Carnivore Reintroductions — Before, During, and After; Variables and Case Studies to Consider for the Northeastern U.S.

The following article is the first of a three-part series, an adaptation of a presentation the author gave at the Northeast Natural History Conference in Burlington, Vermont, Earth Day weekend, 2023. Note: “Before, During, and After” refers to both 1) phases during species reintroduction programs in the United States that are executed, underway, and currently considered, as well as 2) phases of a hypothetical puma (Puma concolor) reintroduction program needed for puma recovery in the Northeast. References follow the article.

Part 1

Wildlife authorities of the United States have a long history of reintroducing functionally extinct or extirpated species to their native ranges. State and federal agencies have implemented reintroduction, or population augmentation programs, across an arc of legislation that empowers them to do so, one that continues to evolve and undergo public debate. Carnivore reintroductions follow this arc, beginning prior to a mandate directing agencies to manage endangered species. How agencies manage extirpated species, i.e. what actions they take and why to recover species missing from human-impacted ecosystems, is an issue that evolves in the negative space of enacted endangered species legislation. What should wildlife authorities do about native species driven to extinction before laws were passed for extant species? And what should they do for species extinct on a state or regional level, but not national? Regional apex carnivore recovery, such as puma (Puma concolor, also known commonly as “cougar”, “mountain lion”, or “panther”) recovery in the Northeast is an example of this murky management void, which can also be seen, clearly by some, as a conservation opportunity, if not morally infused duty. The following case studies of apex, or keystone carnivore reintroductions in the U.S., can help inform a growing movement to restore central and eastern American landscapes by both supporting passively recovering populations, and actively reintroducing species such as the puma responsibly to these regions.

Black Bear (Ursus americanus) in Arkansas   Arkansas’ Game and Fish Commission ended black bear hunting in 1927, and by the late 1950s, only a small population persisted in and around the White River National Wildlife Refuge¹. Between about 1958-1968 — state records are spotty — around 250 young males and females captured in Minnesota, and Manitoba, Canada, were reintroduced^1,2. Bears were trapped in barrel traps and initially grouped together for transport resulting in debilitating injuries and death. Adjusting, project personnel then transported 6 bears per pick-up while keeping them individually contained¹. Bears were released into the interior highlands of Ouachita and Ozark National Forests with minimal public input^1,2. Hugh Hackler, the Commission’s director commanded: “There is to be no publicity”¹. Staff placed feeding troughs at each release site provisioned with commercial dog food¹. The reintroduced interior population established^1,2, and by 1973 the Commission began a program to relocate nuisance bears. The State reopened its black bear hunting season in 1980¹.

Red Wolf (Canis rufus) in North Carolina   The U.S. passed its first comprehensive law concerning rare and endangered species in 1966: the Endangered Species Preservation Act. Wildlife authorities then generated the first list of endangered wildlife after its enactment³. Red wolves, persisting in southeastern Texas and southwestern Louisiana at the time, were placed on this historic list, known then as C. niger^3,4. The U.S. Fish and Wildlife Service (USFWS) trapped over 400 canids from the relic population between 1973-1980 to create a captive breeding program. Congress passed today’s Endangered Species Act (ESA) in 1973, and authorities declared red wolves extinct in the wild in 1980. Port Defiance Zoo, in Washington, began a red wolf captive breeding program with 14 animals, which became an official, cooperative Species Survival Plan® in 1984. The USFWS designated a Non-Essential Experimental Population area (an ESA mechanism) in coastal North Carolina (NC) in 1986, and began reintroducing wolves to the Alligator National Wildlife Refuge in 1987.

This 50-year-old program features laudable achievements, much adaptive management, and wrenching challenges. Highlights include: An island facility was established in Florida in 1990 for a breeding pair to raise pups in a wilder but controlled environment before their translocation north. Red wolves formed packs, maintained territories, and bred successfully in the wild by the mid-1990s. Coyotes (C. latrans) continued their eastward range expansion entering NC in the 1980s⁵; the NC Wildlife Resources Commission (NCWRC) subsequently allowed coyote hunting in 1993 (C. Olfenbuttel, NCWRC, pers. comm.), at grave risk to wolves. Much conflict ensued, and conflict resolution work. Complicating matters, coyotes and red wolves can hybridize. The reintroduced NC population peaked at 120 animals over a decade ago. It remains endangered. Its population estimate was 15-17 in spring 2023. Human-caused mortality was the leading cause of population decline. A revised recovery plan is due this September (2023).

