS of these taxa may be warranted. To interpret and implement the DPS provision of the Act and congressional guidance, the Service and the National Marine Fisheries Service (NMFS) published a policy regarding the identification of distinct vertebrate population segments under the Act (Policy Regarding the Recognition of Distinct Vertebrate Population Segments Under the Endangered Species Act, 61 FR 4722, February 7, 1996) (hereafter DPS Policy). Under the DPS policy, two factors are considered in a decision regarding the potential identification of a DPS: (1) Discreteness of the population segment in relation to the remainder of the taxon, and (2) the significance of the population segment to the taxon to which it belongs. If a population meets both tests, it can be identified as a DPS. Then a third factor, the DPS's conservation status, is evaluated in relation to the Act's standards for listing, delisting, or reclassification, meaning that we undertake an analysis to determine whether the DPS is endangered or threatened or does not meet the criteria for listing. All three steps are necessary components of a complete DPS analysis.

Past Practice and History of Using DPSs

As of February 1, 2011, of the 392 native vertebrate listings, 85 are listed as less than an entire taxonomic species or subspecies (henceforth referred to in this discussion as populations) under one of several authorities, including the “distinct population segment” language in the Act's definition of species (section 3(16)). Thirty-three of these 85 populations, which span 52 different taxa, predate the 1996 DPS Policy; as such, the final listing determinations for these populations did not include formal policy-based analyses or expressly designate the listed entity as a DPS. In several instances, however, the Service and National Marine Fisheries Service (NMFS) have established a DPS and revised the List of Endangered and Threatened Wildlife in a single action, as shown in the following examples.

In February 1985, the Service delisted the brown pelican (

Pelecanus occidentalis

) in the southeastern United States and continued to identify it as endangered throughout the remainder of its range (50 FR 4938). In June 1994, NMFS revised the entry for the gray whale (

Eschrichtius robustus

) to remove the eastern North Pacific population from the List while retaining the western North Pacific population as endangered (59 FR 31094). In July 2003, the Service established two DPSs of the Columbian white-tailed deer (

Odocoileus virginianus leucurus

)—the Douglas County DPS and the Columbia River DPS—and delisted only the Douglas County DPS, while listing the Columbia River DPS (68 FR 43647). In March 2007, the Service established a DPS of the grizzly bear (

Ursus arctos horribilis

) for the Greater Yellowstone Area and surrounding area within the existing grizzly bear listing in the lower 48 States, and delisted this DPS (72 FR 14865). Also in March 2007, the Service identified the American crocodile (

Crocodylus acutus

) in Florida as a DPS within the existing endangered listing of the American crocodile in the United States and reclassified the Florida DPS from endangered to threatened (71 FR 13027). Revising and delisting the WGL DPS of wolves is consistent with the Service's past practice and does not represent a change in agency position.

Proposed Western Great Lakes Distinct Population Segment

In 1978, based on what was at that time the “best available biological data,” the Service stated that there were two “species” of gray wolves in the conterminous United States: “For purposes of this rulemaking, the gray wolf (

Canis lupus

) group in Mexico and the 48 conterminous States of the United States, other than Minnesota, is being considered as one “species,” and the gray wolf group in Minnesota is being considered as another “species.” (43 FR 9607, 9610, March 9, 1978). The Service then assigned a different status under the Act to each of those two “species,” finding the Minnesota gray wolf “species” to be threatened, while the other gray wolf “species” (the 48 conterminous States, except Minnesota, and in Mexico) to be endangered. The 1978 rule referred to the Minnesota listing as the listing of a “species” when, clearly, based on the information available at that time, the Minnesota wolves did not taxonomically constitute a separate species of wolf. Therefore, the 1978 listing either effectively established a Minnesota DPS or listed an entity in a portion of its broader range.

The DPS Policy (61 FR 4725, February 7, 1996) expressly provides for reexamining pre-policy DPS listings: “Any DPS of a vertebrate taxon that was listed prior to implementation of this policy will be reevaluated on a case-by-case basis as recommendations are made to change the listing status for that distinct population segment. The appropriate application of the policy will also be considered in the 5-year reviews of the status of listed species required by section 4(c)(2) of the Act.” Based on this provision, we are, within this proposed rule, (1) recognizing that a Minnesota DPS was established in 1978, (2) reevaluating that DPS listing, and (3) proposing to revise that DPS to meet the criteria in the DPS policy and to reflect the “best available biological data.”

A gray wolf DPS that includes only Minnesota does not meet the criteria in the DPS policy because it is not discrete “* * * in relation to the remainder of the species to which it belongs” (61 FR 4725, February 7, 1996). The Minnesota wolf population has expanded beyond State boundaries and is connected to the wolf population in Wisconsin and Michigan, as evidenced by frequent movements of wolves among the States (Van Deelen 2009, p. 140; Treves

at al.

