2013 NORTHEASTERN NATURALIST 20(2):289–308 History and Current Status of the Black Bear in Kentucky David E. Unger1,*, John J. Cox2, Hannah B. Harris3, Jeffery L. Larkin4, Ben Augustine2, Steven Dobey5, Joseph M. Guthrie2, John T. Hast2, Rebekah Jensen2, Sean Murphy2, Jason Plaxico5, and David S. Maehr2,6 Abstract - Once abundant in Kentucky, Ursus americanus (American Black Bear) were extirpated from the state by the late 19th century because of overharvest and habitat alteration. Regenerating deciduous forests, increased human tolerance, and source population growth and expansion in neighboring states have facilitated Black Bear recolonization in parts of southeastern Kentucky since the 1980s. As of 2012, <500 Black Bears were estimated to occur in Kentucky, with most individuals found in two successfully reproducing, geographically separate, and genetically distinct core populations in the southeastern part of the state. Our research suggests that population growth and expansion of Black Bears within Kentucky is occurring and abundant suitable habitat exists to support further increases in range and numbers. Potential impediments to further population growth and recolonization include roads, overexploitation primarily from illegal harvest, and habitat loss and fragmentation. The recolonization of Kentucky by the Black Bear represents an important case study of population growth and expansion of large mammals in the eastern US that has widespread ecological and economic implications. Introduction Large body size, low density, habitat specialization, slow growth rates, carnivory, and herd or pack formation are life-history attributes of large mammals that make them vulnerable to persecution and overexploitation by sympatric humans who often feel threatened by their existence (Forbes and Boyd 1996, Kellert et al. 1996, Mladenoff et al. 1995, Purvis et al. 2000, Treves and Karanth 2003, Woodroofe and Ginsburg 1998). These characteristics, coupled with an increasingly human populated and modified global landscape, also make it difficult for large mammals to naturally recolonize areas within their historic range. Nonetheless, several important examples of large-mammal recolonization and successful reestablishment have occurred during the past half century that has had important ecological and sociological impacts. For example, Canis lupus L. (Gray Wolf) recolonized 8 areas of North America and Europe, in two cases negotiating agricultural landscapes to do so (Wydevan et al. 1998). In Europe, Ursus arctos L. (Brown Bear) has recolonized the northern two-thirds of Sweden and northeastern Norway after being reduced to approximately 130 animals in the 1930s (Swenson et al. 1998). 1502 E. Lamar Alexander Parkway, Division of Natural Sciences, Maryville College, Maryville, TN 37804. 2Department of Forestry, University of Kentucky, Lexington, KY 40546-0073. 3Department of Animal and Food Sciences, University of Kentucky, Lexington, KY 40546-0073. 4Department of Biology, Indiana University of Pennsylvania, Indiana, PA 15765. 5Kentucky Department of Fish and Wildlife Resources, #1 Sportsman’s Lane, Frankfort, KY 40601. 6Deceased. *Corresponding author - dave.unger@ maryvillecollege.edu. 290 Northeastern Naturalist Vol. 20, No. 2 Bears in North America have been extirpated at different spatial scales due to heavy overexploitation, habitat loss and alteration, and human-bear conflicts (Hellgren and Maehr 1992, Servheen 1990). In contrast to populations in Scandinavia, the Brown Bear has regained little of its former range in the US during the past half century because of direct conflicts with humans. Ursus americanus Pallus (American Black Bear) has experienced a 41% overall loss of its historical range in North America since European settlement (Laliberte and Ripple 2004) and as much as a 90% loss of habitat in the United States (Pelton 1996) including the southeast (Maehr and Brady 1984). Unlike most of North America’s other large carnivores, the Black Bear currently appears to be thriving in large portions of its existing range (Hristienko and McDonald 2007, Servheen 1990, Servheen et al. 1999). For example, the Black Bear recently recolonized the Trans Pecos Region of Texas (Onorato et al. 2004) and, following reintroduction elsewhere, parts of Arkansas (Smith and Clark 1994), Oklahoma (Bales et al. 2005), and Mississippi (Ursus americanus and Ursus americanus luteolus; MDWFP 2012). In some areas, wildlife managers assert that Black Bear numbers are sufficient to sustain a regulated harvest (Garshelis and Hristienko 2006), while elsewhere, smaller populations face threats of extinction because of habitat loss, poaching, and vehicle collisions (Maehr et al. 2003). In the past two decades, the Black Bear has become reestablished in the southeastern portion of Kentucky via colonization from eastern border states and, to a lesser degree, from population growth after a limited translocation that occurred on the state border with Tennessee during the late 1990s. The immigration and successful reestablishment of the Black Bear into Kentucky is an important case study of large-mammal population growth and expansion in