Mexican Gray Wolf (Canis lupus baileyi) in Arizona, New Mexico   USFWS listed Mexican gray wolves as endangered in 1976. Extinct in the U.S.⁶, it began a trapping effort in Mexico to again seed a captive breeding program. Ultimately, 7 wolves from 3 separate lineages, sourced both in Mexico and the U.S. (D. Parsons, USFWS, retired, pers. comm.), became founders of today’s population; the species went through a severe genetic bottleneck. The first Recovery Plan was published in 1982. It was not revised until 2017, but a successful legal challenge over inadequate attention to human-caused mortality resulted in a court-ordered remand. As a result, USFWS published a second revision in 2022. It too is currently challenged over inadequate genetic management⁷.

Reintroduction began in 1998 with 11 wolves: 3 bonded pairs with yearling offspring into Apache National Forest, Arizona. Wolves were released into acclimation pens as family groups for about 60 days; pens were at least 10 miles apart. New Mexico’s first release was of 1 family group in 2000 into the Gila Wilderness Area of Gila National Forest. Wolves were transported by horseback, and released into a nylon mesh pen they could chew out of (D. Parsons, pers. comm.). Like the red wolf, the “lobo” is still managed under an SSP program. Cross-fostering of captive pups by wild surrogate mothers is used as a genetic management technique. The “Interagency Field Team” reintroduces, monitors, and manages the U.S. population, which crossed 200 animals in 2022.

Canada Lynx (Lynx canadensis) in New York   The density and abundance of Canada lynx in northern New York (NY) prior to their extirpation is not well known. Rainer Brocke, PhD cited A.S. Bergstrom (1977) when reporting NY maintained a bounty on lynx until 1970⁸. Joint bobcat (Lynx rufus)-lynx studies began in 1976⁸. Brocke, a senior research associate at the State University of NY’s College of Environmental Science and Forestry, led the background research for lynx restoration/reintroduction — he used both terms — to the Adirondack Park. He built support for the program, and led the actual reintroduction that occurred over 3 field seasons from 1989-1992. His team carried, then delivered by helicopter, 83 wild lynx (49 females, 34 males) transported from the Canadian Yukon⁹ into a 670 square mile “Potential Lynx Restoration Area” of high-elevation conifer forest with, Brocke estimated, sufficient snowshoe hare density⁸. Lynx remained crated for 5 days provisioned with domestic rabbit carcasses⁹, then were released (the plan, as per the NY Times, was within a half mile of one another¹⁰). Post-release monitoring showed lynx failed to establish in the desired PLRA. Individuals roamed far beyond NY and to lower elevations; they suffered high rates of death by vehicular trauma, they were shot, and some even starved. No sign of breeding was ever confirmed⁹. Today, Canada lynx are considered extirpated by NY’s Department of Environmental Conservation.

Grizzly Bear (Ursus arctos horribilus) in Montana, Washington   Grizzly bears, or brown bears, are another species on the first federal endangered species list, published in 1967³. Categorized as threatened in the lower 48 states in 1975¹¹, their first recovery plan was also generated in 1982¹². In 1993, USFWS revised the recovery plan to include 6 recovery zones, core areas within 6 larger “ecosystems” where habitat and population criteria for recovery are measured. “Each recovery zone will include an area large enough and of sufficient habitat quality to support a recovered grizzly bear population.” The plan articulated recovery criteria for 4 zones¹² with the remaining 2 completed in 1996¹³ and 1997¹⁴. Brief discussion of 2 zones follows.

• In 1990, a pilot augmentation program began in Montana’s Cabinet (Mountain)-Yaak (River) Ecosystem. Four subadult females were translocated from southeast British Columbia, Canada to the Cabinet Mountains over 4 years. Genetic analysis of hair snags demonstrated successful breeding, so a formal, collaborative population augmentation program developed. From 2005-2021, 10 female and 8 male grizzlies were translocated within-state from the Flathead River region to the Cabinet Mountains. During 2020, 45 individual grizzly bears were detected in the Cabinet-Yaak Ecosystem using 5 methods¹⁵.
• In 2022, USFWS and the National Park Service (NPS) published a Notice of Intent to prepare an Environmental Impact Statement (EIS) for grizzly bear restoration to the North Cascades Ecosystem in Washington, where the species remains functionally extinct. Authorities plan to translocate grizzlies from British Columbia’s interior or the Northern Continental Divide Ecosystem. They aim to introduce 3-7 animals per year for 5-10 years with an initial population target of 25 to help restore biodiversity. The Draft EIS is due this summer (2023)¹⁶.