2009, pp. 192-195) and genetic analyses

that demonstrate the Wisconsin and Michigan wolves are mostly from the same genetic mix as Minnesota wolves (Wheeldon and White 2009, p. 4; Fain

et al.

2010). Therefore, we are proposing to revise the boundaries of the Minnesota DPS to meet the criteria in the DPS policy as discussed under the

Distinct Population Segment Analysis,

below.

Geographical Area of the Proposed Western Great Lakes DPS

The geographical area of the proposed WGL DPS is shown in figure 1, below, and is described as all of Minnesota, Wisconsin, and Michigan; the portion of North Dakota north and east of the Missouri River upstream to Lake Sakakawea and east of the centerline of Highway 83 from Lake Sakakawea to the Canadian border; the portion of South Dakota north and east of the Missouri River; the portions of Iowa, Illinois, and Indiana north of the centerline of Interstate Highway 80; and the portion of Ohio north of the centerline of Interstate Highway 80 and west of the Maumee River at Toledo.

BILLING CODE 4310-55-P

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Distinct Population Segment Analysis

Analysis for Discreteness

Under the 1996 DPS Policy (61 FR 4722), a population segment of a vertebrate taxon may be considered discrete if it satisfies either of the following conditions: (1) it is markedly separated from other populations of the same taxon as a consequence of physical, physiological, ecological, or behavioral factors (quantitative measures of genetic or morphological discontinuity may provide evidence of this separation); or (2) it is delimited by international governmental boundaries within which differences in control of exploitation, management of habitat, conservation status, or regulatory mechanisms exist that are significant in light of section 4(a)(1)(D) of the Act.

Markedly Separated from Other Populations of the Same Taxon

—The western boundaries of the proposed WGL DPS are approximately 400 mi (644 km) from the nearest known gray wolf packs in Wyoming and Montana. The distance between those western packs and the nearest packs within the proposed WGL DPS is nearly 600 mi

(966 km). The area between Minnesota packs and northern Rocky Mountain (NRM) packs largely consists of unsuitable habitat, with only scattered islands of possibly suitable habitat, such as the Black Hills of eastern Wyoming and western South Dakota. There are no known populations of gray wolves to the south or east of the proposed WGL DPS within the United States.

As discussed in the previous section, wolves are known to disperse over vast distances, but straight line documented dispersals of 400 mi (644 km) or more are very rare. Although we cannot rule out the possibility of a WGL wolf traveling 600 mi (966 km) or more and joining or establishing a pack in the northern Rockies, such a movement has not been documented and is expected to happen very infrequently, if at all. Similar movements from the NRM wolf population into the proposed WGL DPS are unknown and are expected to happen infrequently. The 2006 Sturgis (South Dakota) wolf is the closest that an NRM wolf has come to entering the proposed WGL DPS (Fain in litt. 2006); however, the Sturgis wolf would still have had to travel over 300 mi (500 km) before encountering the nearest wolf pack in the proposed WGL DPS. As the discreteness criterion requires that the DPS be “markedly separated” from other populations of the taxon rather than requiring complete isolation, this high degree of physical separation between the WGL DPS and the northern Rocky Mountains satisfies the discreteness criterion.

Delimited by International Boundaries with Significant Management Differences

—The DPS policy allows us to use international borders to delineate the boundaries of a DPS if there are differences in control of exploitation, conservation status, or regulatory mechanisms between the countries. The border between the United States and Canada has been used as the northern boundary of the listed entity since gray wolves were reclassified in the lower 48 States and Mexico in 1978. There remain significant cross-border differences in exploitation, management, conservation status, and regulatory mechanisms. About 52,000 to 60,000 wolves occur in Canada, where suitable habitat is abundant (Boitani 2003, p. 322). Because of this abundance, wolves in Canada are not protected by Federal laws and are only minimally protected in most Canadian provinces (Pletscher

et al.

1991, p. 546). In the United States, unlike Canada, Federal protection and intensive management has been necessary to recover the wolf (Carbyn 1983).

In general, Canadian gray wolf populations are sufficiently large and healthy so that population regulation, rather than protection and close monitoring, is the management focus. There are an estimated 4,000 wolves in Manitoba (Manitoba Conservation undated). Hunting is allowed nearly province-wide, including in those provincial hunting zones adjoining northwestern Minnesota, with last year's season running from August 31, 2009, through March 31, 2010 (Manitoba Conservation 2009a). Trapping wolves is allowed province-wide, except in and immediately around Riding Mountain National Park (southwestern Manitoba), with last year's season running from October 14, 2008, through February 28 or March 31, 2009 (varies with trapping zone) (Manitoba Conservation 2009b).

