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  • Northeastern Gray WolfDistinct Population Segment

    By Certified Mail

    1 April 2003

    U. S. FISH AND WILDLIFE SERVICEUNITED STATES DEPARTMENT OF THE INTERIOR

    Defenders of Wildlife )1101 14th St. NW, Suite 1400 )Washington, D.C. 20005 )Tel: (202) 682-9400 )

    )and )Sierra Club )Legislative Office )408 C St., N.E. )Washington, DC 20002 )

    )and )RESTORE: The North Woods )P.O. Box 1099 )Concord, MA 01742 )

    )and )The Wildlands Project )P.O. Box 455 )Richmond, VT 05477 )

    Petition to list a distinct population segment ofgray wolves (16 U.S.C. § 1533 and 5U.S.C. § 553) generally recognized as theNortheastern United States

    Submitted by,

    Martin E. SmithCarnivore Biologist

  • Defenders of Wildlife

    I. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3A. The Petitioners . . . . . . . . . . . . . . . . . . . 3B. Current Legal Status . . . . . . . . . . . . . . . 5C. DPS and ESA Criteria . . . . . . . . . . . . . . 6D. Overview and Current Issues . . . . . . . . 7

    II. NATURAL HISTORY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8A. Description of the Species . . . . . . . . . . 8

    Physical description . . . . . . . . . . . . 8Pack Behavior . . . . . . . . . . . . . . . . 9Reproduction . . . . . . . . . . . . . . . . . 9Communication . . . . . . . . . . . . . . . 9

    B. Taxonomy . . . . . . . . . . . . . . . . . . . . . . 10C. Historical Distribution in the Northeast 11

    III. WOLF ECOLOGY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12A. The Role of the Wolf as a Top Carnivore12

    Energy transfer between trophiclevels . . . . . . . . . . . . . . . . 12

    Mesopredator release . . . . . . . . . . 12Regulation of prey genetic health 12Wolf behavior and population

    dynamics . . . . . . . . . . . . . 13B. Ecological Importance of Dispersal and

    Management Considerations . . . . 13Gray wolf dispersal . . . . . . . . . . . 13Landscape characteristics of

    dispersal paths . . . . . . . . . 14Dispersal mortality . . . . . . . . . . . . 14Dispersal distance . . . . . . . . . . . . 15

    C. Minimum Viable Population Size . . . . 16

    IV. NORTHEASTERN GRAY WOLF DPS PETITION PROPOSAL . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17A. Distinct Population Segments under the

    Endangered Species Act . . . . . . . 17Discreteness and significance . . . 17Determination of federal protections17

    B. DPS Boundaries and Habitat Description17C. Suitability of the Northeast for Gray

    Wolf Restoration . . . . . . . . . . . . . 19Land availability . . . . . . . . . . . . . 19Road density . . . . . . . . . . . . . . . . 19Prey base . . . . . . . . . . . . . . . . . . . 20Human Attitudes . . . . . . . . . . . . . 20

  • Ecosystem Impacts . . . . . . . . . . . . 20Economic Impacts . . . . . . . . . . . . 21

    D. Qualifications of the Northeastern GrayWolf Population as a DPS . . . . . . 22Significance. . . . . . . . . . . . . . . . . 22

  • E. Northeastern Gray Wolf DPSQualifications for ESA Listing . . 23Conservation Status . . . . . . . . . . . 23

    1. The present or threateneddestruction,modification, or curtailment of itshabitat or range. . . 24

    2. Overutilization forcommercial,recreational,scientific, or educational purposes24

    3. Disease or predation . . . 254. The inadequacy of existing

    regulatorymechanisms . . . . . 25

    5. Other natural or manmadefactors . . . . . . . . . 25

    Environmental Stochastity . . . . . . 25Negative Human Attitudes . . . . . . 26Development and human population

    growth . . . . . . . . . . . . . . . 27

    IV. SUMMARY AND CONCLUSIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

    V. LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

  • Northeastern Gray Wolf DPS - 1 April 2003 5

    I. INTRODUCTION

    Defenders of Wildlife, Sierra Club,RESTORE: The North Woods, and theWildlands Project (Petitioners) herebypetition the U. S. Fish and Wildlife Service(FWS) to list a Distinct Population Segment(DPS) of wolves as endangered under theEndangered Species Act (ESA) (16 U.S.C. §1533) and the Administrative Procedure Act(5 U.S.C. § 553). The DPS is defined inSection III but generally represents thenortheastern United States. The gray wolf(Canis lupus) in the northeastern U.S.(Maine, Vermont, New York, NewHampshire) has recently been designated apart of the Eastern Gray Wolf DPS and iscurrently classified as threatened under theESA (Fed. Reg. Vol. 68, No. 62, Tuesday,April 1, 2003, pp. 15804-15875). Inaddition, the FWS, in an advance notice ofproposed rulemaking, has indicated its intentto delist the species and to forego an activerecovery effort in this region (Fed. Reg. Vol.68, No. 62, Tuesday, April 1, 2003, pp.15876-15879).

    In this petition, we will present severalfactors that establish the significance anddiscreteness of a Northeast population to theconservation of gray wolves in the lower 48states. We will also present documentationof vast areas of suitable habitat andfavorable conditions for the establishment ofviable populations of wolves in this region.Recent studies indicate that suitable habitatand sufficient prey exist for wolves in NewEngland, from northern Maine acrossnorthern New Hampshire and Vermont toAdirondack Park in New York (Mladenoff1998, Wydeven et al. 1998, Harrison andChapin 1997, Hosack 1996). These studiessuggest that the Northeast could support atleast 1,200 wolves and perhaps as many as

  • Northeastern Gray Wolf DPS - 1 April 2003 6

    1,800. In the Northeast, the distance fromextant wolf populations in the Great Lakesregion, combined with anthropogenic andgeographic barriers between the U.S. andCanadian wolf populations, preclude thereasonable expectation that wolves willnaturally recolonize the region (Wydeven etal 1998, Harrison and Chapin 1997). It is tothese areas that we wish to see active wolfrecovery initiated by the U. S. Fish andWildlife Service. Finally, we will show thata wolf population located in the northeasternU.S. would qualify as an endangered speciesunder the ESA. We believe that the FWS islegally obligated to establish a NortheastDPS and to expeditiously complete andimplement a recovery plan that addressesthe entire geographic area encompassed bythis proposed DPS.

    A. The PetitionersDefenders of Wildlife (Defenders) is a non-profit, science-based, conservationorganization with more than 430,000members and an extensive involvement inwolf restoration and protection in NorthAmerica. For more than 30 years Defendershas been directly involved in making graywolf recovery a reality in the lower 48states. Our activities in this arena include:

    < lobbying Congress and variousadministrations for wolf recovery actionsand funding;

    < litigating on behalf of wolves as well asintervening on behalf of the government toprotect the Yellowstone and Mexican graywolf recovery efforts;

    < operating a privately funded wolfcompensation trust in the northern Rockiesand other regions since 1987;

    < offering and paying rewards for informationleading to the conviction of illegal wolfkillers;

  • Northeastern Gray Wolf DPS - 1 April 2003 7

    < working with current and potentialcooperating tribes often providing technicaltraining and funding for equipment orpersonnel;

    < funding and training field staff to manageand protect wolves in recovery areas;

    < sponsoring educational symposia andactivities such as the annual North AmericanInteragency Wolf Conference and WolfAwareness Week to educate and organizewolf supporters and others;

    < financing and participating in numerousscientific studies to gauge habitat suitabilityand public support for wolf recovery,documenting wolf-related ecologicalphenomenon, and testing the efficacy ofmany management approaches andtechniques;

    < providing emergency funding and staffduring the government shutdown of 1996 tocomplete the second Yellowstonereintroduction; and

    < providing support for captive breedingfacilities.

    In December 1999, Defenders published Places for Wolves: A Blueprint for Restorationand Long-term Recovery in the Lower 48States (Ferris et al. 1999) as our formal anddetailed response to early drafts of the FWSgray wolf reclassification proposal. Thisdocument, which was recognized as theNatural Resource Council of America's 1999Conservation Publication of the Year, laysout our science-based vision for whatfederally-led wolf recovery should entail.The publication identifies several areas thatoffer outstanding opportunities for wolfrecovery, including the northeastern UnitedStates. To help enable wolf recovery in thisarea, Defenders has agreed to extend TheBailey Wildlife Foundation WolfCompensation Trust to this region untilwolves no longer require federal protection.We have launched a national public

  • Northeastern Gray Wolf DPS - 1 April 2003 8

    education and outreach program thatincludes traveling education booths, a wolfcurriculum and a biennial internationalpredator conference.

    RESTORE: The North Woods (RESTORE) is a regional non-profit conservation groupdevoted to restoring and preservingwilderness and wildlife in the North Woodsof New England. Their major programsinclude: (1) promoting grassroots efforts torestore imperiled wildlife species; (2)rallying public support and purchasing landto create a 3.2-million-acre Maine WoodsNational Park; and (3) inspiring a wildernessrevival by raising awareness of and workingto protect wild forests in New England.

    Since the organization was founded in 1992,RESTORE has become a leading voice forendangered wildlife in this region. Theywere the first local organization to launch acampaign to restore the eastern wolf to thenortheastern United States. They initiated acitizen petition to protect the Atlanticsalmon, which led to a federal protection ofthe species as endangered. And, they areworking with a national coalition of groupsthat used administrative and legal means towin Endangered Species Act protection forthe Canada lynx. RESTORE is alsospearheading efforts to protect core habitatfor the full range of native wildlife. Towardthis end, they have worked with aphilanthropist to purchase and protect nearly15,000 acres in the Maine Woods.RESTORE has more than 1,500 members,runs offices in Concord, Massachusetts andHallowell, Maine and a seasonal MaineWoods Visitor Center in Bar Harbor, Maine,and employs seven staff members.