the eastern US during the past century. The return of the Black Bear to Kentucky has restored an important ecological component to an increasingly denatured, yet still highly diverse ecosystem. Additionally, the Black Bear has already had socioeconomic impacts on local communities as a wildlife icon used to promote local ecotourism areas that have experienced decades of economic hardship (Harris 2011). As an important case study of large-mammal recolonization, we provide a brief history of the Black Bear in Kentucky, its ecological and economic importance, and discuss factors that may challenge further colonization and long-term population persistence. History of the Black Bear in Kentucky Post-Pleistocene through 1900 The Black Bear and its ancestors have existed in North America since the Pliocene and precede all other extant species of bear on the continent (Kurten and Anderson 1980). The evolutionary success of the Black Bear can be evaluated both in terms of persistence and distribution; it has survived where other Bears have been extirpated, first during the ice ages (Wooding and Ward 1997) and more recently in the face of human encroachment (Laliberte and Ripple 2004). The pre-European settlement distribution of the Black Bear ranged from Mexico to Alaska and from the Atlantic Ocean to the Pacific Ocean (Kays and Wilson 2002). The Black Bear is a habitat generalist that can adapt to a wide variety 2013 D.E. Unger, et al. 291 of environmental conditions and inhabit a diversity of habitats from swamps to desert scrub (Onorato et al. 2004, Whittaker and Hamilton 1998). Regardless of the vegetation type, the Black Bear requires escape cover (Pelton 2000), which has led to it occasionally being labeled as a forest obligate (Maehr et al. 2003) in particular ecosystems. Immediately prior to European settlement, much of the eastern US, including what is now Kentucky, was forested (Wharton and Barbour 1973) and supported a post-Pleistocene large-mammal fauna that included Bison bison L. (American Bison), Cervus elaphus L. (Elk), Canis sp. (wolf), Odocoileus virginianus Zimmerman (White-tailed Deer), Puma concolor L. (Mountain Lion), and Black Bear (Barbour and Davis 1974, Whittaker and Hamilton 1998). Kentucky’s diverse, mixed-mesophytic forests in the eastern one-third of the state are highly productive, and may have harbored one of the highest densities of Black Bears in North America before European settlement. These forests contained vast numbers of mast-producing canopy trees (Braun 1950) such as the now functionally extirpated Castanea dentata (Marsh.) Borkh. (American Chestnut) as well as Quercus spp. (oaks), Diospyros virginiana L. (Persimmon), Nyssa sylvatica Marsh. (Blackgum), Sassafras albidum (Nutt.) Nees (Sassafras), Fagus grandifolia Ehrh. (American Beech), Carya spp. (hickories), and Juglans spp. (walnut). Understory fruiting woody plants such as Amelanchier arborea (Michx. F.) Fernald (Serviceberry), Vaccinium constablaei L. (Highbush Blueberry), V. pallidum Ait. (Lowbush Blueberry), Rubus spp. (blackberry), and Asimina triloba (L.) Dunal (Paw-Paw), were also abundant and undoubtedly offered important food items to Black Bears and other wildlife species. Likewise, abundant rock shelters and hollow trees interspersed among Rhododendron spp. (rhododendron) and Kalmia latifolia L. (Mountain Laurel) likely provided important cover for escape, thermoregulation, and denning. Although the Black Bear was likely most abundant in the Southern Appalachian regions of Kentucky before European settlement, their range was not confined to those mountainous habitats. Archaeological evidence indicates the Black Bear was also widespread in the savanna-woodland-cane landscape mosaic that characterized the Bluegrass Region where the same plant species or ecological surrogates provided abundant forage (Hall 1981, Wharton and Barbour 1991). Although there are few accounts by early Europeans describing western Kentucky flora and fauna, local place names suggest Black Bears may have also been present in the central hardwood forests, barrens, and swamps of this region. The use of fire by Native Americans beginning around 7500 years ago had major impacts on forest ecosystems and large mammal abundance and distribution in the eastern US by creating a more heterogeneous landscape favorable to a number of game species including Black Bear (Delcourt et al. 1998, Lyons et al. 2003, Russell 1983, Williams 1989). Many shade-intolerant species that produce soft mast fruit often thrive in fire-maintained forests. Such early successional plants are valuable to wildlife, providing food to both birds and mammals (Martin et al. 1951). Fire can offer a competitive advantage to hard-mast producing trees, such as oaks, which provide important seasonal food to Bears and other wildlife (Brose et al. 2001, Wright and Bailey 1982). 