Florida Panther (Puma concolor coryi) in Florida   Florida (FL) panthers, the sole population of Puma concolor that persisted in the eastern U.S. through the 1900s, were also 1 of 14 mammals placed on our nation’s first endangered species list³. Declining due to inbreeding as management transitioned from sanctioned persecution to conservation, the World Wildlife Fund hired a private tracker and trapper in 1972¹⁷ to investigate the presence of a remnant population. Finding sign then live cats¹⁷, a FL Panther Recovery Team formed by 1976. The 1981 Recovery Plan was revised by 1987. Then, a feasibility study was conducted in north FL from 1993-1995 investigating the behavior of cats translocated from Texas, of cats wild-caught in Texas but temporarily captive, and captive-born cats from White Oak Conservation Center. Males were vasectomized, and all cats released for observation. Fifteen of 19 panthers established at least 1 home range, so the FL Game and Fresh Water Fish Commission decided re-establishment of additional panther populations was biologically feasible¹⁸. During the study, the Commission also convened with IUCN’s SSC Conservation Breeding Specialist Group to design a strategy for genetic rescue. Based on the estimate of 30-50 breeding adults, experts concluded introducing 8 young adult, non-pregnant females could meet the level of genetic introgression needed to counter genetic drift and inbreeding’s effects. They also stressed maintaining restored genetic health would necessitate further translocation to mimic an immigration rate of at least 1 new breeding panther per generation into the isolated population¹⁹.

Eight female pumas translocated from Texas were released as recommended into protected areas of southern FL in 1995. Five produced litters sired by FL males, and at least 20 first-generation kittens were born. Multiple health metrics indicate introgression worked, and the population grew to approximately 120-130 panthers by 2022²⁰. The FL panther is still isolated in need of more habitat and habitat corridors, and gene flow to grow. Vehicular trauma is the leading cause of death²⁰. The cause of feline leukomyelopathy, a disease that emerged in 2017 affecting the central nervous system of bobcats and panthers, is still unknown²¹. FL panthers remain endangered.

Gray Wolf (Canis lupus) in Idaho, Montana, Wyoming   By 1978 all extant gray wolf subspecies in the lower 48 states except for Minnesota’s population were federally protected. USFWS approved a recovery plan in 1980 for the Northern Rocky Mountain region, which was revised in 1987. Both discussed 3 recovery areas in Idaho, Montana, and the Yellowstone National Park (YNP) vicinity. Congress allocated funds in 1991 for an EIS directing USFWS to complete it by January 1994, and to choose management action that conformed with the law. USFWS chose “to establish an experimental population rule and reintroduce gray wolves to YNP and central Idaho, if two naturally occurring wolf packs cannot be located in either area”²². USFWS and NPS began reintroductions in 1995. Wolves came from western Canada from 1995-1996, then northwest Montana in 1997 where packs had begun to recover. YNP received 41 wolves, and Forest Service-administered lands in central Idaho received 35. Wolves underwent soft releases using guarded acclimation pens about 1 acre in size per family group. In 2011, Congress took unprecedented action by removing the gray wolf from the endangered species list in Idaho and Montana through a budget rider. USFWS delisted wolves in Wyoming in 2017. Gray wolf management and protection across the U.S. remains politicized and dynamic. USFWS announced in February 2023 that by 2 February 2024, it intends to submit a proposed rule concerning the listing status of gray wolves throughout the lower 48 states²³.

Canada Lynx in Colorado   Colorado Division of Wildlife (CDW) began planning for Canada lynx reintroduction in 1997. State biologists felt sufficient habitat existed in the San Juan portion of the southern Rocky Mountains to foster a self-sustaining population²⁴. Reintroduction occurred from 1999-2006. USFWS declared the Canada lynx threatened within the lower 48 states in 2000 to address what it considered inadequate protections in federal land management plans²⁵. Colorado at that point needed a permit to continue its reintroduction program, which it secured²⁶. Lynx were sourced from Alaska, and 3 provinces of Canada and the Yukon (E. Odell, CDW, pers. comm.). They were held in captivity for only days at first to minimize contact with humans; however, post-release mortality was high including incidences of starvation²⁷. Biologists began to extend the captivity period, ultimately revising protocol to retain lynx and feed them a high-quality diet for at least 3 weeks in captivity. Releases could then generally coincide with annual snow melt in late March or April. Initial post-release survival improved using the revised protocol²⁸. CDW also completed a formal Conservation Plan for USFWS in late 2002 — 5 years into the program — addressing multiple ways lynx could be taken, i.e. captured or killed, given their close resemblance to bobcats, a native game species²⁴. CDW documented its first lynx litters in spring 2003. Ultimately it translocated 218 cats²⁸. The reintroduced population eventually established and is monitored non-invasively during winters.