    Sierra Club is the oldest and largest grassroots conservation organization in the UnitedStates with more than 700,000 members inall 50 States and Puerto Rico. The Sierra

  • Northeastern Gray Wolf DPS - 1 April 2003 9

    Club works to explore, enjoy and protect thewild places of the Earth; practice andpromote the responsible use of the Earth'secosystems and resources; and educate andenlist humanity to protect and restore thequality of the natural and human environment.

    The Wildlands Project is working to restore and protect the natural heritage of NorthAmerica. Through advocacy, education,scientific consultation, and cooperation withmany partners, they are designing andhelping to create systems of interconnectedwilderness areas that can sustain thediversity of life. Wild Earth—the quarterlypublication of the WildlandsProject—inspires effective action for wildnature by communicating the latest thinkingin conservation science, philosophy, policy,and activism, and serves as a forum fordiverse views within the conservationmovement.

    B. Current Legal StatusUntil recently, under provisions of the ESA(43 Fed. Reg. 9607-9615 March 9, 1978), allgray wolves south of the UnitedStates–Canada border (including Mexico)were listed as endangered, except inMinnesota (where they were listed asthreatened) and in the three non-essentialand experimental areas of Yellowstone,central Idaho, and Arizona. Since its initiallisting, the gray wolf has made someprogress in parts of its historical range. InJuly, 2000, the FWS proposed areclassification of gray wolves under theESA that would establish four DistinctPopulation Segments (DPSs) covering all orparts of 19 states and Mexico (65 Fed. Reg.43450 - 43496, July 13, 2000). Theseproposed DPSs were: Western Gray WolfDPS (threatened status: WA, OR, ID, MT,WY, UT, CO, northern NM, and northernAZ); Southwestern (Mexican) Gray Wolf

  • Northeastern Gray Wolf DPS - 1 April 2003 10

    DPS: (endangered status: southern AZ,southern NM, west TX, Mexico); WesternGreat Lakes Gray Wolf DPS (threatenedstatus, ND, SD, MN, WI, MI); andNortheastern Gray Wolf DPS (threatenedstatus: NY, NH, ME, and VT).

    On April 1, 2003, the FWS released its final rule on gray wolf reclassification, whichdiffered substantially from the proposedrule. In the final rule, the FWS reclassifiedthe gray wolf under three DPSs: the EasternGray Wolf DPS (Minnesota, Wisconsin,Michigan, North Dakota, South Dakota,Kansas, Nebraska, Iowa, Missouri, Illinois,Indiana, Ohio, Pennsylvania, New York,New Jersey, Connecticut, Rhode Island,Massachusetts, Vermont, New Hampshireand Maine); the Western Gray Wolf DPS(Montana, Idaho, Wyoming, Washington,Oregon, California, Nevada, northern Utah,and northern Colorado); and theSouthwestern Gray Wolf DPS (Arizona,New Mexico, southern Colorado, southernUtah, western Oklahoma, western Texas,and Mexico). For the Eastern and WesternGray Wolf DPSs, the gray wolf has beendownlisted to threatened, while in theSouthwestern DPS, the classificationremains endangered. Gray wolves lose allESA protection in any state not includedwithin one of these DPSs. On April 1, 2003,the FWS also published an advance noticeof proposed rulemaking, announcing theagency’s intent to propose delisting both theEastern and Western Gray Wolf DPSswithin the next two years. In effect, thismeans that when populations arereestablished in no more than 6 of the 48conterminous United States, the gray wolfwill lose all federal protections under theproposed plan (Fed. Reg. Vol. 68, No. 62,April 1, 2003, pp. 15804-15882).

  • Northeastern Gray Wolf DPS - 1 April 2003 11

    The goal of endangered species protection is to recover endangered species. Beforewolves in the lower 48 States can beconsidered recovered and then delisted,continued threats to and negative attitudestoward the wolf must be adequatelyaddressed. The FWS decision to removeand reduce federal protections for the graywolf will increase the probability ofextinction for many existing small andisolated populations due primarily to thewolf’s dispersal behavior. Regardless of theprotection a wolf population has within theboundaries of a recovery zone, naturereserve, or park, wolves will disperse to newterritories beyond reserve boundaries insearch of prey and mates–areas where theywill not receive adequate legal protection(Mladenoff et al. 1995, Gese and Mech1991).

    While many wildlife biologists and conservationists cautiously celebrate the success ofthe gray wolf in the northern RockyMountains and Great Lakes recovery areas,population sizes are still far from what theywere prior to the twentieth century. In thelower 48 states, wolves currently occupyless than 2 percent of their historic range.For example, early estimates of theYellowstone region wolf population exceedthirty-five thousand animals, while today,recovering populations number only a fewhundred (Fischer 1995). In addition to thelow numbers of wolves currently foundthroughout the species’ former historicalrange, present-day wolves are separated byenormous distances and numerousanthropogenic barriers that impede naturaldispersal and movement betweenpopulations (65 Fed. Reg. 43450 - 43496,July 13, 2000).

    For recovery purposes, the FWS final rule combines the Northeast region with existingwolf populations in the Great Lakes, eventhough the two regions are geographically

  • Northeastern Gray Wolf DPS - 1 April 2003 12

    disconnected. Since the FWS considerswolves to be sufficiently recovered inMinnesota, Wisconsin and Michigan,federal protection will be downgraded in allstates within the proposed Eastern GrayWolf DPS–despite the fact that wolves areabsent from most states in this region.Further, wolf recovery in the states wherewolves do currently exist, will continue tobe hampered by curtailment of the species’range, negative human attitudes, andenvironmental stochastity. Becausemetapopulations, such as those of the graywolf, are less likely to become extinct whenthere are more local populations established(Gotelli 1998), the FWS should promoterecovery over a larger geographic area andstrive to protect dispersing individualsrecolonizing new areas. For these reasons,we propose to create a Northeastern GrayWolf DPS.

    C. DPS and ESA CriteriaUnder the FWS DPS policy, 61 Fed. Reg.4722-25 (Feb. 7, 1996), three elements areconsidered in a decision of whether to list aDPS as threatened or endangered under theESA. First, the population must be discretebased on one of the following criteria: (1)the population is markedly separated fromother populations of the same taxon, or (2)the population is delimited by internationalgovernmental boundaries. Second, apopulation’s significance can be establishedbased on the following factors: (1)persistence of the DPS in an ecologicalsetting unusual or unique for the taxon, (2)evidence that loss of the DPS would resultin a significant gap in the range of the taxon,(3) evidence that the DPS represents theonly surviving natural occurrence of a taxonwithin its historic range, or (4) evidence thatthe discrete population segment differsmarkedly from other populations of the

  • Northeastern Gray Wolf DPS - 1 April 2003 13

    species in its genetic characteristics. Lastly,if a population is determined to be bothdiscrete and significant and therefore a“species” under the ESA, its status asendangered or threatened is then evaluated. The standard for listing species under theESA is fairly straight forward, 16 U.S.C. §1533 (a)(1); 50 C.F.R. § 424.11. The ESArequires the Secretary to determine, "solelyon the basis of the best scientific andcommercial data available..." whether aspecies is endangered or threatened based onany one or a combination of five factors: (1)the present or threatened destruction,modification, or curtailment of its habitat orrange; (2) overutilization for commercial,recreational, scientific, or educationalpurposes; (3) disease or predation; (4) theinadequacy of existing regulatorymechanisms; and (5) other natural ormanmade factors affecting its continuedexistence.

    D. Overview and Current IssuesCurrently, the northeastern states arecontained within an Eastern Gray Wolf DPSand are part of this region’s recovery plan.In accordance with the 1992 Recovery Planfor the Eastern Timber Wolf recoverycriteria, at least two viable populations ofwolves within the eastern range of the graywolf must be re-established in order toproceed with any reclassification or de-listing objectives. The Petitioners believethat wolves present in Wisconsin andMichigan are too closely associated andgeographically proximate to threatened wolfpopulations in Minnesota to be considered aseparate and viable population. Althoughwe support the down-listing of the gray wolfin portions of the Eastern Gray Wolf DPS(Wisconsin, Michigan), we cannot supportexpanding these criteria to states outside therealm of the current recovery area.

  • Northeastern Gray Wolf DPS - 1 April 2003 14

    Inclusion of the northeastern states ofMaine, New Hampshire, Vermont and NewYork in the delisting process withoutadditional and site-specific recovery goals inthese states is unacceptable.

    Delisting of the Eastern Gray Wolf DPS without the creation of a separate NortheasternDPS would leave an area about 26 millionacres devoid of wolves, a region thathistorically maintained healthy and viablewolf populations. Gray wolves are veryunlikely to recolonize the region on theirown because of the distance (about 500miles) and the multiple anthropogenicbarriers (highways, farmland, and urbandevelopment) between the northeasternstates and existing wolf populations in theGreat Lakes states. The northern boundaryof the Northeast region is legally isolatedalong the international border with Ontarioand Quebec, Canada. The closestsignificant wolf populations to the Northeastare found in Algonquin Park, Ontario andthe Laurentides region in Quebec. The wolfpopulations in these regions of Canada arequite low at this time and are continuallyunder pressure from lenient hunting andtrapping regulations, so probably would notprovide a source population for naturalrecolonization.