292 Northeastern Naturalist Vol. 20, No. 2 Kentucky was considered a rich hunting ground and was shared by several neighboring Native American tribes (Arnow 1960, Raitz et al. 1998). In fact, the Cumberland Plateau ecoregion of eastern Kentucky and neighboring states in Central Appalachia was regarded as the “Middle Hunting Ground” (Arnow 1960:58) due to its abundant and diverse game. Swanton (1979) speculated that the Black Bear was perhaps the second most important game species for Native Americans. Historical evidence suggests that Black Bears were abundant in Kentucky, as their claws, skulls, and teeth were used by Native Americans and have been found at numerous archeological sites in the state (Funkhouser and Webb 1928, MacCord 1953, Mills 1904). Hallowell (1926) reported the use of dogs by Native Americans to locate and kill hibernating bears in winter, and Dr. Thomas Walker noted the killing of 53 Black Bears during their 1750 journey into Kentucky (Walker 1750). Other early European settlers describe the prevalence of Black Bear and the use of a substance called “butter” that scholars believe was probably fat collected from harvested Black Bears (Davis 2000). The hundreds of place names dotting the Kentucky landscape that reference bear, such as Bruin, Bear Creek, and Bear Branch, also suggest a statewide range and abundance of the species (Cox et al. 2002). Despite early subsistence use by native peoples, the first large-scale impact of hunting on Black Bear likely began in the more southern US with the increased fur trade following Spanish settlement in the 16th century. Trade in fur among tribes had existed for some time, but growing European demand in the 17th through the early 19th century quickly outpaced supply (Cronon 1983). European settlers may initially have lacked the experience to efficiently harvest furbearers, but high demand produced an incentive for native hunters to increase their harvest significantly above subsistence levels (Arnow 1960, Cronon 1983). Although the native peoples of Appalachia were slower to become involved in trade with the Spanish than groups living farther south, historical records indicate they were actively involved in the trade of animal hides by 1650 (Davis 2000). Around a century later, native peoples radically changed the nature of Black Bear hunting by adopting firearms and European hunting dogs (Arnow 1960, Davi s 2000). Bear hides are not mentioned specifically in most accounts of early European traders, as deer, Bison, and Castor canadensis Kuhl (Beaver) appear to have been the dominant species traded (Davis 2000). Nonetheless, the Black Bear was one of the most sought after game animals because of the many products that could be derived from its carcass (Aron 1990). Early accounts by pioneers detail the use of Black Bear hides as bedding, coats, rugs, food preparation bowls, and moccasin linings (Arnow 1960, Belue 1996, Eslinger 2004). Black Bear hides were also used to wrap and protect deerskins in bundles of 50 bound with buffalo hide ties for shipment (Belue 1996). Black Bear bladders were used to carry oil, or were themselves used as oil cloth to wrap letters and packages (Arnow 1960). Black Bear fat was used for cooking oil, shortening, candles (Belue 1996, Eslinger 2004), and medicinally for people (Belue 2003) and Equus caballus L. (Horse; Arnow 1960). Black Bear meat was thought to increase a man’s sexual potency (Arnow 1960). Bear bacon was highly sought after for its flavor and because it took less salt (a precious item to pioneers) to cure than its domestic counterparts 2013 D.E. Unger, et al. 293 (Belue 2003), and the heart and kidneys were considered delicacies. Even bear cubs were consumed (Belue 2003). From 1805–1807 over 8000 Black Bear hides from the vicinity of the Big Sandy River in Lawrence County, KY were sold and exported to Europe to make grenadiers’ hats for the British armies (Collins 1882). Consequently, the lucrative supply of Black Bear products coming from Kentucky led to its Colonial period nickname as the “Bear State”. This would soon change as the more commonly traded species became regionally scarce or extirpated and bears became even more sought after. Dr. Walker’s journey into eastern Kentucky and the subsequent publication of his enumerated animal observations would spur “one of the greatest human migrations in history” (Jillison 1931:11), and have consequences that would determine the fate of Black Bears in Kentucky and lands westward for the next two centuries. Between 1775 and 1810, nearly 300,000 settlers entered southeastern Kentucky through the Cumberland Gap (Burns 2000); the state was considered settled by 1830. Among the first were the Kentucky “long hunters”, including Daniel Boone, who arrived soon after Dr. Walker’s journey and brought market hunting with them (Belue 2003). The meat and fur of Kentucky’s abundant wildlife were a boon for areas east of the Appalachians, which had already lost many large mammal species (including Black Bear) as early as 1710 (Dunaway 1996). Bison were extirpated from Virginia by the 1730s (Belue 1996), and even deer were scarce by the 1750s (Dunaway 1996). When long hunters arrived in Kentucky, “Buffalo, Elk, and bear were said to be rolling fat, and weary for the rifle shot” (quoted in Aron 1990:55). Market hunting, which peaked between 1760 and 1780, greatly reduced the region’s wildlife. By 1820, Bison were extirpated from Kentucky (Belue 1996). In addition, European