Gray Wolf in Colorado   In 2019, Colorado (CO) citizens submitted Ballot Initiative 107 to CO’s Secretary of State soliciting a democratic vote on gray wolf reintroduction west of the Continental Divide. This became Proposition 114, the Gray Wolf Reintroduction Initiative. Sam Brasch of CO’s Public Radio reported: “The Colorado ballot measure is the first time voters in any state will choose whether to bring back an endangered species”²⁹. Prop. 114 ordered the CO Parks and Wildlife Commission (CPWC) to create a plan to reintroduce and manage wolves, and to begin reintroduction by 31 Dec 2023. It also ordered the CPWC to create a livestock depredation compensation program funded by CO’s state legislature. Prop. 114 passed by a 1.8% margin in November 2020 and is now state statute. CPWC approved a final Wolf Restoration and Management Plan on 3 May 2023, and works now to meet statute deadlines with USFWS. CPWC plans to translocate 30-50 gray wolves over 3-5 years during the fall and winter from “Northern Rocky Mountain” states. Wolves will undergo a hard release; they will simply be freed from transport crates upon arrival at designated release sites within suitable habitat. Releases will occur at least 60 miles from adjoining state borders and sovereign tribal lands in the State’s southwest³⁰.

So what can the American electorate, particularly in the Northeast — from citizens to conservation practitioners to civil servants with decision-making power — and what can our youth learning ecological literacy, take away from these case studies? 1) The U.S. is highly experienced at reintroducing and rescuing carnivore populations, including large-bodied, far-ranging species; we have many programs to learn from. 2) Though states have differing mechanisms for voters to drive change, citizens, and state and federal wildlife agencies, can all act effectively to reintroduce native species. 3) The Northeast has experienced a lynx reintroduction attempt; reintroducing an apex carnivore to the Northeast would not be novel. 4) We have an ongoing recovery and management program in FL for its “panther”, a conspecific of pumas, that included, and may again, translocation of animals for genetic introgression. 5) The Northeast has exceptional biologists in its academic institutions, agencies, and non-profits, and the greater conservation community who can contribute expertise to a well-conceived, planned, and implemented puma reintroduction program. 6) Ecological consequences of a missing guild of species (“apex carnivores”) in northeastern ecosystems are well-researched and documented (more on this in Part 3). 7) We have mechanisms for transparency, communication, and accountability in the public realm; a serious consideration of puma reintroduction to the Northeast would occur in the public realm. 8) We have a growing and more vocal base of stakeholders who care for animals and the health and resiliency of our natural world.

Part 2 of this series transitions into planning for a hypothetical puma reintroduction program in the Northeast, and plausible ways operations could unfold during reintroduction and post-release monitoring.

Literature Cited^*
*Reference material found readily on authoritative websites is not listed

¹ Smith, K.G., J.D. Clark, and P.S. Gipson, 1990. History of Black Bears in Arkansas: Over-Exploitation, Near Elimination, and Successful Reintroduction. Proceedings of the Tenth Eastern Workshop on Black Bear Research and Management, 1990, pp 5-14. University of Arkansas Press, Fayetteville, AR.

² Smith, K.G. and J.D. Clark. 1994. Black Bears in Arkansas, Characteristics of a Successful Translocation. Journal of Mammalogy, 75(2): 309–320.

³ General Services Administration. 1967. Department of the Interior, Office of the Secretary, Native Fish and Wildlife, Endangered Species. Federal Register 32(48): 4,001. Washington, D.C.

⁴ McCarley, H. 1962. The Taxonomic Status of Wild Canis (Canidae) in the South Central United States. The Southwestern Naturalist, 7(3-4): 227-235.

⁵ Albers, G., C. Olfenbuttel, P.W. Sumner, and J. Thomas. 2017. North Carolina Wildlife Profiles: Coyote. North Carolina Wildlife Resources Commission. Raleigh, N.C.

⁶ U.S. Fish and Wildlife Service. 2022. Mexican Wolf Recovery Program, Progress Report #24 [2021]. Albuquerque, NM. 51pp.

⁷ Center for Biological Diversity and Defenders of Wildlife v. Haaland and USFWS. 2022. Case 4:22-cv-00303-JAS. https://earthjustice.org/wp-content/uploads/mexican_wolves_-_22-07-12_doc._1_complaint.pdf

⁸ Brocke, R.H. 1983. Restoration of the Lynx Lynx canadensis in Adirondack Park: A Problem Analysis and Recommendations. Final Report 1982: Federal Aid Projects E-1-3 and W-105-R, New York State Department of Environmental Conservation, Study XII, Job 5. NY DEC, Albany, NY. 77pp.