The Ontario Ministry of Natural Resources estimates there are 8,850 wolves in the province, based on prey composition and abundance, topography, and climate and wolf numbers in most parts of the province are believed to be stable or increasing since about 1993 (Ontario MNR 2005a, pp. 7-9). In 2005, Ontario limited hunting and trapping of wolves by closing the season from April 1 through September 14 in central and northern Ontario (Ontario MNR 2005b). In southern Ontario, the portion of the province that is adjacent to the proposed WGL DPS, wolf hunting and trapping is permitted year round (Ontario MNR 2005c). If delisted, Minnesota, Wisconsin, and Michigan would carefully monitor and manage wolves to retain populations at or above the recovery goal (see Factor D). Therefore, even though biologically the WGL wolf population is simply a well-connected southern extension of wolves in Canada, we will continue to use the United States-Canada border to mark the northern boundary of the DPS due to the difference in control of exploitation, conservation status, and regulatory mechanisms between the two countries.

Conclusion

—We find, based on our analysis of the best available scientific information, that the proposed WGL DPS is markedly separated from other United States populations of gray wolves and difference in control of exploitation, conservation status, and regulatory mechanisms justifies discreteness between United States and Canadian wolf populations. Therefore, the proposed WGL DPS meets the criterion for discreteness under the DPS policy.

Analysis for Significance

If we determine that a population segment is discrete, we next consider available scientific evidence of its significance to the taxon to which it belongs. Our DPS policy states that this consideration may include, but is not limited to, the following: (1) Persistence of the discrete population segment in an ecological setting unusual or unique for the taxon; (2) evidence that loss of the discrete population segment would result in a significant gap in the range of the taxon; (3) evidence that the discrete population segment represents the only surviving natural occurrence of a taxon that may be more abundant elsewhere as an introduced population outside its historic range; and/or (4) evidence that the discrete population segment differs markedly from other populations of the species in its genetic characteristics. Factor 2 applies to the proposed WGL DPS and is included in our analysis for significance. Factors 1, 3, and 4 do not apply to the proposed WGL DPS and thus are not included in our analysis for significance.

Significant Gap in the Range of the Taxon

—Wolves once lived throughout most of North America. Wolves have been extirpated from most of the southern portions of their historical North American range. The successful restoration of a viable wolf metapopulation to large parts of Minnesota, Wisconsin, and Michigan has filled a significant gap in the holarctic range of gray wolves in the United States, and it provides an important extension of the range of gray wolves in North America. The loss of the WGL wolf population would, therefore, represent a significant gap in the species' holarctic range in that the WGL wolf population is the only wolf population in the conterminous States east of the Rocky Mountains, except for the red wolves (a different species) being restored along the Atlantic Coast, and currently holds about 40 percent of North American gray wolves known to occur south of Canada.

Finding

—We find, based on our analysis of the best available scientific information, that the proposed WGL DPS is significant to the taxon to which it belongs because its loss would result in a significant gap in the range of the taxon. Therefore, the proposed WGL DPS meets the criterion for significance under the DPS policy.

Discrete Vertebrate Population Segment Conclusion

We propose, based on our review of the best available scientific data, that the WGL DPS is discrete from other gray

wolf populations as a result of physical separation from other gray wolf populations in the United States and the international border with Canada. The DPS is significant to the taxon to which it belongs because it contains a wolf metapopulation that fills a large gap in the historical range of the taxon in the conterminous States. Therefore, we have determined that this population segment of wolves satisfies the discreteness and significance criteria required for a DPS. The evaluation of the appropriate conservation status for the proposed WGL DPS is found below.

Delineating the Boundaries of the Proposed WGL Gray Wolf DPS

In contrast to a species or a subspecies, a DPS is a biological population that is delineated by a boundary that is based on something other than established taxonomic distinctions. Therefore, the starting point for delineating a DPS is the biological population or metapopulation, and a geographical delineation of the DPS must reasonably represent the population or metapopulation and its biological characteristics and recovery needs.

To delineate the boundary of the proposed WGL DPS, we considered the current distribution of wolves in the Midwest and the characteristic movements of those wolves and of wolves elsewhere. We examined the best available scientific data on long-distance movements, including long-distance movements followed by return movements to the vicinity of the natal pack. We concluded that wolf behavior and the nature of wolf populations require that we include within the area of the DPS some subset of known long-distance movement locations. However, as explained below, wolf biology and common sense argue against including all known or potential long-distance movements within the DPS's boundaries.