markets for pelts pushed prices to levels where a backcountry hunter could make several hundred English pounds per year. Hunters and trappers were subject to fluctuating market demands for furs and regularly had pelts stolen or ruined by fellow trappers, Native Americans, and predators; all these factors encouraged more intensive hunting (Aron 1990, Belue 1996). Additionally, Native Americans increasingly hunted wildlife with firearms acquired from colonists as many tribes sought to become a part of the new European political and economic order (Dunaway 1994, Hudson 1981). Furs allowed Native Americans to trade for European and colonial goods, acquire additional guns for protection and warfare, and served as bargaining chips during negotiations (Hudson 1981). By May of 1774, the harvest of wildlife by settlers in Kentucky was so prodigious that delegates from four settlements attempted to pass laws preserving game (Cotterill 1917). By 1781, the effects of overhunting were apparent, as hunters were forced to travel up to 40 km to find game (Arnow 1960). Like many other areas in the US during the 18th and 19th centuries, agriculture caused the rapid clearing of forests, savannas, grasslands, and draining of wetlands. This land development in turn led to the loss of wildlife, particularly in the arable western two-thirds of the state. Viable Black Bear populations that remained in Kentucky during the latter half of the 19th century were likely found in the more rugged southeastern mountains. Although less human populated, ecosystems in these areas were heavily impacted by smaller farms. Here, 294 Northeastern Naturalist Vol. 20, No. 2 Sus scrofa domesticus L. (Domestic Pig) and other livestock were often allowed to free-range forage in forests where they consumed massive amounts of acorns, chestnuts, and beechnuts, putting them in direct competition with Black Bears and other wildlife (Aron 1990, Cox 2005, Yarnell 1998), as well as slowing recruitment of important mast trees. In this region, subsistence hunting supplemented agriculture, diversified the diet, and became a core part of eastern Kentucky and central Appalachian culture that lasted well into the 20th century. Black Bear meat was considered a staple food in eastern Kentucky well into the 1800s (Arnow 1960), and as late as 1871, Black Bears were still being hunted along Pine and Black Mountain (Phillips 1991). Large-scale habitat loss, degradation, and fragmentation of old-growth forests from commercial logging that occurred between 1880 and 1920, and continued overharvest, likely led to the extirpation of the few remaining Black Bears in Kentucky around the beginning of the 20th century. Genetic evidence and early accounts suggest that the Black Bear’s range was largely reduced in the eastern US to remnant strongholds in the most rugged and inaccessible mountains of central Appalachia (LaFollette 1974, VDGIF 2002, Wathen 1983). Stabilization, population growth, and recolonization (1900–2010) During the late 19th and early 20th centuries, the overexploitation of natural resources and the potential consequences of inaction had become increasingly apparent to the US public. Consequently, the conservation movement in the US had gained significant political influence and social acceptance during this period. From 1900–1920, governments at multiple levels established a resource conservation model by enacting a number of federal and state laws that outlawed market hunting, established harvest limits of wildlife and timber, and created protected areas to conserve these resources. Most state wildlife agencies, including the Kentucky Department of Fish and Wildlife Resources (KDFWR), were created during this era and began enforcement of laws designed to primarily protect game species; however, by that time, the Black Bear had already been extirpated from the state. During the early 20th century, the Black Bear was present in limited numbers in Kentucky’s neighboring states of Virginia, West Virginia, and Tennessee, as well as nearby North Carolina (Maehr and Brady 1984). The establishment of public protected areas and reforestation of much of their early 20th-century range likely saved the Black Bear from extirpation in Appalachia (Clark and Pelton 1999, Frary et al. 2011). At the turn of the 20th century, rampant destruction of forests began to produce concerns by the newly formed American Forestry Association (AFA) and influential socialites who viewed the mountains as a place for rest and recreation. As early as 1901, recommendations for the establishment of an eastern forest reserve in the Appalachian Mountains were introduced to the US Congress (Eller 1982). During this same period, human emigration to urban areas caused the abandonment of many farms throughout the region, allowing reforestation to occur, and creating opportunities for large-scale land acquisition by natural resource agencies (Clark and Pelton 1999, VDGIF 2002, Yarnell 1998). The Weeks Act of 1911, and later the Clark-McNary Act of 1924, resulted 2013 D.E. Unger, et al. 295 in the establishment of a number of US National Forests (Cherokee, George Washington, Jefferson, and