    Delisting the Eastern Gray Wolf DPS would remove federal protection for gray wolvesand give responsibility for their continuedprotection to the individual states includedunder that DPS region. None of the fournortheastern states have any laws in place toprotect wolves if federal ESA protectionsare lifted nor do any of these states have anymanagement plans in place for wolves ifthey ever to return to the region. In Maine,eastern timber wolves (Canis lupus lycaon)are listed as a “Species of Special Concern”(Maine Dept. of Inland Fisheries andWildlife, 2002) but have no designated state

  • Northeastern Gray Wolf DPS - 1 April 2003 15

    endangered or threatened classification. InMay, 2001, the Maine state legislature ruledthat no reintroduction of gray wolves (Canislupus) can take place without the consent ofboth houses of the legislature and theCommissioner of Inland Fisheries andWildlife. It also declared that the U.S. Fishand Wildlife Service must allow thelegislature to prohibit the reintroduction ofwolves into Maine if it so decides (LD-736,Maine Legislature). In New Hampshire,legislators passed a bill in 1999 making itillegal to introduce any wolf populations tothis state (HB 240 - New HampshireLegislature). At the same time, the Houseissued a Joint Resolution asking the U.S.Fish and Wildlife Service to prohibit thereintroduction of wolves in the Northeast,including New Hampshire. The gray wolf isnot protected by any New Hampshire statewildlife laws. In New York, the gray wolf islisted as a species that no longer occurs in awild state within the State. There are noendangered or threatened designations forthis species in New York. Two countiesbordering Adirondack Park have banned thereintroduction of gray wolves within theirboundaries. In Vermont, the gray wolf isnot listed under the Vermont EndangeredSpecies Law as endangered or threatened,nor is it listed as a rare or uncommon animalunder the State Non-game and NaturalHeritage Program. Several efforts have beenmade within the Vermont legislature to banthe reintroduction of wolves into this state,but no law has yet been passed. While thesestate laws are relatively meaninglessbecause they would be superceded byfederal authority, they nonethelessdemonstrate the prevailing negative attitudestowards wolves by state officials.

    Little can be done to significantly increase the amount of suitable habitat available forwolf recovery in the lower 48 states.

  • Northeastern Gray Wolf DPS - 1 April 2003 16

    Consequently, the best that can be done forthe wolf is to make the most use of whathabitat remains. The only way to maximizethe species’ chances of long-term survival isto utilize remaining habitat to the extentpossible to restore populations that canprovide adequate representation, resiliency,and redundancy (Shaffer and Stein 2000).Representation refers to establishingpopulations across the full array ofappropriate potential habitats. Resiliencyrefers to maintaining populations in eachhabitat at levels large enough to survive anynegative consequences of demographicstochasticity and inbreeding. Redundancyrefers to providing several populations ineach habitat type as a hedge against extremeenvironmental events (Shaffer and Stein2000). Wolf populations should beestablished in remaining habitat based onthese principles in order to maximize thelong-term viability of the gray wolf in thelower 48 states. In practice, the abovewould call for a minimum of two (preferablythree or more) populations of not less thanseveral hundred wolves in each ecologicallyor environmentally distinct area of itsformer range.

    With the states showing no indication of restoring wolves, and considering theimportance of the Northeast to overall wolfrecovery, the only solution for recovery of aviable long-term population of gray wolvesis through continued federal oversightincluding the establishment of aNortheastern Gray Wolf DPS. The FWSshould develop a comprehensive recoveryplan for this region and follow this actionwith whatever steps are deemed necessary toencourage the restoration of this species.

    II. NATURAL HISTORY

    A. Description of the Species

  • Northeastern Gray Wolf DPS - 1 April 2003 17

    Physical description – Gray wolves are thelargest member of the dog family Canidae(Mech 1970). Female average weightranges from 80 - 85 lbs. and males averagefrom 95 - 100 lbs. (Mech 1970), thoughconsiderable clinal variation in size and peltcolor exists from the Arctic to centralMexico (Young and Goldman 1944). Theheaviest recorded wolf was a 175- poundmale from east-central Alaska, though malesseldom exceed 120 lbs. and females areseldom over 100 lbs. (Mech 1970).

    Pack Behavior – Wolves live, travel, andhunt in packs averaging four to sevenanimals, consisting of an alpha, or dominantpair, their pups, and several othersubordinate or young animals. The alphafemale and male are the pack leaders,tracking and hunting prey, choosing densites, and establishing the pack's territory(Mech 1970). Wolves prey mainly onungulates, such as deer, elk, moose, caribou,bison, bighorn sheep and muskoxen. Theyalso eat smaller prey such as snowshoe hare,beaver, rabbits, opossums and rodents. Wolves will prey on livestock, althoughwild prey are their preferred food (Mech1970).

    Wolf pups romp and play fight with each other from a very young age. Scientistsspeculate that even these early encountersestablish hierarchies that will help determinewhich members of the litter will grow up tobe pack leaders. All adults share parentalresponsibilities for the pups. They feed thepups by regurgitating food for them from thetime the pups are about four weeks old untilthey learn to hunt with the pack. Pupsremain with their parents for at least theirfirst year, while they learn to hunt. Duringtheir second year of life, when the parentsare raising a new set of pups, young wolvescan remain with the pack, or spend periods

  • Northeastern Gray Wolf DPS - 1 April 2003 18

    of time on their own. Frequently, they returnin autumn to spend their second winter withthe pack (Mech 1970).

    By the time wolves are two years old, they leave the pack permanently to find mates andterritories of their own. Not all the pups in alitter live to the age of dispersal. Biologistshave determined that only one or two ofevery five pups born live to the age of 10months, and only about half of thoseremaining survive to the time when theywould leave the pack and find their ownmates. Adult wolves, on the other hand,have fairly high rates of survival. A sevenyear-old wolf is considered to be relativelyold, and the maximum lifespan is about 16years (Young and Goldman 1944).

    Reproduction.– The alpha pair mate inJanuary or February and give birth in spring,after a gestation period of about 65 days.Litters can contain from one to nine pups,but usually consist of around six. Pups haveblue eyes at birth and weigh about onepound. Their eyes open when they are abouttwo weeks old, and a week later begin towalk and explore the area around the den. Wolf pups grow rapidly, reaching 20 poundsat two months. A wolf pup is the same sizeas an adult by the time he or she is about ayear old, and reaches reproductive maturityby about two years of age (Mech 1970).

    Communication.– Wolves communicatethrough facial expressions and bodypostures, scent-marking, growls, barks,whimpers and howls. Howling can meanmany things: a greeting, a rallying cry togather the pack together or to get ready for ahunt, an advertisement of their presence towarn other wolves away from their territory,spontaneous play or bonding. Pups begin tohowl at one month old. The howl of the wolfcan be heard for up to six miles. When

  • Northeastern Gray Wolf DPS - 1 April 2003 19

    wolves in a pack communicate with eachother, they use their entire bodies:expressions of the eyes and mouth, set of theears, tail, head, and hackles, and generalbody posture combine to expressexcitement, anxiety, aggression, oracquiescence. The wolf’s acute hearing andexceptional sense of smell—up to 100 timesmore sensitive than that of humans—makethem well-adapted to their surroundings andto finding food (Mech 1970).

    Wolves wrestle, rub cheeks and noses, nip, nuzzle, and lick each other. They also leave"messages" for themselves and each otherby urinating, defecating, or scratching theground to leave scent marks. These markscan set the boundaries of territories, recordtrails, warn off other wolves, or help lonewolves find unoccupied territory. No oneknows how wolves get all this informationfrom smelling scent marks, but it is likelythat wolves are very effective atdistinguishing between many similar odors.

    B. TaxonomyConfusion and disagreement exist over theidentity and taxonomy of the wolf thathistorically occupied the Northeast region ofthe United States (Fascione et al. 2001). Goldman (1937) classified the eastern timberwolf as Canis lupus lycaon, a smallersubspecies of gray wolf, and for years itshistoric range was thought to be thenortheastern United States as far west as theGreat Lakes states and north into southernOntario and Quebec (Goldman 1944, FWS1992, Nowak 1995). In 2001, Wilson et al.,based on DNA analysis of several specimens,hypothesized that C. l. lycaon had a commonorigin with the red wolf (Canis rufus), aspecies whose historic range has long beenconsidered to be the southeastern U.S. as farwest as Texas and as far north asPennsylvania (Nowak 1995). The ranges of

  • Northeastern Gray Wolf DPS - 1 April 2003 20

    the eastern wolf and the red wolf would haveoriginally overlapped in the mid-Atlanticstates (Nowak 1995). The most recentmorphological evidence, presented by Nowak(2002), speculates that C. l. lycaon might haveresulted from the hybridization between C.rufus and C. lupus nubilus, the larger wolfthat formerly inhabited the western U.S. andmuch of Canada. The regions in far northernOntario contain predominantly western graywolf, but the boundary between the easternwolf and the western gray wolf is not wellestablished (P. Wilson, personalcommunication).

    To confound the issue, the eastern timber wolf has shown a tendency to hybridize withcoyotes (Canis latrans) (Wilson et al. 2000).Coyotes were historically absent from theEast but moved in from the West by the 1930s(Parker 1995). Genetic testing of the relativelylarge coyotes from the Adirondack Park andcentral New York indicates a history ofinterbreeding with wolves. The degree ofwolf genetic material varies across thesesamples, with some being more “wolf-like”than others (Chambers 2000). Genetic testingon northern New England coyotes showsinterbreeding as well, though more samplingis needed (Fascione et al. 2001). In theFrontenac Axis region, southeast ofAlgonquin Park, a slightly larger canidcommonly called the “Tweed wolf,” isprobably a hybrid containing more wolf genesthan coyote genes (Edwins et al. 2000). Acoyote-wolf mix is commonly found west ofAlgonquin Park (Wilson et al. 2000). Theresult is a canid genetic gradient in thenortheastern U.S. and southeastern Canada,with a mix of eastern timber wolf, westerngray wolf and coyote genes (Fascione et al.2001).