The analysis detailed below resulted in the proposed boundaries of the WGL DPS that are shown in figure 1. This DPS has been delineated to include the core recovered wolf population plus a wolf movement zone around the core wolf populations. This geographic delineation is not intended to include all areas to which wolves have moved from the Great Lakes population. Rather, it includes the area currently occupied by wolf packs in Minnesota, Wisconsin, and Michigan; the nearby areas in these States in which wolf packs may become established in the foreseeable future; and a surrounding area into which Minnesota, Wisconsin, and Michigan wolves occasionally move but where persistent packs are not expected to be established because suitable habitat is rare and exists only as small patches. The area surrounding the core wolf populations includes the locations of most known dispersers from the core populations, especially the shorter and medium-distance movements from which wolves are most likely to return to the core areas and contribute to the wolf population. Therefore, the DPS encompasses the current range of the population, which is considered to be viable, including the primary range and the peripheral range.

The WGL areas that are regularly occupied by wolf packs are well documented in Minnesota (Erb and Benson 2004, p. 12, fig. 3; Erb and Don Carlos 2009, pp. 57-60), Wisconsin (Wydeven

et al.

2006, p. 33, fig. 1; Wydeven

et al.

2009c, pp. 93-98), and the UP of Michigan (Huntzinger

et al.

2005, pp. 25-27, figs. 4-6; Beyer

et al.

2009, pp. 73-75). Wolves have successfully colonized most, perhaps all, suitable habitat in Minnesota. Minnesota data from the winter of 2007-08 indicate that wolf numbers and density have stabilized since 1997-98, and there was no expansion of occupied range in the State (Erb 2008, pp. 5-7). Wisconsin wolves now occupy most habitat areas believed to have a high probability of wolf occurrence except for some areas of northeastern Wisconsin, and the State's wolf population continues to annually increase in numbers and, to a lesser degree, in area (Wydeven and Wiedenhoeft 2009, p. 2). The UP of Michigan has wolf packs throughout the peninsula. In the last 22 years, the wolf population in the UP has grown every year except 1997 and 2010 (Roell 2010, pers. comm.). Over the past 5 years, the average annual growth has been about 7 percent. While the population trend continues to increase, the rate of increase has slowed, consistent with any population expanding into and then filling available habitat. The population may continue to grow or remain steady; however, a small or even negative growth rate may occur any year and should be considered a natural fluctuation seen in any wildlife population.

When delineating the proposed WGL DPS, we had to consider the high degree of mobility shown by wolves. The dispersal of wolves from their natal packs and territories is a normal and important behavioral attribute of the species that facilitates the formation of new packs, the occupancy of vacant territories, and the expansion of occupied range by the “colonization” of vacant habitat. Data on wolf dispersal rates from numerous North American studies (summarized in Fuller

et al.

2003, p. 179, Table. 6.6; Boyd and Pletscher 1999, p. 1102, Table 6) show dispersal rates of 13 to 48 percent of the individuals in a pack. Sometimes the movements are temporary, and the wolf returns to a location in or near its natal territory. In some cases, a wolf may continue its movement for scores or even hundreds of miles until it locates suitable habitat, where it may establish a territory or join an existing pack. In other cases, a wolf is found dead at a distance from its original territory, leaving unanswered the questions of how far it would have gone and whether it eventually would have returned to its natal area or population.

Minnesota

—The current record for a documented movement by a wolf in North America is held by a Minnesota wolf that moved a minimum (that is, the straight-line distance from known starting point to most distant point) of at least 550 mi (886 km) northwest into Saskatchewan (Fritts 1983, pp. 166-167). Nineteen other primarily Minnesota movements summarized by Mech (in litt. 2005) averaged 154 mi (248 km). Their minimum distance of travel ranged from 32 to 532 mi (53-886 km) with the minimum dispersal distance shown by known returning wolves ranging from 54 mi (90 km) to 307 mi (494 km).

Wisconsin

—In 2004, a wolf tagged in Michigan was killed by a vehicle in Rusk County in northwestern Wisconsin, 295 mi (475 km) west of his original capture location in the eastern UP (Wydeven

et al.

2005b, p. 4). A north-central Wisconsin yearling female wolf traveled a similar distance (298 mi, 480 km) to the Rainy Lake region of Ontario during 1988-89 (Wydeven

et al.

1995, p. 149).

Michigan

—Drummer

et al.

(2002, pp. 14-15) reported 10 long-distance dispersal events involving UP wolves. One of these wolves moved to north-central Missouri and another to southeastern Wisconsin, both beyond the core wolf areas in the WGL. The average straight-line distance traveled by those two wolves was 377 mi (608 km), while the average straight-line distance for all 10 of these wolves was 232 mi (373 km). Their straight-line distances ranged from 41 to 468 mi (66 to 753 km).