Monongahela) in central Appalachia that, with Great Smoky Mountains (GSMNP) and Shenandoah National Parks, would prove critical for harboring remnant Black Bear populations that would later serve as source populations for recolonization. Tourism in the southern Appalachians increased rapidly from 1945 through 1960, resulting in increased political power of environmental interest groups (Yarnell 1998) and increased interest in protecting the Black Bear beyond park boundaries. The desire for Black Bear population recovery for both recreational and ecological reasons led to the creation of Black Bear sanctuaries in Tennessee (four in 1973 and two in 1997), North Carolina (28 beginning in 1971), and West Virginia (1967 and 1971) to serve as source populations for growth and colonization. Yet as forests were beginning to regenerate in the mid-20th century, the most consistent mast-producing tree throughout the Southern Appalachian Mountains, American Chestnut, began to succumb to Cryphonectria parasitica (Murrill) Barr (Chestnut Blight Fungus). The blight was first noted in New York in 1904, and by the 1950s had caused a near complete functional loss of the American Chestnut in Appalachia that continues today (Diamond et al. 2000, Leopold et al. 1998, Schibig et al. 2005). Consequently, less consistent mast-producing species such as oaks, hickories, and the American Beech gained dominance in regenerating forests throughout the Appalachian region. Decline of the American Chestnut may have affected the overall carrying capacity for a number of species inhabiting eastern forests, altered Black Bear ecology in many regions (Clark and Pelton 1999, Pelton 1996), and likely slowed recolonization in several areas of eastern North America. Where extant in several eastern states, the Black Bear was often declared a game animal, providing it some protection through the establishment of bag limits and hunting seasons (Bolgiano 1998, Clark and Pelton 1999), although these were relatively unrestrictive until the early 1970s. After hunting seasons were shifted to later in the season to allow females to den and as forests continued to mature, Black Bear populations began to slowly increase and expand in Tennessee, West Virginia, and Virginia (Clark and Pelton 1999, Cowan 1972, Spiker and Bittner 2004, VDGIF 2002, Wolgast et al. 2005); however, these populations only began to exponentially grow after 1980 (Table 1). Yet while Black Bears recolonized other portions of Appalachia, observations in Kentucky remained rare until the late 1980s because major source populations from neighboring states remained geographically distant until the latter part of the 20th century. Although unconfirmed sightings of Bears in scattered portions of eastern Kentucky were reported from 1900–1980 (Barbour and Davis 1974, Funkhouser 1925, Harris 2011), an established breeding population was never confirmed during this period. Collectively, most observations of Black Bears until the mid-1980s appear to have been subadult males that had dispersed from neighboring states with existing breeding populations (KDFWR, Frankfort, KY, unpubl. data). This pattern typifies the colonizing front of many carnivores, and is driven by the tendency of young males to potentially disperse hundreds of kilometers from natal areas, while young females are highly 296 Northeastern Naturalist Vol. 20, No. 2 philopatric, typically establishing territories within and adjacent to the matriarchal home range (Rogers 1987). Beginning in the early 1980s, confirmed Black Bear sightings were reported by KDFWR personnel in counties bordering West Virginia and Virginia (Maehr 1984; Fig. 1). The first capture of a Black Bear in recent history occurred in southern Pike County near Kentucky’s border with Virginia in 1987. Black Bear sightings increased to 69 in 1989 and 161 in 1992. Bear observations continued to increase from the mid-1990s, with KDFWR personnel handling 26 Bears (25 M, 1 F) between 1987 and 2000 (Plaxico and Bonney 2001). In 2001, cubs were observed in Harlan County, confirming the first resident breeding Black Bear in eastern Kentucky in over a century. In addition to natural recolonization, the Black Bear was reintroduced by the National Park Service (NPS) in 1996 and 1997 to the Big South Fork National River and Recreation Area (BSF) in Tennessee within a few miles of McCreary County, KY (Eastridge 2000, Eastridge and Clark 2001; Fig. 1). Twenty-seven Black Bears (14 adult females, 13 cubs) were translocated from GSMNP to BSF. Three additional cubs were born in winter 1997 from one of the translocated females, and five more cubs were born from relocated females in 1999 (Eastridge 2000). From 1987–2010, Black Bears were observed in 54 Kentucky counties, although most of these individual reports were indicative of transient males far from the resident breeding populations (Fig. 1). The apparent population increase and documented reproduction of Black Bears prompted initiation of formal research studies through a University of Kentucky (UK)-KDFWR partnership beginning in 2002. From 2000–2010, KDFWR personnel handled 83 (64 M, 7 F, and 12 unknown gender) subadult or older Black Bears, or nearly three times as many from the previous 14 years of capture effort; all but three of those were located within a two-county distance from West Virginia, Virginia, or Tennessee. From 2003–2010, UK researchers captured 138 (93 M, 45 F; 122 in southeast Kentucky, 16 in BSF) subadult or older Black Bears. These collective capture efforts and radio-telemetry studies identified two core areas of reproduction: Table 1. Harvest trends in states that border Kentucky’s two core Black Bear populations. State Date Number Harvested Tennessee1 1951 29 1981 21 2011 581 West Virginia2 1950 32 1980 47 2011 2007 Virginia3 1950 <200 1980 ≈250 2011–2012 1997 1TWRA (2012). 