    Given the radical changes that have occurred in the Northeast ecosystem since colonialtimes, and the lack of remaining physical

  • Northeastern Gray Wolf DPS - 1 April 2003 21

    evidence of the presence of wolves, it isdifficult to determine which species may havebeen present historically. For now, scientistscan only speculate, but it is generally acceptedthat the northeastern U.S. was primarily amoose-caribou ecosystem before Europeansettlement (D. Harrison, personalcommunication). It is questionable whetherthe deer-adapted eastern timber wolf wouldhave thrived in this environment, indicatingthat perhaps both canids— the larger graywolf and the smaller eastern wolf—mighthave inhabited the northeast at various points(P. Wilson, personal communication).

    Because of the fluid nature of gray wolf taxonomy and the FWS’s goal to affordprotection to all gray wolves south of theU.S.-Canada border, the FWS listed all graywolves as threatened (Minnesota) orendangered (remaining 47 contiguous statesand Mexico) at the species (Canis lupus)level in 1978 (43 Fed. Reg. 9607-9615,March 9, 1978). In its 2000 proposal toreclassify gray wolves by DPSs, the FWSstates: “We recognize that gray wolftaxonomy at the subspecies level is subjectto conflicting opinions and continuingmodification. For this reason, we will notbase our gray wolf recovery efforts on anyparticular portrayal of gray wolfsubspeciation. Instead we have identifiedgeographic areas where wolf recovery isoccurring or is feasible, and we will focusrecovery efforts on those geographicentities, regardless of the subspecificaffiliation of current or historical graywolves in those areas.” The FWS also statesthat “it is likely that a separate form of thegray wolf historically occupied thenortheastern United States and adjacentCanada. Establishing a Northeastern DPSmaximizes the ability of the Service, States,and Tribes to reestablish this form, or itscurrent-day equivalent. The wolves in

  • Northeastern Gray Wolf DPS - 1 April 2003 22

    Canada...are thought to be taxonomicallyand genetically similar to the wolves thatonce populated the northeastern UnitedStates,” 65 Fed. Reg. 43451-43452 (July 13,2000). Any wolf restoration in the Northeastwould have to include a detailed analysis ofthe best source population as part of therecovery process.

    C. Historical Distribution in theNortheastThe historic range of the eastern wolf, oneof the smallest subspecies of Canis lupus,once extended throughout the entirenortheastern United States, from HudsonBay to northern Florida and west intoMinnesota (Mech 1970). As Europeansettlers first arrived and began to “tame” theforests of the Northeast by cutting, burningand plowing, the persecution of wolves wasset into motion (McKibben 2000).Beginning in 1630 in the Massachusetts BayColony, a system of government-sponsoredwolf bounty payments was developed,which pervasively spread throughout theNortheast, until all of these large predatorshad been exterminated. The last report of awolf in Maine was approximately 1880(Maine Dept. of Inland Fisheries andWildlife) and the last known northeasternwolf was killed in upstate New York in1897 (Fascione et al. 2000).

    Today, the eastern wolf can only be found in parts of Quebec, Ontario, and the GreatLakes states (Mech 1970). A fewobservations of wolves or wolf-like animalshave been made in recent years throughoutthe Northeast, including Vermont, NewYork and Maine, but most of these reportsare anecdotal and have not been readilyverified. Recently, a large canid was killedin northern New York and initial DNAtesting indicates that the animal was a wolfand might have originated in the Great

  • Northeastern Gray Wolf DPS - 1 April 2003 23

    Lakes region (Inslerman, pers. comm) Awolf killed by a Quebec trapper in 2001,was apparently traveling with a few otherwolves just north of the Maine-Canadaborder (Associated Press, 2002). In Maine,there have been several reported sightings oflarge canids (Smith, personalcommunication), indicating the possibilitythat wolves might be present in this state. Alarge female wolf was shot in 1993,although physical and behavioral evidencesuggested the animal had a history ofcaptivity. In 1996, an 86-pound canid wascaptured; genetic testing provedinconclusive (Fascione et al. 2000, MaineDept. of Inland Fisheries and Wildlife). Noevidence has yet been found of anestablished wolf population in Maine orelsewhere in the Northeast. Indeed, severalstudies have indicated that the barriers awolf would have to cross to establish apopulation in the Northeast are almostinsurmountable (Harrison and Chapin 1997).For the few individuals that might actuallymake it across the border, the result wouldlikely be either persecution by humans orgenetic swamping with the larger easterncoyote (Wydeven et al. 1998.)

    III. WOLF ECOLOGY

    A. The Role of the Wolf as a TopCarnivoreEnergy transfer between trophic levels.–Thegray wolf, along with other top predatorssuch as the bear, cougar, and coyote, helpregulate prey populations such that alandscape may support multiple trophiclevels in a healthy ecosystem. Whenpopulations of large herbivores are kept incheck by predators, the amount of primaryproduction available to smaller animalsincreases, allowing for increasedbiodiversity. A basic principle of ecology

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    states that only ten percent of the vegetableor prey’s biomass is retained in the biomassof the grazer or predator, respectively. Forexample, only one tenth of the vegetationconsumed by an ungulate in a year will beassimilated into the molecular buildingblocks of the ungulate itself. The otherninety percent is converted into energy usedfor metabolic processes and is eventuallylost to the environment. Large browsingherbivores such as deer and elk require agreat quantity of woody stems, herbaceousplants and lichens to fuel their metabolismsand to reproduce. Left uncontrolled, largeherbivores that require such a large amountof primary productivity to survive willdeplete a landscape of its primaryproductivity. Without predators to regulatethe number of ungulates, ecosystems aresimplified. Ungulate population explosionssimplify the food web and ultimately reducebiodiversity. (Terbough et al. 1999).

    Mesopredator release.– Another harmfulecological effect of removing wolves froman ecosystem is the expanded niche formesopredators such as the coyote, raccoonand fox. Mesopredator release can beresponsible for decreased biodiversitybecause mesopredators tend to prey heavilyon a wide variety of smaller animals,including songbirds. Coyotes and othermesopredators are generalists and cansurvive after they deplete a preferred foodsource. Normally, when wolves are present,coyote populations are suppressed byterritorial aggression and by predation(Crabtree and Sheldon 1999), relievingsmall mammals and birds from the risk ofcoyote predation (Fischer 1998, Wilkenson1997). Additionally, wolves increase theamount of carrion available in an ecosystem,potentially benefiting scavenger speciessuch as bear, foxes, weasels, and raptors.

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    Availability of carrion may increasebiodiversity, as it provides an alternate foodsource for generalist mesopredators(Crabtree and Sheldon 1999).

    Regulation of prey genetic health.– Inaddition to the role carnivores play inincreasing biodiversity, they also improvethe gene pool of their prey species over timeby culling genetically inferior individuals. The gray wolf, in particular, exerts thispositive force on the prey gene pool, as itoften chases after a herd of ungulates until aslower animal is left behind. This“coursing” technique may more effectivelyreduce the chance of a genetically weakanimal from reproducing than other huntingstrategies (Mech 1970). A cougar, bycontrast, will usually hide in a hunting beduntil its prey comes within springingdistance. The prey in this case is almost aslikely to be healthy as it is to be weak. Because all carnivores occupy a distinctniche in an ecosystem and employ differenthunting strategies, they play a unique role inthe management of the lower trophic levels. The wolf, however, may have a more directeffect on prey gene flow than othercarnivores.

    Wolf behavior and population dynamics.–Wolves hunt, live, and travel in packsaveraging four to seven animals consistingof an alpha, or dominant pair, their pups,and several other subordinate or younganimals. The alpha male and female are thepack leaders, whose role it is to track andhunt prey, choose den sites and to establishthe pack’s territory (Mech 1970). Wolvesprey primarily on ungulates such as deer,elk, moose, caribou, bison, bighorn sheep,and muskoxen depending on the distributionof these prey species. They also eat smallerprey such as lagomorphs, beaver, opossums,

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    and rodents. Wolves will take livestock,though wild prey is their preferred food(Mech 1970). Pups will remain with theirparents for at least their first year while theylearn to hunt. During the second year whenparents may be raising a new set of pups,young wolves can remain with the pack orspend time on their own, frequentlyreturning in autumn to spend their secondwinter with the pack. Biologists havedetermined that only one or two of everyfive pups born live to the age of ten months,and only about half of those survive to theage of dispersal (Young and Goldman1944).

    By the time wolves are two years old, young wolves leave the pack permanently to find mates and territories of their own. Becausedispersal is an important stage in wolfdevelopment, wolf populations can beviewed as a dynamic mosaic of populationsor a metapopulation (65 Fed. Reg). Metapopulations are simply a group ofsmaller populations linked by immigrationand emigration (Levins 1970). The basicstructure of wolf populations at small andlarge scales allows wolves to be described inthis way. On a small scale, wolves travel infamily units, but depend on dispersal to findmates and new territories. On a large scale,especially in the Northern Rocky Mountainrecovery areas, wolves disperse from onepopulation center to another, and wildlifeofficials expect individuals will interbreedwith members of populations in distant areas(65 Fed Reg).