⁹ Brocke, R.H. 2009. Wildlife for a Wilderness, Restoring Large Predators in the Adirondacks in The Great Experiment in Conservation: Voices from the Adirondack Park, pp 169-190. Porter, W.F., Erickson, J.D., and Whaley, R.S., eds. Syracuse University Press, Syracuse, NY.

¹⁰ Halpern, S. 1989. Missing Lynx Return to the Wild. New York Times, New York City. NY.

¹¹ U.S. Fish and Wildlife Service. 2022. Grizzly Bear Recovery Program, 2021 Annual Report. Missoula, MT. 22pp.

¹² U.S. Fish and Wildlife Service. 1993. Grizzly Bear Recovery Plan. Missoula, MT. 181pp.

¹³ U.S. Fish and Wildlife Service. 1996. Grizzly Bear Recovery Plan, Supplement: Bitterroot Ecosystem Recovery Plan Chapter. Missoula, MT. 27pp.

¹⁴ U.S. Fish and Wildlife Service. 1997. Grizzly Bear Recovery Plan, Supplement: North Cascades Ecosystem Recovery Plan Chapter. Missoula, MT. 62pp.

¹⁵ U.S. Fish and Wildlife Service. 2022. Cabinet-Yaak Grizzly Bear Recovery Area 2021 Research and Monitoring Progress Report. Missoula, MT. 114pp.

¹⁶ National Archives and Records Administration. 2022. Department of the Interior, National Park Service, Fish and Wildlife Service. Federal Register 87(218): 68,190-68,192. Washington, D.C.

¹⁷ O’Connor, M.R. 2015. Resurrection Science: Conservation, De-Extinction, and the Precarious Future of Wild Things. Saint Martin’s Press. New York City, NY. 272pp.

¹⁸ Belden, R.C. and J.W. McCown. 1996. Florida Panther Reintroduction Feasibility Study, Final Report, Study Number 7507. Bureau of Wildlife Research, Division of Wildlife, FL Game and Fresh Water Fish Commission. Tallahassee, FL. 70pp.

¹⁹ Seal, U.S., ed. 1994. A Plan for Genetic Restoration and Management of the Florida Panther (Puma concolor coryi). Conservation Breeding Specialist Group, SSC, IUCN. Florida GFC Contract # 94027 Report. Apple Valley, MN. 23pp.

²⁰ Florida Fish and Wildlife Conservation Commission. 2022. Annual Report on the Research and Management of Florida Panthers: 2021-2022. Fish and Wildlife Research Institute & Division of Habitat and Species Conservation, Naples, FL. 86pp.

²¹ Florida Fish and Wildlife Conservation Commission. 2023. Updates, March 30, 2023. https://myfwc.com/wildlifehabitats/wildlife/panther/disorder/updates/. Accessed ~7 Apr 2023.

²² U.S. Fish and Wildlife Service. 1994. The Reintroduction of Gray Wolves to Yellowstone National Park and Central Idaho, Final Environmental Impact Statement. Helena, MT. 414pp.

²³ U.S. Fish and Wildlife Service. 2023. Statement on the Gray Wolf in the Lower-48 United States [Published 13 Feb 2023]. Washington, D.C. 1p.

²⁴ Colorado Division of Wildlife. 2002. Final Conservation Plan for Canada Lynx in Colorado. Fort Collins, CO. 10pp.

²⁵ National Archives and Records Administration. 2000. Department of the Interior, Fish and Wildlife Service. Federal Register 65(58):16,052-16,086. Washington, D.C.

²⁶ Colorado Division of Wildlife. 2000. Colorado Lynx Recovery Project 2000 Progress Report to the U.S. Fish and Wildlife Service. Glenwood, CO.16pp.

²⁷ Boster, S. 2019. 20 Years Later, Colorado Lynx Reintroduction Heralded a Success — but Threats Loom. Colorado Springs Gazette. Colorado Springs, CO.

²⁸ Colorado Division of Wildlife. 2010. Success of the Colorado Division of Wildlife’s Lynx Reintroduction Program. Gunnison, CO. 4pp.

²⁹ Brasch, S. 2020. It’s Official: Wolves Are Headed To Colorado’s 2020 Ballot. Colorado Public Radio. https://www.cpr.org/2020/01/06/its-official-wolves-are-headed-to-colorados-2020-ballot/. Accessed 20 Sep 2020.

³⁰ Colorado Parks and Wildlife. 2023. Colorado Wolf Restoration and Management Plan. Denver, CO. 261pp.