Illinois and Indiana

—In December 2002, a Marshall County (Illinois) wolf likely dispersed from the Wisconsin wolf population, nearly 200 mi (322 km) to the north (Great Lakes Directory 2003). The Randolph County (Indiana) wolf had traveled a minimum distance of at least 428 mi (689 km) to get around

Lake Michigan from its central Wisconsin birthplace; it likely traveled much farther than that unless it went through the city or suburbs of Chicago (Wydeven

et al.

2004, pp. 10-11; Treves

et al.

2009, p. 194). The Pike County (Illinois) wolf that was shot in late 2005 was about 300 mi (180 km) from the nearest wolf packs in central Wisconsin.

North Dakota, South Dakota, and Nebraska

—Licht and Fritts (1994, p. 77) tabulated seven wolves found dead in North Dakota and South Dakota from 1981 through 1992 that are believed to have originated from Minnesota, based on skull morphometrics. Although none of these wolves were marked or radio-tracked, making it impossible to determine the point of initiation of their journey, a minimum travel distance for the seven can be determined from the nearest wolf breeding range in Minnesota. For the seven, the average distance to the nearest wolf breeding range was 160 mi (257 km) and ranged from 29 to 329 mi (46 to 530 km). One of these seven wolves moved west of the Missouri River before it died.

Genetic analysis of a wolf killed in Harding County, in extreme northwestern South Dakota, in 2001 indicated that it originated from the Minnesota-Wisconsin-Michigan wolf populations (Fain in litt. 2006). The straight-line travel distance to the nearest Minnesota wolf pack is nearly 400 mi (644 km).

The wolf from the Greater Yellowstone area that was killed by a vehicle on Interstate 90 near Sturgis, South Dakota, in March of 2006 traveled a minimum straight-line distance of about 270 mi (435 km) from the nearest known Greater Yellowstone pack before it died (USFWS

et al.

2006, in USFWS Program Report, Figure 1).

A large canid was shot by a Boyd County (Nebraska) rancher in late 1994 or early 1995, likely after crossing the frozen Missouri River from South Dakota (Anschutz in litt. 2006, Jobman in litt. 1995). It was determined to be a wolf that originated from the Great Lakes wolf populations (Fain in litt. 2006), whose nearest pack would have been about 300 mi (480 km) away. A wolf illegally killed near Spalding, Nebraska, in December of 2002 also originated from the Minnesota-Wisconsin-Michigan wolf population, as determined by genetic analysis (Anschutz in litt. 2003, Fain in litt. 2006). The nearest Minnesota wolf pack is nearly 350 mi (563 km) from this location.

Other notable extra-territorial movements

—The extra-territorial movements of several wolves were radio-tracked in sufficient detail to provide insight into their actual travel routes and total travel distances for each trek, rather than only documenting straight-line distance from beginning to end-point. Merrill and Mech (2000, pp. 429-431) reported on four such Minnesota wolves with documented travel distances ranging from 305 to 2,640 mi (490 to 4,251 km) and an average travel route length of 988 mi (1590 km). Wydeven (1994, pp. 20-22) described a Wisconsin wolf that moved from northwestern Wisconsin to the northern suburbs of St. Paul, Minnesota, for 2 weeks (apparently not seen or reported to authorities by the local residents), then moved back to north-central Wisconsin. The total travel distance was 278 mi (447km) from her natal pack into Minnesota and on to the north-central Wisconsin location where she settled down.

While investigating the origins of Scandinavian wolf populations, Linnell

et al.

(2005, p. 387) compiled wolf dispersal data from 21 published studies, including many cited separately here. Twenty-two of 298 compiled dispersals (7.4 percent) were over 300 km (186 mi). Eleven dispersals (3.7 percent) were over 500 km (311 mi). Because of the likelihood that many long-distance dispersers are never reported, they conclude that the proportion of long-distance dispersers is probably severely underestimated.

From these extra-territorial movement records, we conclude that wolf movements of over 200 mi (320 km) straight-line distance have been documented on numerous occasions, while shorter distance movements are more frequent. Movements of 300 mi (480 km) straight-line distance or more are less common, but include one Minnesota wolf that journeyed a straight-line distance of 300 mi (480 km) and a known minimum-travel distance of 2,640 mi (4,251 km) before it reversed direction, as determined by its satellite-tracked collar. This wolf ultimately returned to a spot only 24 mi (40 km) from its natal territory (Merrill and Mech 2000, p. 430). Although much longer movements have been documented, including some by midwestern wolves, return movements to the vicinity of natal territories have not been documented for extra-territorial movements beyond 300 mi (480 km).

Based on these extra-territorial movement data, we conclude that affiliation with the midwestern wolf population is diminished and essentially lost when dispersal takes a Midwest wolf a distance of 250 to 300 mi (400 to 480 km) beyond the outer edge of the areas that are continuously occupied by wolf packs. Although some WGL wolves will move beyond this distance, available data indicate that longer distance dispersers are unlikely to return to their natal population. Therefore, they have lost their functional connection with, and potential conservation value to, the WGL wolf population.