    B. Ecological Importance of Dispersaland Management ConsiderationsGray wolf dispersal.– Dispersal is afundamental aspect of wolf ecology thatshould be addressed when designing wildlifereserves or planning for wolf management.Pups remain with their parents for at least

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    their first year while they are learning tohunt and then both sexes disperse with thesame frequency (Gese and Mech 1991). During the second year when parents maybe raising a new litter of pups, young wolvescan remain with the pack or spend time ontheir own making predispersal forays (Geseand Mech 1991), frequently returning inautumn to spend their second winter withthe pack. Adults disperse as well asyearlings and pups, but not with the samefrequency. When they do disperse, they areoften more successful than younger wolvesin finding vacant territory within a shorterdistance of their natal territory (Gese andMech 1991). Some researchers estimate thata wolf can run as fast as 40 miles an hour.Wolves have been known to travel 120miles in a day, but they usually travel anaverage of 10 to 15 miles a day (Mech1970). Wolf dispersal rate (and overall wolfabundance) is negatively correlated to preyabundance (Fuller 1989, Gese and Mech1991, Messier 1985, Ballard et al. 1987,Peterson and Page 1988, Hayes andHarestad 2000). The number of wolveswithin a region can be estimated by findingthe number of prey in the area, according tothe prey biomass: wolf index developed byFuller (1989) and tested by other studies(Hayes and Harestad 2000).

    Movements and population fluctuations of prey are the major causes of wolf dispersaland the determining factors determiningdispersal distance. Even when prey densityis adequate, however, most wolves dispersefrom their natal pack territories as pups oryearlings (Gese and Mech 1991). Factorsother than prey density that can influencedispersal are social strife within a pack andthe unavailability of genetically unrelatedmates. The search for mates may be one ofthe most important reasons for dispersalbecause wolves avoid inbreeding to the

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    point where they will not breed if theycannot locate a mate from a sufficientlydistant pack. For this reason the number ofpacks occupying an area partiallydetermines dispersal distance and dispersalsuccess. If there are very few packs in anarea, a dispersing wolf might not be able tolocate a mate that is genetically unrelated,and therefore may have to disperse longdistances to find a suitable mate.

    Landscape characteristics of dispersalpaths.– Because of this close relationshipbetween wolves and their prey, designingwildlife reserves for wolves is difficult. Ungulates, the most common prey speciesfor wolves (Mladenoff et al. 1995), are oftenmore dense at the edges of wilderness areasthan they are within the boundaries ofpreserves because disturbed areas usuallyprovide more browse. Although they mayhave habitat preferences when prey densityis high and they are able to select certainhabitat characteristics over others, wolvesare not specific to any one particular habitattype and are able to survive in almost anytype of landscape as long as there isadequate prey and contact with humans isminimal.

    Dispersal mortality.– Because wolves willstray beyond the boundaries of protectedhabitat if their prey moves to other areas,and because prey species such as deer oftenmove across the landscape seasonally,wolves can be expected to travel seasonallyas well. As wolves follow their prey intodisturbed areas where deer are able to findmore browse and cover from winterconditions, they may be moving towardhuman population centers (Haight et al.1998). This behavior poses a severe threatto individual wolves and to overallpopulation numbers within protected areas.

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    Dispersal mortality greatly affects overallpopulation size and probability of localextinction (Fritts and Carbyn 1995). Ifwolves within protected areas must strayinto agricultural or semi-rural regions inorder to find prey, they may not survive longenough to return when prey moves onceagain into protected wolf habitat.

    If human threats to wolves were limited by federal protection, dispersal patterns ofwolves would not be such a danger tooverall populations. However, wolves areoften extremely vulnerable to human-causedmortality because they are able to adapt toalmost any type of environment, includingareas with relatively high humanpopulations. A dispersing wolf mayencounter many dangers it had not beenexposed to while living within a protectedhabitat. One of the biggest threats to adispersing wolf may be human-causedmortality in the form of illegal taking andlegal depredation control measures(Mladenoff et al. 1995, Mech 1970, Fuller1989).

    Because wolves are able to adapt to almost any type of environment, including areas withrelatively high human populations, wolvesdispersing out of protected reserves willlikely encounter humans if they do notquickly settle. Mortality caused by humansincludes accidental killing by motorvehicles, legal depredation controlmeasures, and illegal takings. Dispersalmortality greatly affects overall populationstability, especially if there are fewimmigrants into the population and if theinitial population size is small (Haight et al.1998). Furthermore, dispersal has beenfound to be a key factor limiting populationgrowth (Hayes and Harestad 2000).

    As wolves recover within the protected boundaries of Yellowstone National Park,individual wolves dispersing into

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    agricultural areas are exposed to increasedthreats posed by humans. Increased wolfmortality during and after wolf recoveryprograms illustrates that the greatest numberof human-caused mortalities occurs at ornear the boundaries of protected reserveswhere wolves disperse into more fragmentedhabitat (Hayes 2001, Mladenoff et al.1995). One reason for this trend is that thepresent reserves in the Midwest and in thenorthern Rockies are in close proximity to,if not completely surrounded by, agriculturalland. Human attitudes of those surroundingthe protected wolf habitat can determinewhether or not wolves will successfullydisperse (Fritts and Carbyn 1995). Whereasthe reserve boundary areas adjacent toagricultural land might actually representsome of the most favorable habitat forwolves due to the presence of foragingungulates at field edges, it also introduceswolves to another, more dangerous type ofprey: domestic livestock. Cattle ranchingand dairy farming puts pressure on wolfpopulations, especially if there are mosaicsof forest interlaced with ranching and farmlands because wolves are enticed to explorefurther away from their natal territorieswhen dispersing (Fritts and Carbyn 1995).

    Human actions account for approximately 80-90 percent of all wolf mortalities (Weaver2001). Between 1995 and July of 2002, 64of the 118 known wolf mortalities within theYellowstone wolf population wereconfirmed to be human-caused, three wereunknown and the remainder were naturaldeaths. Illegal shooting or poisoning wasresponsible for 10 deaths, legal controlclaimed 37 wolves, road kills caused another12 deaths and five of the remaining losseswere human-related (Maughan 2002). Similarly, when wolf populations increasedin Minnesota between the years of 1988 and1993, the number of wolves killed through

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    government-sponsored programs fordepredation control increased by 223percent (Mech 1995).

    Dispersal distance.– The risk of mortalityduring dispersal increases as dispersaldistance increases. There is muchvariability in wolf dispersal distances. While some wolves travel very shortdistances before settling, most wolves seemto travel several hundred kilometers beforefinding suitable vacant territory (Gese andMech 1991, Wabakken et al. 2001). Wolvesare capable of dispersing hundreds of miles,with the longest known dispersal exceeding550 miles (Fritts 1983). Gese and Mech(1991) found that the mean dispersaldistance for 316 dispersing gray wolves was47.8 miles, with a range of 5-220 miles. Dispersal distance is important to thesurvival probability of a wolf populationbecause individual wolves have a greaterchance of survival if their dispersal distanceis short (Weaver 2001). Long dispersaldistances increase the risk of mortality dueto conflicts with humans or starvation andreduce the chance that a disperser will settleand find a mate.

    The factors that initiate dispersal are the same that ultimately determine the distance a

    wolf will disperse: social interactionsbetween wolves, availability of mates,spatial distribution of available territories,and prey density (Gese and Mech 1991,Hayes and Harestad 2000, Fuller 1989). Ifconditions are favorable, a dispersing wolfmay only need to travel to nearby areas tosuccessfully establish a new territory. However, if a wolf travels through areaswith low prey abundance and few potentialmates, it will search much longer beforelocating food and a mate, becomingincreasingly vulnerable to human-caused ornatural mortality (Gese and Mech 1991,

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    Fritts and Carbyn 1995, Mladenoff et al.1995 Mech 1995, Fuller 1989).

    C. Minimum Viable Population SizeAlthough there is some debate over what theminimum viable population size is for graywolves and how much gene flow throughimmigration is required to maintain geneticdiversity, most wildlife ecologists agree thatthe probability of population extinction ishigh when the number of individuals is low(Franklin 1980, Thomas 1990, Wabakken etal. 2001). Minimum viable population isdefined as the number of individualsnecessary to insure the population’s survivaland genetic diversity over a specified timeperiod, regardless of harsh environmentalconditions, fluctuating prey base, successionof forest plant species, and dispersal (Frittsand Carbyn 1995).

    If a population of wolves is isolated without frequent genetic exchange with immigratingwolves, an ideal population size should beanywhere from several hundred to twothousand individuals (Franklin 1980,Wabakken et al. 2001, Lande andBarrowclough 1987, Soule 1980, Thomas1990). Five to six hundred individuals, orapproximately 100 breeding pairs, may besufficient to maintain genetic diversitywithin a population closed to immigration(Wabakken et al. 2001, Soule 1980, Frittsand Carbyn 1995), but dispersal andenvironmental stochastity may strain apopulation of this small size (Franklin 1980,Thomas 1990).

    For a relatively small population of 280 to 300 wolves in Italy, which is approximatelythe same size of that in the northernRockies, population viability analysisshowed that populations of this size arevulnerable to extinction if there are anydramatic changes in percent adult mortality. If adult mortality increased beyond 10

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    percent, the model showed that thepopulation would likely become extinctwithin 60 to 100 years (Ciucci and Boitani1991). Small populations that barely meetthe minimum viable populationrequirements such as the Italian populationand that of the northern Rockies are morevulnerable to extinction when the mortalityrate increases even by a small amount(Wabakken et al. 2001). Also, these smallpopulations may experience inbreedingpressure, as there is probably a significantdecline in genetic variability over time(Wabakken et al. 2001).