2West Virginia Division of Natural Resources, South Charleston, WV (unpubl. data). 3VDGIF (2012). 2013 D.E. Unger, et al. 297 a southeastern population that primarily inhabited the Pine, Black, and Cumberland Mountain region of Bell, Harlan, and Letcher counties, and a separate breeding population located in the McCreary County area within the BSF adjacent to Tennessee (Frary et al. 2011, Hast 2010). As with many other large, cryptic mammals, estimating the Black Bear population size can be difficult given its low population density and forest habitat. The distribution and abundance of Black Bears in North America and Kentucky have often been estimated based on surveys sent to state agency personnel, informed opinion, and conjecture (Maehr 1984, Pelton and Van Manen 1994, Rounds 1987). Maehr (1984) indicated only occasional Black Bear sightings in Kentucky, while Rounds (1987) suggested that only a small portion of the easternmost edge of Kentucky adjacent to West Virginia and Virginia was occupied by Black Bears. Pelton and Van Manen (1994) indicated a much wider Black Bear distribution in Kentucky, highlighting much of the Cumberland Plateau as occupied range. Littleton (1994) published the first Kentucky distribution map based on field data with confirmed Black Bear sightings in 25 counties (Fig. 1). Williamson (2002) reported that <25% of Kentucky Black Bear habitat was occupied, but offered no distribution map. Collectively, several studies estimated that from 1970–1996, Black Bears likely numbered 100 or fewer individuals and were primarily found in eastern Kentucky (Pelton 2001, Pelton and Van Manen 1994, Pelton et al. 1999, Williamson 2002). Figure 1. Indices of Black Bear distribution in Kentucky, 1987–2010. No visual sightings were reported from 1995–1997. 298 Northeastern Naturalist Vol. 20, No. 2 Non-invasive approaches (Mowat and Paetkau 2002) have become a preferred empirically based approach to estimate population sizes than less standardized ones (e.g., roadkill surveys). Frary et al. (2011) performed the first systematic survey of Black Bear abundance in an 11-county area of southeastern Kentucky and used density estimates to calculate a Black Bear population size of 130 individuals (95% C.I. = 92–165) in 2007. The majority of identified Black Bear hair samples from that study were found in Bell, Harlan, and Letcher counties immediately bordering Virginia and Tennessee in far southeastern Kentucky (Fig. 1). A more recent genetic analysis from a hair snare survey of McCreary County estimated approximately 40 (95% C.I. = 30–113) Black Bears within the one-county study area in 2010 (Murphy 2011). Based on those empirical mark-recapture estimators and associated biological data, it is estimated that the statewide population at the end of 2011 numbered 250–500 individuals primarily located in eastern Kentucky (KDFWR, Frankfort, KY, unpubl. data). Using DNA microsatellite loci data, Hast (2010) determined that Black Bears in Kentucky can effectively be divided into 2 primary populations: the Cumberland Plateau population, sourced by recolonizing individuals with genotypes unique to or an admixture of those from West Virginia and Virginia (but not Tennessee), and the separate BSF population, sourced almost exclusively from translocated individuals and their descendants from a limited reintroduction effort in 1996 and 1997. Collectively, radiotelemetry data and microsatellite DNA analyses indicate that connectivity between the Cumberland Plateau and BSF populations appears to be very limited thus far. Hast (2010) and Frary et al. (2011) found that genetic diversity within the Kentucky Black Bear population is similar to others in North America. Current and Future Management Issues for the Kentucky Black Bear Given the relatively small size of Kentucky’s two distinct Black Bear populations, there are two major factors that could slow or impede further colonization, and ultimately lead to population declines: 1) habitat fragmentation and loss, primarily from roads, urbanization, agriculture, and large-scale surface mining, and 2) overexploitation from legal harvest (Clark and Eastridge 2010) and illegal harvest driven by the continuing black market demand for Black Bear parts in eastern Asia (Mills and Serveen 1991) and a growing market in the US (Mann 2012). The Black Bear typically requires relatively large (>5 km2) tracts of forest habitat (Smith and Pelton 1990). A recent habitat analysis revealed over 8500 km2 of well-connected, suitable forestland in eastern Kentucky (Unger 2007). From a landscape perspective, it would therefore appear that Kentucky has sufficient resources to facilitate continued colonization of much of the state where forest size, composition, and connectivity is adequate. Yet despite the capacity of an individual Black Bear to colonize an area far from its natal range, Hast (2010) and Frary et al. (2011) found that females primarily occupied areas very near the state border or at BSF within ≈10 km of the original release sites. This characteristic philopatric behavior indicates that the reproductive population core lags far behind a colonization front comprised almost entirely of juvenile males. 