Wolves moving substantial distances outward from the core areas of Minnesota, Wisconsin, and Michigan will encounter landscape features that are at least partial barriers to further wolf movement and that may, if crossed, impede attempts of wolves to return toward the WGL core areas. If such partial barriers are in a location that has separate utility in delineating the biological extent of a wolf population, they can and should be used to delineate the DPS boundary. Such landscape features are the Missouri River in North Dakota and downstream to Omaha, Nebraska, and Interstate Highway 80 from Omaha eastward through Illinois, Indiana, and into Ohio, ending where this highway crosses the Maumee River in Toledo, Ohio. We do not believe these are absolute barriers to wolf movement. There is evidence that several Minnesota-origin wolves have crossed the Missouri River (Licht and Fritts 1994, pp. 75 & 77, Fig. 1 and Table 1; Anschutz in litt. 2003, 2006) and some Midwest wolves have crossed interstate highways (Merrill and Mech 2000, p. 430). There is also evidence that some wolves are hesitant to cross highways (Whittington

et al.

2004, pp. 7, 9; Wydeven

et al.

2005b, p. 5; but see Blanco

et al.

2005, pp. 315-316, 319-320 and Kohn

et al.

2000, p. 22). Interstate highways and smaller roads are a known mortality factor for wolves and, therefore, pose a partial barrier to wolf movements (Blanco

et al.

2005, p. 320). The death of a NRM wolf near Sturgis in western South Dakota (Fain in litt. 2006) suggests that the area of the Dakotas west of the Missouri River may be traversed by a small number of wolves coming from both the NRM and WGL wolf populations, as well as wolves from Canada (Licht and Fritts 1994, pp. 75-77). Wolves in this area cannot be assumed to belong to the WGL wolf population, supporting our belief that the boundary should not be designed to include the locations of all known dispersers.

Summary of Factors Affecting the Species

Section 4 of the Act and its implementing regulations (50 CFR part 424) set forth the procedures for listing species, reclassifying species, or removing species from listed status.

“Species” is defined by the Act as including any species or subspecies of fish or wildlife or plants, and any distinct vertebrate population segment of fish or wildlife that interbreeds when mature (16 U.S.C. 1532(16)). Once the “species” is identified, we then evaluate whether that species may be endangered or threatened because of one or more of the five factors described in section 4(a)(1) of the Act. We must consider these same five factors in delisting a species. We may delist a species according to 50 CFR 424.11(d) if the best available scientific and commercial data indicate that the species is neither endangered nor threatened because (1) the species is extinct, (2) the species has recovered and is no longer endangered or threatened, or (3) the original scientific data used at the time the species was classified were in error.

A recovered species is one that no longer meets the Act's definition of threatened or endangered. The analysis for a delisting due to recovery must be based on the five factors outlined in section 4(a)(1) of the Act. This analysis must include an evaluation of threats that existed at the time of listing, those that currently exist, and those that could potentially affect the species once the protections of the Act are removed.

In the context of the Act, the term “threatened species” means any species or subspecies or, for vertebrates, Distinct Population Segment (DPS) that is likely to become an endangered species within the foreseeable future throughout all or a significant portion of its range. The term “endangered species” means any species that is in danger of extinction throughout all or a significant portion of its range. The Act does not define the term “foreseeable future.” For the purpose of this proposal, 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 WGL DPS. For the proposed WGL DPS, the foreseeable future differs for each factor potentially affecting the DPS.

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 in the proposed WGL DPS. 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 States have had a solid history of cooperating and assisting in wolf recovery and have made a commitment, through legislative actions, to continue these activities. We believe 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. As explained previously, our analysis of the factors affecting the WGL DPS refer to the gray wolf (

C. lupus

), because that is the named entity currently on the List of Endangered and Threatened Wildlife (see

Procedural Aspects of Proposal Applying to the Gray Wolf

above).

A. The Present or Threatened Destruction, Modification, or Curtailment of Its Habitat or Range

A common misconception is that wolves inhabit only remote pristine forests or mountainous areas, where human developments and other activities have produced negligible change to the natural landscape. Their extirpation south of Canada and Alaska, except for the heavily forested portions of northeastern Minnesota, reinforced this popular belief. However, the primary reason wolves survived in those areas was not because of habitat conditions, but, rather, because remote areas were sufficiently free of the human persecution that elsewhere killed wolves faster than the species could reproduce (Mech 1995a, p. 271).