  • Northeastern Gray Wolf DPS - 1 April 2003 34

    IV. NORTHEASTERN GRAY WOLF DPS PETITION PROPOSAL

    A. Distinct Population Segments underthe Endangered Species ActDiscreteness and significance.–Individualpopulations of a species should be managedseparately if there is sufficient reason tobelieve that there are factors threateningtheir persistence, according to the FWS DPSpolicy (61 Fed. Reg. 4722-25 Feb. 7 1996). To be designated as a DPS, a populationmust be discrete based on the fact that thepopulation is markedly separated from otherpopulations of the same taxon, or it isdelimited by international or governmentalboundaries. If a population is determined tobe discrete, then it must meet one or anycombination of the following factors toprove that it is significant to the overalltaxon: 1) the species is persisting in anecological setting that is unusual or uniquefor its taxon, 2) there is evidence that loss ofthe particular population would result in asignificant gap in the range of the taxon, 3)there is evidence that the populationrepresents the only surviving naturaloccurrence of a taxon within its historicrange, and 4) there is evidence that thepopulation differs markedly from otherpopulations of its species in its geneticcharacteristics.

    Determination of federal protections.– If apopulation is determined to be discrete andsignificant based on these criteria, theSecretary will then determine the level ofFederal protections given to the DistinctPopulation Segment. Congress then cangive a mandate to the FWS to developrecovery plans where appropriate and wherechance of success is high. Recovery effortsshould give priority to areas with adequateresources. The Endangered Species Actrequires the Secretary of the Interior to

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    determine if a species should be listed asendangered or threatened according towhether they meet one or any combinationof the following five criteria (16U.S.C.§1533 (a) (1); 50 C.F.R. § 424.11 ): 1) there is present or threatened destruction,modification, or curtailment of its habitat orrange, 2) the species is over-utilized forcommercial, recreational, scientific, oreducational purposes, 3) the species isgreatly threatened by disease or predation,4) existing regulatory mechanisms areinadequate, and 5) other natural or manmadefactors are affecting its continued existence.

    B. DPS Boundaries and HabitatDescriptionTo build a landscape scale metapopulation itis necessary to manage each distinctsubpopulation separately because of thedifferences in regional landscape attributessuch as local climate, geology, and preybase. Therefore, the northeastern UnitedStates and the few wolves that may inhabitthis vast and favorable habitat should bedesignated as a DPS under the ESA. Theregion that should be defined by theNortheastern Gray Wolf DPS includes thestates of Maine, New Hampshire, New Yorkand Vermont. Several areas of core habitat,containing few roads and low humanpopulation density are located within theproposed DPS (Mladenoff and Sickley1998, Harrison and Chapin 1997).

    The Northeastern DPS, encompasses an area know as the Northern Forest, a 26 millionacre tract of land, comprised of remote,pristine lakes, rugged mountain ranges, andthe headwaters of a few large eastern rivers.This region stretches from Maine’s St. CroixRiver westward through the WhiteMountains of New Hampshire, the GreenMountains of Vermont and into New York’sAdirondack Mountains.

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    Maine, the largest of the New England states, stretches from the Atlantic Ocean in theeast to Canada in the north and NewHampshire to the west and south, and isapproximately 89 percent forested. Most ofthe topography is relatively flat with a meanelevation of 600 feet, although Baxter StatePark hosts one of the highest points in theregion, 5268-foot Mt. Katahdin. Maine canbe divided into three major geographicalareas: the coastal lowlands, an areacharacterized by sandy beaches, small inlets,salt marshes and tidal creeks; the easternNew England uplands, a region that risesfrom sea level to about 2000 feet in the westand is marked by lakes, fast streams, andfertile soil; and the White Mountains, anextension of New Hampshire’s range thatcover a portion of northwestern Maine. TheAllagash Wilderness Waterway stretchesalong a 92-mile corridor of lakes and riversand connects several large public reservedland units in northern Maine.

    New Hampshire, one of the smallest states in the U.S., features rugged mountains, clearlakes, and a sandy coastline. With an meanelevation of 1000 feet, this state is borderedby Vermont on the west, the Atlantic Oceanand Maine on the east, Canada to the northand Massachusetts on the south. LikeMaine, New Hampshire is influenced bythree main geographic areas: the CoastalLowlands, which characterize thesoutheastern section of the state and extendfrom 15 to 20 miles inland; the Eastern NewEngland Upland, a region comprised of thefertile and hilly Merrimack River Valley, theHills and Lakes region that includes most ofthe state’s major lakes, including the largest,Lake Winnipesaukee, and the ConnecticutRiver Valley, a region comprised of fertilefarmland and hardwood forests; and theWhite Mountain Region, which covers thenorthern portion of the state and includes

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    rugged mountains, narrow valleys and theWhite Mountain National Forest. MountWashington, part of the Presidential Rangeand the highest point in New England, is6,288 feet high and home to some of theworst weather in the world.

    Vermont, the Green Mountain State, is known for its fertile valleys and picturesquemountain ranges. Bordered by Canada onthe north, New York to the west, NewHampshire to the east and Massachusetts tothe south, Vermont has an average elevationof 1000 feet and is approximately 80 percentforested. The Green Mountains, animportant source of minerals such as graniteand marble, cover most of central Vermont,and are home to the highest peaks in thestate and to the Green Mountain NationalForest. The northern region of the state isknown as the northeast highlands, a regionthat is characterized by granite mountainsthat reach heights of 2,700 to 3,330 feet andare divided by swift flowing streams. LakeChamplain comprises much of thenortheastern part of the state and thelowland areas surrounding the lake aredominated by fertile agricultural lands. Insouthern Vermont, the Taconic Mountainrange stretches in to Massachusetts and is aregion of rolling hills, lakes and streams.Most of eastern Vermont is covered by thewestern New England upland, a geographicland area that is covered by the fertilelowlands of the Connecticut River Valleyand populated with many lakes in the north.

    New York, the largest of the proposed DPS states, has the most diverse geography.Bordered by two of the Great Lakes to thenorth and west, by Canada to the north andthe Atlantic Ocean to the east, New York’stopography averages approximately 1000feet in elevation. In the northern part of thestate, between Lake Champlain and LakeOntario, stand the highest and most rugged

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    mountains in the state, the AdirondackMountains, that are part of the 6.1 millionacre Adirondack State Park. Mt. Marcy,located in the High Peaks region, has anelevation of 5,344 feet. South of theAdirondack mountains, lies the Hudson-Mohawk lowlands and in the southeast, isthe Atlantic coastal plain. In the westernpart of the state, west of the Hudson River,are the Appalachian Highlands, whichinclude the Catskill Mountains and theFinger Lakes region. The St. Lawrence-Champlain lowlands can be found on theshores of Lake Ontario and runningnortheast along the St. Lawrence River andthe Canadian border.

    Two distinct forest community types are represented within the DPS region (Marchand1987, Kricher 1998, Thompson andSorenson 2000). The Northern hardwoodforest, generally present at elevations lessthan 3000 feet, is dominated by yellow birch(Betula lutea), sugar maple (Acersaccharum) and American beech (Fagusgrandifolia) with a well-developedunderstory of striped maple (Acerpennsylvanicum) and Hobble bush(Viburnum alnifolium), including manywildflower and fern species. Easternhemlock (Tsuga canadensis) and white pine(Pinus strobus) can also be found inabundance within this community type. Thenorthernmost part of the DPS is includedwithin a Boreal Forest community type,generally found above 3000' elevations. Thisarea is characterized by large tracts ofbalsam fir (Abies balsamea) and whitespruce (Picea glauca) intermingled withspecies such as paper birch (Betulapapyrifera) and aspen (Populus spp.). Thesoil is in this community type is highlyacidic and winters are usually prolongedwith extended snow cover. In the ecotonewhere the boreal forest meets the northern

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    hardwood forest, there is considerablespecies overlap although this region isgenerally characterized by spruce-fir-northern hardwood associations (Thompsonand Sorenson 2000). At higher elevations,above timberline, the “krummholz”naturalcommunity consists of gnarled and stuntedtrees that manage to survive the harsh winterelements on the exposed mountainsides(Marchand 1987).

    C. Suitability of the Northeast for GrayWolf RestorationLand availability.– Studies havedemonstrated that significant areas ofpotential gray wolf habitat exist throughoutthe proposed Northeast DPS region(Mladenoff and Sickley 1998, Wydeven etal. 1998, Harrison and Chapin 1997, Hosack1996.) Harrison and Chapin (1997)estimated that as much as 78,000 km2 couldpotentially be used as core and dispersalhabitat for gray wolves, with the majority ofthis habitat, almost 69,000 km2, containedwithin Maine and New York. In the 1992Eastern Timber Wolf Recovery Plan, theFWS proposed three areas for potential wolfrecovery in the Northeast: Adirondack Park,approximately 11,300 square miles innorthwestern Maine and northern NewHampshire, and a portion of eastern Maineconsisting of approximately 2,500 squaremiles. Potential wolf habitat in theNortheast is contiguous throughout Maine,northern New Hampshire and northernVermont, however, habitat in theAdirondack Mountain region is relativelyisolated (Mladenoff and Sickley 1998,Harrison and Chapin 1997.) While much ofthe potential habitat in the Northeast isunder either state or private ownership, mostof it is uninhabited and relativelyinaccessible.

  • Northeastern Gray Wolf DPS - 1 April 2003 40

    Road density.–Another benefit of this regionas favorable wolf habitat is that roads arerelatively sparse over most of the terrain. Ofthe potential suitable habitat areas identifiedin several studies on northeast wolfrestoration, 84 percent of the total area wasidentified as core habitat with road densitiesof less than 0.7 km of roads/km2 passable by2-wheel-drive vehicles and less than 4human residents/km2. Sixteen percent of thepotential habitat was identified as suitablewolf dispersal habitat with similar roaddensity criteria and less than 10 humanresidents/km2 (Harrison and Chapin 1997,Mladenoff and Sickley 1998).