2013 D.E. Unger, et al. 299 Although primary road density is relatively low in southeastern Kentucky, the proposed construction of Interstate 66 in this area has the potential to be a major movement barrier. Jensen (2009) found that roads affected Black Bear movement and space use in Kentucky; they avoided roads at the local and landscape level, and patterns of road avoidance and crossing varied by gender (males displayed stronger road avoidance than females, while females had greater selectivity in road crossing than males). Yet despite their apparent road wariness, at least 40 Black Bears have died in vehicle collisions since the late 1980s in Kentucky. Jensen (2009) suggested a potential barrier effect by roads could slow the continued Black Bear recolonization into Kentucky by inhibiting dispersal of females westward into the Cumberland Plateau. Given the relative abundance of forest cover and recent regional declines in coal production and timber prices that have negatively impacted the level of these extractive activities in Kentucky, overexploitation may far outweigh habitat-related factors concerning near-term population viability and growth. Humans are responsible for nearly all mortality in Black Bears over two years of age (Rogers 1976). Although legal hunting is the major cause of Black Bear mortality elsewhere, it is not yet a significant factor in Kentucky, which held its first hunt in 2009. The first three limited seasons from 2009–2011 resulted in the harvest of only six bears; however, at least 24 Bears have been poached in the past two decades. Other types of human-caused deaths are still problematic in Kentucky, if population growth and long-term viability are desired. Bear populations are often not able to sustain heavy adult mortality (Cowan 1972, Freedman et al. 2003, Miller 1990). Thus, limiting legal harvest and curtailing poaching while the population is relatively small are important actions to sustain population growth. Protection of adult female Black Bears was suggested to promote continued population growth and expansion (Unger 2007), and KDFWR has incorporated a Black Bear sanctuary within its three-county hunting zone to help achieve that objective. The Black Bear is a habitat generalist, tolerant of human intrusion, and well-adapted to take advantage of anthropogenic food sources. Consequently, the Black Bear is better able to coexist with people than most other bear species or large mammals (Cowan 1972, Herrero 2002). Nonetheless, Black Bears sometimes aggregate in the urban-wildland interface to take advantage of the abundant anthropogenic food sources (Beckmann and Berger 2003). Bear-human conflicts most often occur in or near human communities or in recreational areas frequented by people. Direct contact with Black Bears occurs when people feed them, or when Black Bears find available foods such as garbage, pet food, or wildlife feeders near human settlements, or food stored by campers in parks (Spencer et al. 2007, Whittaker and Knight 1998). Because it is difficult to change the behavior of Black Bears conditioned to human food and feeding (McCarthy and Seavoy 1994), these animals can quickly become a public nuisance and subsequent targets for wildlife hazing or euthanasia. Nuisance Black Bears are at increased risk of being killed illegally by local residents or euthanized by wildlife agencies for reasons of public safety (Gray et al. 2004). Human-Black Bear conflict often escalates during periods of natural food shortages or fluctuations 300 Northeastern Naturalist Vol. 20, No. 2 in seasonal availability, environmental degradation, and drought (Delozier and Stiver 2001). Black Bears that have recolonized Kentucky must cope with a more urbanized, populous, yet perhaps more tolerant human society than their predecessors faced a century ago. In Kentucky, like most states, Black Bears have widespread access to anthropogenic food sources. As the Black Bear population has increased in size and range, so has the number of bear nuisance complaints. In state and national parks in Kentucky, Black Bears have routinely raided trash cans in picnic areas and campgrounds, and been intentionally fed by people eager to interact with this charismatic animal (Harris 2011). In urban areas and surface mines, Bears have frequented garbage dumpsters and trash cans, damaged crops and apiaries, fed from wildlife feeders, and occasionally entered homes or vehicles in search of food. Since 