In the western Great Lakes region, wolves in the densely forested northeastern corner of Minnesota have expanded into the more agricultural portions of central and northwestern Minnesota, northern and central Wisconsin, and the entire UP of Michigan. Habitats currently being used by wolves span the broad range from the mixed hardwood-coniferous forest wilderness area of northern Minnesota, through sparsely settled, but similar habitats in Michigan's UP and northern Wisconsin, and into more intensively cultivated and livestock-producing portions of central and northwestern Minnesota and central Wisconsin.

Wolf research and the expansion of wolf range over the last three decades have shown that wolves can successfully occupy a wide range of habitats, and they are not dependent on wilderness areas for their survival. In the past, for instance, wolf populations occupied nearly every type of habitat north of mid-Mexico that contained large ungulate prey species, including bison, elk, white-tailed deer, mule deer, moose, and woodland caribou; thus, wolves historically occupied the entire Midwest. Inadequate prey density or high levels of human-caused mortality appear to be the only factors that limit wolf distribution (Mech 1995a, p 271; 1995b, p. 544).

Suitable Habitat Within the Proposed Western Great Lakes DPS

Various researchers have investigated habitat suitability for wolves in the central and eastern portions of the United States. In recent years, 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, p. 769-770; Wydeven

et al.

2001a, 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-related 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 miles per sq mi (0.6 to 0.7 km per sq km) (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 sq mi (0.45 km per sq km) (Mladenoff

et al.

1995, p. 289). However, recent work in the UP 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 UP. In Minnesota, a combination of road density and human density is used by MN DNR to model suitable habitat. Areas with a human density up to 8 people per sq km are suitable if they also have a road density less than 0.5 km per sq km. Areas with a human density of less than 4 people per sq km are suitable if they have road densities up to 0.7 km per sq km (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-related 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 sq mi (0.84 km per sq km), but human-related factors produced 71 percent of the wolf deaths in areas with higher road densities (Wydeven

et al.

2001a, pp. 112-113).

Some researchers have used a road density of 1 mi per sq mi (0.6 km per sq km) 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 sq mi (0.45 km per sq km) 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 sq mi (0.6 km per sq km) 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 (WI DNR 1997, 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 and, therefore, less-suitable areas (Mech 1989, pp. 387-388; Wydeven

et al.

2001a, 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 has been completed (Mladenoff

et al.

2009). This new 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 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 (see Figure 2, below; 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 high and moderate probability of occupancy (primary and secondary wolf habitat). The late-winter 2008-09 Wisconsin wolf survey identified packs occurring throughout the central Wisconsin forest area (Wolf Management Zone 2, Figure 3) and across the northern forest zone (Zone 1, Figure 3), with highest pack densities in the northwest and north-central forest; pack densities are lower, but increasing, in the northeastern corner of the State (Wydeven and Wiedenhoeft 2009, Figure 1).

Michigan wolf surveys in winter 2009-10 continue to show wolf pairs or packs (defined by Michigan DNR as two or more wolves traveling together) in every UP county except Keweenaw County (Huntzinger

et al.

2005, p. 6; Roell 2011, pers. comm.), which probably lacks a suitable ungulate prey base during winter months (Potvin

et al.

2005, p. 1665).

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 LP 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 LP (Potvin 2003, p. 39). The northern LP is separated from the UP by the Straits of Mackinac, whose 4-mile (6.4-km) width freezes during mid- and late-winter in some years. In recent years there have been several documented occurrences of wolves in the northern LP, but until 2010, there had been no indication of persistence beyond several months. Prior to those occurrences, the last recorded wolf in the LP was in 1910.

In the first instance a radio-collared female wolf from the eastern UP was trapped and killed by a coyote trapper in Presque Isle County in late October 2004. In late November 2004, tracks from two wolves were verified in the same northern LP county. Follow-up winter surveys by the DNR in early 2005 failed to find additional wolf tracks in the northern LP (Huntzinger

et al.

2005, p. 7); additional surveys conducted in 2006-10 also failed to find evidence of continued northern LP wolf presence (Roell

et al.

2009, p. 5; Roell 2010, pers. comm.). A video of a single wolf was taken near Mackinac City in Cheboygan County in May 2009, and another trail-camera video-recorded a wolf in Presque Isle County in July 2009. These two sightings may have been the same animal (Roell 2009, pers. comm.). In 2010, USDA Wildlife Services and MI DNR staff confirmed a single breeding pair with pups in Cheboygan County in the northern LP (MI DNR 2010).