    Prey base.–Wolves rely primarily onungulates for food and to a lesser degree,beaver and snowshoe hare (Paquet et al.1999). Ungulate populations in thenortheastern states fluctuate depending onwinter severity, harvest levels, disease, andpredation. In Maine, the white-tail deerpopulation has increased to 255,000wintering deer with deer abundance rangesfrom 2 to 5 deer/mile2 in the north to 15 to25 deer/mile2 in central and southern areas.Some locations, in which access torecreational deer hunters has been limited ordenied entirely, support deer populations of40 to 100 deer/mile2 (Maine Dept. of InlandFisheries and Wildlife). In the AdirondackMountains of New York, deer densitiesrange from 2-3/km2 and in the WhiteMountains of New Hampshire, from 3-4/km2 (Mladenoff and Sickley 1998). Moose populations in the Northeast werevery low in the late 1800's to early 1900'sdue to unregulated hunting. Since the1970's, moose populations have beenincreasing steadily in all the northeasternstates. From a low of 2,000 in the early1900's, Maine’s moose population iscurrently estimated at 29,000. Moose are

  • Northeastern Gray Wolf DPS - 1 April 2003 41

    beginning to re-colonize the Adirondackregion in New York State, and NewHampshire estimates its moose population at9,600 individuals. Overall, populationestimates indicate that moose densities are0.1-0.5/km2 in New England (Wydeven etal. 1998).

    Recent studies have documented that the proposed recovery areas in the Northeast couldsupport a minimum of 1,200 wolves and upto 1,800 wolves, based primarily on preydensity levels (Harrison and Chapin 1997,Mladenoff and Sickley 1998). With thecurrent prey biomass, the potential wolfrecovery areas could support wolf densitiesof 10 or fewer wolves/km2 (Wydeven et al.1998).

    Human Attitudes.– A review by Williams etal. (2002) of quantitative surveys conductedin the past thirty years on public attitudestowards wolves reported that 60 percentsupported wolf restoration. Another reviewby Buckley (2000) clearly shows a nationaltrend of growing support for restoration ofviable wolf populations. Specific to theNortheast, a December 2002 opinion pollconducted for the Henry P. KendallFoundation found that 63 percent ofnorthern New England residents believe it isimportant to reintroduce the wolf back to theregion for the balance of nature (BRS 2002). Kellert (1995) found that only 36 percent ofthe general public had a negative attitudetowards wolves, while 40 percent had apositive attitude and 20.9 percent neitherliked or disliked the species.. According to apoll conducted by Responsive Management(1996), 85 percent of New England residentsand 80 percent of New York residentssupported wolf reintroduction in AdirondackPark. A later survey showed that 60 percentof New York residents supported therestoration of wolves into the park while 34

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    percent neither approved or disapproved(Enck et al. 2000).

    Ecosystem Impacts.– The impacts of wolvesin ecosystems have never beencomprehensively studied, due to thedifficulty of establishing controls andreplication (Smith et al. 1999). It has beennoted, however, that removal of largepredators releases herbivores andmesopredators, causing overgrazing,vegetation recruitment failure, declines inground-nesting birds, and in general,ecosystem simplification, extinctions, anddecreased biodiversity. (Terbough et al.1999). Wolf effects on their herbivore preyspecies, as well as the resultant vegetationresponse, have been investigated. In three-level trophic systems, wolves areresponsible for maintaining vegetationlevels; for instance, on Isle Royale in LakeSuperior, predation by wolves releasesbalsam fir from browsing by moose(McLaren and Peterson 1994). Theinterruption of these trophic cascadeinteractions have been speculated as causingthe decline of aspen (Populus spp.) trees inYellowstone National Park following wolfextirpation in the 1920s. However, it is toosoon to determine if there has been avegetation recruitment response inYellowstone Park since wolf reintroduction(Ripple and Larsen 2000).

    Estimates based on population size indicated that wolf presence in Yellowstone Parkwould triple available carrion (Garton et al.1990), with potentially positive effects for awide range of scavenging species, includingfoxes, bears, weasels and raptors (Crabtree& Sheldon 1999). Wolves have killed anumber of coyotes in Yellowstone andaltered their behavior and home ranges(Crabtree and Sheldon 1999). Once theecosystem is released from extreme coyote

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    and ungulate pressure it has been speculatedto give a positive impact on numbers ofground squirrels, pocket gophers, hawks,owls, eagles, pronghorn, beaver, wetlands,moose, aspen, willows, and songbirds(Fischer 1998, Wilkinson 1997).

    Economic Impacts.– Eco-tourism is quicklymoving to the forefront of recreationalactivities. In areas where many animalsroam freely, none is more sought after thanthe elusive wolf. The longing to see thismagnificent animal has created an economicboom in areas such as Yellowstone NationalPark, Algonquin Provincial Park, NorthCarolina, and northern Minnesota.

    Since wolves returned to Yellowstone National Park in 1995, they have stimulatedsignificant economic activity. Visitors tothe park now rank the wolf as the numberone animal they come to see, therebycreating new demand for lodging, guidedwolf-watching tours and a variety of wolf-related merchandise. In Cooke City,adjacent to the northeast entrance, 22percent more tourists passed through thetown the summer after wolves were restoredthan the year before. In a survey of thecity’s business owners, 71 percent thoughtwolf recovery was responsible for increasedtourist traffic. It is estimated that the wolfreintroduction program has brought anadditional $23 million annually toYellowstone Park.

    Algonquin Provincial Park in Ontario, Canada, has had success in using wolves to attractvisitors to the park. Since 1963, one of themost popular events in Algonquin has beenpublic wolf howls. During any givensummer evening, an audience of about 2,400people will attend one of these events. A1997 study of North Carolina indicates that71 percent of those interviewed wereinterested in visiting the red wolf recovery

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    area. More significantly, respondents wouldbe less likely to visit the recovery area if redwolves were removed from the region. More recent research by Cornell Universitysuggests that red wolves have benefittednortheastern North Carolina’s economyanywhere between $40 million and $184million directly due to increased tourism. Innorthern Minnesota, 56 percent of touristsvisiting the town of Ely had visited theInternational Wolf Center. The Centerproduced a $3 million impact on the localeconomy in one year and either directly orindirectly provides the equivalent of 66 full-time jobs. Wolves attract a great deal ofattention and any area with a wolfpopulation will likely see an increase intourism revenues.

    D. Qualifications of the NortheasternGray Wolf Population as a DPSDiscreteness.–Northeastern populations ofeastern gray wolves should be designated asa DPS based on the criteria determined bythe FWS (61 Fed. Reg. 4722-25 Feb. 71996). The gray wolf population in thenortheastern United States is discreteaccording to the language of the FWS policybecause it is markedly separated from thenatural and recolonizing populations in theGreat Lakes due to distance separating thepopulations by 500 -1000 miles. In addition,the Service states that “the existinggeographic isolation of wolf populationsbetween these four areas [proposed DPSs]fully satisfies the Vertebrate PopulationPolicy’s criterion for discreteness of eachDPS,” (65 Fed. Reg. 43450 - 43496, July 13,2000. The distance separating the GreatLakes recovery areas and the westernmostpart of the proposed Eastern Gray Wolf DPSis 6-7 times the average dispersal distancefor a gray wolf (Gese and Mech 1991). While occasional wolves may disperse

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    across this distance, recolonization of thenortheastern states is very unlikely withinthe foreseeable future.

    Any gray wolves that appear in the northeastern United States may be lone dispersersfrom Canada. While several wolfpopulations exist in Canada within dispersaldistance of the Northeast, the numerousobstacles, both geographic andanthropogenic, these animals face as theyattempt to move into unoccupied habitat inthe U.S., are virtually insurmountable. Since no wolves have formed packs orestablished territories over the course of thepast few decades in the northeast region,there is little reason to believe that they willdo so in the future. Wolves in Canada aresubject to less protective wildlifemanagement plans and the ESA recognizespopulation discreteness based oninternational borders. Also, the FWS criteriafor “discreteness” states that a populationcan be considered discrete if it is isolatedfrom other populations of its taxon byinternational boundaries.

    Significance.–The Northeast represents asignificant portion of the wolf’s historicalhabitat and represents the easternmost extentof the wolf’s range in the lower 48 states.Wolves once roamed throughout NewEngland but were eliminated from theregion by the late 1800's. The regioncurrently encompasses over 26 million acresof suitable habitat and sufficient prey forwolves in a swath of forest that includesnorthern Maine, New Hampshire, Vermontand New York’s Adirondack Park. Studieshave shown that the Northeast could supportat least 1,200 wolves and perhaps as manyas 2000 (Mladenoff 1998, Harrison andChapin 1997). The current classification ofthis region under the Eastern Gray Wolf

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    DPS ignores the potential of this vast regionto wolf recovery and restoration.

    The FWS developed its 1992 Eastern Timber Wolf Recovery Plan to specifically recoverC. l. lycaon throughout its historic range,which included the Northeast. In its 2001Proposed Rule to Reclassify the Gray Wolfin the Conterminous U.S., the FWSacknowledges the complexities associatedwith gray wolf taxonomy but emphasizedthe agency’s commitment not to base graywolf recovery efforts on any particularportrayal of gray wolf subspecies, but toinstead identify geographic areas where wolfrecovery is feasible, regardless ofsubspecific affiliation of current or historicalgray wolves in the recovery areas. Theproposed rule also states that the FWSrecognizes the benefits to the species offocusing recovery efforts across a largeexpanse of the species’ range in order torecover and retain as much of the remaininggenetic variation as is feasible (65 Fed. Reg.43451-43452, July 13, 2000).