These northern LP patches of potentially suitable habitat contain a great deal of private land, are small in comparison to the occupied habitat on the UP and in Minnesota and Wisconsin, and are intermixed with agricultural and higher road density areas (Gehring and Potter 2005, p. 1240). Therefore, continuing wolf immigration from the UP may be necessary to maintain a future northern LP population. The Gehring and Potter study (2005, p. 1239) predicted 850 sq mi (2,198 sq km) of suitable habitat (areas with greater than a 50 percent probability of wolf occupancy) in the northern LP. Potvin (2003, p. 21), using deer density in addition to road density, believes there are about 3,090 sq mi (8,000 sq km) of suitable habitat in the northern LP. Gehring and Potter (2005, p. 1239) exclude from their calculations those northern LP low-road-density patches that are less than 19 sq mi (50 sq km), while Potvin (2003, pp. 10-15) does not limit habitat patch size in his calculations. Both of these area estimates are well below the minimum area described in the Revised Recovery Plan, which states that 10,000 sq mi (25,600 sq km) of contiguous suitable habitat is needed for a viable isolated gray wolf population, and half that area (5,000 sq mi or 12,800 sq km) is needed to maintain a viable wolf population that is subject to wolf immigration from a nearby population (USFWS 1992, pp. 25-26).

Based on the above-described studies and the guidance of the 1992 Revised Recovery Plan, the Service has concluded that suitable habitat for wolves in the proposed WGL DPS can be determined by considering four

factors: Road density, human density, prey base, and size. An adequate prey base is an absolute requirement, but in much of the proposed WGL DPS the white-tailed deer density is well above adequate levels, causing the other factors to become the determinants of suitable habitat. Prey base is primarily of concern in the UP where severe winter conditions cause deer to move away from some lakeshore areas, making otherwise suitable areas locally and seasonally unsuitable. Road density and human density frequently are highly correlated; therefore, road density is the best single predictor of habitat suitability. However, areas with higher road density may still be suitable if the human density is very low, so a consideration of both factors is sometimes useful (Erb and Benson 2004, p. 2).

Finally, although the territory of individual wolf packs can be relatively small, packs are not likely to persist as a viable population if they occupy a small isolated island of otherwise unsuitable habitat. The 1992 Revised Recovery Plan indicates that a wolf population needs to occupy at least 10,000 contiguous sq mi (25,600 sq km) to be considered viable if it is isolated from other wolf populations, and must occupy at least half that area if it is not isolated from another self-sustaining population (USFWS 1992, pp. 25-26).

Based on the information discussed above, we conclude that Minnesota Wolf Management Zone A (Federal Wolf Management Zones 1-4, Figure 2), Wisconsin Wolf Zones 1 and 2 (Figure 3), and the UP of Michigan contain a sufficient amount of suitable wolf habitat. The other areas within the DPS are unsuitable habitat, or are potentially habitat that is too small or too fragmented to be suitable for maintaining a viable wolf population.

Wolf Populations on Federal Lands

National forests, and the prey species found in their various habitats, have been important to wolf conservation and recovery in the core areas of the proposed WGL DPS. There are five national forests in Minnesota, Wisconsin, and Michigan (Superior, Chippewa, Chequamegon-Nicolet, Ottawa, and Hiawatha National Forests) with wolf packs that exclusively or partially reside on them. Their wolf populations range from approximately 484 on the Superior National Forest in northeastern Minnesota, to an estimated 182 on the UP's Ottawa National Forest, 164 on the Chequamegon-Nicolet National Forest in northeastern Wisconsin, and another estimated 49 on the Hiawatha National Forest in the eastern UP (Delphey 2009, pers. comm.; Eklund 2009, pers. comm.; Roell 2011, pers. comm., Wydeven 2011, pers. comm.).

Voyageurs National Park, along Minnesota's northern border, has a land base of nearly 340 sq mi (882 sq km). As of the last survey in 2008, there were 31 to 46 wolves within 7 to 9 packs that exclusively or partially reside within the park, and at least 5 packs are located wholly inside the Park boundaries (Ethier

et al.

2008, p. 5). The 2008 estimates fall within the range of wolf estimates for the Park from the 1990s (Gogan

et al.

2004) and early 2000s (Fox

et al.

2001, pp. 6-7).

Within the boundaries of the proposed WGL DPS, we currently manage seven units within the National Wildlife Refuge System with significant wolf activity. Primary among these are Agassiz National Wildlife Refuge (NWR), Tamarac NWR, and Rice Lake NWR in Minnesota; Seney NWR in the UP of Michigan; and Necedah NWR in central Wisconsin. Agassiz NWR has had as many as 20 wolves in 2 to 3 packs in recent years. Although in 1999 mange and illegal shootings reduced them to a single pack of five wolves and a separate lone wolf, since 2001, two packs with a total of 10 to 12 wolves have been using the Refuge. About 60 percent of the packs' territories are located on the Refuge or on an adjacent State-owned wildlife management area (Huschle in litt. 2005).

Data collected by Agassiz NWR staff during winter wolf sign surveys conducted in cooper

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