    The northeastern states - Maine, New Hampshire, New York, and Vermont - arecurrently considered part of the EasternGray Wolf DPS and are combined withexisting wolf populations in the Great Lakesstates. The FWS believes that wolves in thisDPS have met the recovery criteria set forthin the 1992 Eastern Timber Wolf RecoveryPlan, which called for a stable and growingwolf population in Minnesota and a secondpopulation outside of Minnesota and IsleRoyale. Wolf populations in Michigan andWisconsin are considered by the FWS to bea separate population from wolves inMinnesota. The Petitioners disagree sincethe movement and dispersal of wolvesamong these three states indicates that this iseffectively only one population of wolves inthe Great Lakes region. In order to meet therequirements of the Recovery Plan, a

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    separate population should be establishedoutside this region and within the DPS. Only the states in the northeastern portion ofthis DPS have enough habitat to sustainanother wolf population. Unfortunately, thelikelihood of wolves naturally recolonizingthe Northeast from the Great Lakes region ishighly unlikely given the extensiveanthropogenic and natural geographicbarriers separating the two regions (Harrisonand Chapin 1997, Wydeven et al. 1998). The only alternative is to develop a separaterecovery plan for the Northeast region.

    E. Northeastern Gray Wolf DPSQualifications for ESA ListingBecause this wolf population meets therequirements of discreteness andsignificance to its taxon, it should bedesignated as the Northeastern Gray WolfDistinct Population Segment and managedaccordingly. To increase the probability thatgray wolf populations will be successful inthis region, the Northeastern Gray WolfDPS should be afforded federal protectionunder the ESA and given endangered status. The threats to gray wolves are significant asthey disperse through the northeastern statesand as they recover within core regions ofMaine and New York. The ESArequirements for listing a DistinctPopulation Segment as threatened orendangered include manmade factors thataffect its continued existence and theinadequacy of existing regulatorymechanisms to sufficiently protect thespecies. If the final version of the proposedFish and Wildlife rule to reduce ESAprotections for wolves includes thenortheastern states, then the regulatorymechanisms will not be adequate tomaintain large enough wolf populations toprotect them from environmental stochastity

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    and the greatest threat to wolf populations:human-caused mortality.

    Conservation Status.-- If a population isdetermined to be discrete and significant(i.e., a Distinct Population Segment), theFWS must then determine whether it meetsthe definition of an endangered orthreatened species under the ESA. Thatdetermination must be based solely on anevaluation of the best available scientificinformation and the ESA’s five listingfactors. Gray wolves in the northeasternUnited States are currently listed asendangered. Before the FWS can legallydowngrade the gray wolf in this area, it mustdemonstrate that progress has been madetoward its recovery, and that threats to itscontinued existence have been reduced orremoved. While there have been sporadicobservations of individual wolves in thisregion over the last 20 years (most likelydomesticated animals or a rare transientfrom Canada), there is no evidence to-dateof reproducing pairs or pack formationsdespite vast areas of suitable habitat andseveral feasibility studies that indicate thepotential for successful restoration. Ananalysis of the ESA’s five listing factors andthe best available scientific evidence supportretaining an endangered classification forthe Northeastern Gray Wolf DPS.

    1. The present or threatened destruction,modification, or curtailment of it’shabitat or range.The Northeast represents an expanse ofsuitable habitat that provides an excellentopportunity to restore significant wolfnumbers and range. However, theavailability and utilization of that existingrange is jeopardized by a number of factors. As in most regions, increasing urbanizationand human populations are reducing the

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    amount of suitable wolf habitat. In the lastten years, population growth in the fournortheastern states included in the proposedDPS has increased by an average of sevenpercent. The areas within this regiondesignated as future recovery areas by theFWS are home to over two million people(U.S. Census Bureau 2001). Withpopulation growth expected to expand in asimilar manner in the next 20 years,development pressure on non-populatedareas will certainly increase. Expertsestimate that we are losing up to 70,000acres of wildlife habitat a year to conversionto human use. A large portion of the land inthe Northeast is privately owned andmanagement of any wildlands within theseholdings is subject to individual or corporatedecision-making, a situation not often infavor of endangered species protection. Inaddition, recreational development in andaround federal forest lands and state parksseverely diminishes the value of these landsfor wolf recovery. There are significantgeographical and legal barriers that preventwolf recolonization from any adjacent areas. The end result is that these available habitatsare not being utilized, which constitutes asignificant curtailment of range.

    2. Overutilization for commercial,recreational, scientific, or educationalpurposesCommercial take of wolves is currentlyillegal, though should wolves lose their ESAprotection it could become a significantfactor in preventing the reestablishment ofwolves within this region. The amount ofpoaching for commercial purposes isunknown but will be totally dependant uponthe regulatory status of the gray wolf (i.e.protected or not). For example, bountiesstill exist on the books in some states thatcould make harvesting wolves profitable.

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    Recreational take is also dependant upon theregulatory status of the wolf. Currently,hunting is restricted but without federalprotections some states have alreadysignified their intention to hunt wolves. Wewould expect a few research-relatedmortalities (capture and handling mortality)though it is unlikely that these will presentany significant impact on the population. All these issues indicate the need forcontinued federal protection under the ESAuntil wolves are clearly established, and theneed for implementing a recovery plan thatcan monitor and regulate the take from theabove factors and make managementadjustments accordingly.

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    3. Disease or predationMany diseases and parasites are foundamong the canids and some of these cancreate significant problems in wolf recovery,and require monitoring and appropriatetreatment to ensure that they do not spreadand impact the entire population. Whilesome individuals may die from diseases,disease is generally not considered asignificant problem for wolf recovery in thenortheast. Most wolves in North Americahave had regular exposure to many of thecanine diseases over the years and survive. Of course, any gray wolves that becomereestablished in the Northeastern Gray WolfDPS should be monitored for disease orparasite problems and treated as necessary. If wolves were reintroduced they would bevaccinated or treated for canine diseases andparasites.

    Natural predation from other wolves, bears and mortality from the defensive tactics ofprey species is relatively rare and would notbe expected to significantly affect gray wolfrecovery. However, the risk of human-caused predation can be substantial evenwhile under federal management andprotection (64 - 96 percent of all mortalityamong the reestablished wolves in theWestern US, 65 Fed. Reg. 43467). Wolfpopulations in the Northeast were extirpatedlargely due to human-caused mortality andthere continues to be a high level ofmalevolence towards the wolf fromrelatively small elements in the private andstate government sectors. Some statescurrently offer bounties for wolf kills andagricultural interests are advocating againstwolf recovery. Clearly the threat of humanpredation has not been reduced oreliminated in any substantive way, thereforewe must have the continued presence offederal management and ESA protectionuntil wolves have achieved some recovery

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    goal as defined by a Northeastern RecoveryPlan.

    4. The inadequacy of existing regulatorymechanismsThe proposed Northeastern Gray Wolf DPScontains a mix of primarily private andstate-owned land with very little land underfederal ownership. None of the appropriatestate agencies have yet addressed wolfmanagement issues or wolf protectionadequately. There is no recovery plan inplace for gray wolves, nor does FWS intendto develop one. Instead, the FWS proposesto downlist gray wolves in this area basedon the attainment of goals in the GreatLakes region identified in Recovery Plan forthe Eastern Timber Wolf (USFWS 1992). Gray wolf recovery in the Northeast is notaddressed in that plan even though theregion is geographically discrete. Any moveto downlist gray wolves in this area in theabsence of a scientifically credible recoveryplan for that area and demonstrable progresstoward the attainment of recovery goalsestablished under such a plan, isinappropriate or illegal. Without acoordinated recovery plan that involveslands controlled by the Forest Service, FWS,state agencies and private landowners, itappears highly unlikely that anymanagement plans for the region willadequately address wolf conservation. Allthis indicates the need for federalmanagement in this area with a specificrecovery plan and continued protection forthe wolves under the ESA.

    5. Other natural or manmade factorsaffecting its continued existenceEnvironmental Stochastity.– Natural wolfpopulations in Montana that were beingmonitored before the northern Rockiesreintroduction were in decline, and it was

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    found that extreme weather in the winter of1996-1997 killed an unusually largepercentage of the ungulate populations,leaving the gray wolf with scarce foodsupplies. The food shortage that began inthat winter resulted in a steep decline inwolf populations, partly due to increasedwolf depredation on livestock. Nearly 50percent of all confirmed wolf depredationsand lethal control actions in the period from1987 to 1999 were documented during thisone harsh winter. In the years since wolveswere reintroduced in Yellowstone andcentral Idaho their populations have beengrowing steadily, but there has not been awinter as severe as that of 1996-1997, so wedo not yet know the effects of a harsh winteron these populations. When the FWSdeveloped recovery plans for the gray wolfand set goals that would determine whenlevel of protections could be reduced, norealistic cycle of environmental stochastitywas considered. All successful wolfrecovery zones are in areas where winterconditions are harsh enough to kill naturalwolf prey, and where there are alternatefood sources of livestock available.

    The strong dependence of wolves on prey density also causes instability in wolfpopulations because ungulate populationsare sensitive to environmental stochastity(McRoberts et al. 1995). In the simplestpredator-prey models, predator populationscycle with prey populations, reacting to andcausing the prey population’s increases anddecreases. This model cannot describewolf-prey relationships because ungulatepopulations are much more dependent onclimate than on pressure from predators(McRoberts et al. 1995). However, asdescribed earlier, wolf populations aregreatly affected by shifts in their prey’spopulations. Therefore, when ungulatepopulations are reduced dramatically by a

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    harsh winter as they were in the winter of1996-1997 wolf populations can be expectedto decline in response due to starvation andby increased contact with humans whileforaging through agricultural land (Fed Reg.65). In the winter of 1996-1997 there was adecline in wolf numbers in the natura