1987, fifteen Black Bears have been euthanized as a result of habituation or aggressive behavior towards humans in Kentucky, and 3 others have been shot and killed as a result of defense of person or property claims. Bears raiding dumpsters located on surface mines seem to be particularly vulnerable to poaching and road collisions (University of Kentucky, Lexington, KY, unpubl. data). Although management and education efforts by KDFWR have increased to address Black Bear problems, wildlife managers, land stewards, and humans that live within Black Bear-occupied areas clearly face major challenges in coexisting with this large mammal. Knowledge of public opinion, values, and behavior towards wildlife can make the difference between success and failure in wildlife management (Kellert et al. 1996). The greater our understanding about the ecological importance of the Black Bear and the situations that precipitate human-Bear conflicts, the more likely it is that the Black Bear will be viewed as a public asset. Human tolerance of Black Bears and their subsequent adoption as ecological and economic flagships has occurred in a few Kentucky communities. For example, as Black Bear watching has increased at Kingdom Come State Park in Harlan County, the city of Cumberland tourism bureau has promoted the return of the species with signage on municipal buildings, police cars, and decorative tiles along pedestrian walkways, as well as rebranding the city as the selfproclaimed “Bear capital of Kentucky”. In addition, the city has formed a Black Bear task force to promote both the species and its community with a Black Bear festival, but also to educate people about Black Bear ecology and ways to reduce human-bear conflicts (Harris 2011). Finally, the Harlan County High School has adopted the Black Bear as the school’s mascot, with a prominent Black Bear paw print marking the adjacent hillside. The future of the Black Bear in Kentucky Return of the Black Bear to Kentucky presents both opportunities and challenges to its citizens. Despite increasing human population growth, habitat fragmentation, resource extraction, environmental degradation, lack of large-scale reintroduction attempts, and a general lack of political interest in landscape-scale conservation issues (Maehr 2001, Noss 2001, Schadt et al. 2002), the Black Bear has reoccupied much of its historic range in the Central 2013 D.E. Unger, et al. 301 Appalachians. This recolonization over the past 25 years is an important example of the ability of large mammals to repopulate areas from which they were extirpated. Such natural success stories are rare without intense human intervention, and even more so for large carnivores (Maehr et al. 2001). Factors that contribute to the success of species reintroduction also provide insights on the Black Bear’s successful return to Kentucky; the Black Bear is a native species and it is omnivorous (Smith and Clark 1994). Kentucky represents former core Black Bear range in the eastern United States (Hall 1981), and forests in this area are maturing after decades of intensive logging. Eastern Kentucky is similar to occupied habitat elsewhere (Leopold et al. 1998), and the recolonizing population appears to be genetically robust (Frary et al. 2011, Hast 2010). Human activity, with the exception of surface mining, is of relatively low intensity in areas of rugged topography (Watkins 1998). Potential source populations (Virginia, West Virginia, and Tennessee) are increasing (TWRA 2012, VDGIF 2012, Williamson 2002, WVDNR 2006), and the Black Bear has no natural predators or close competitors in the state. Perhaps most importantly, the primary threats of historical extirpation, overharvest and habitat destruction, have been reduced or eliminated with our better understanding of Black Bear population dynamics, habitat management, and the implementation of more sustainable harvest strategies. While the natural recolonization and continued expansion of the Black Bear in Kentucky is impressive, the species has returned to a landscape profoundly changed by extractive industries and human settlement. Decades of timber removal and subsequent forest recovery, changes in the fire regime, coal mining, introduction of exotic species, and human development have created an environment that is distinctly different from pre-Columbian eastern Kentucky, which was nearly entirely forested (Delcourt 2002). Anthropogenic environmental changes may have far-reaching and multifaceted effects beyond the immediately visible impact. Many diseases have or threaten to alter forest composition and structure in ways that reduce the availability and types of forage and denning structures. Surface mining has altered topography and stream ecology and created vast swaths of exotic grasslands in an otherwise forested landscape. Although changes to the Kentucky landscape over the past two centuries are profound, the recolonizing Black Bears are dispersing from populations that survived similar environmental changes and pressures in adjacent states. Consequently, these populations may have adapted to have a higher tolerance for human-altered environments that have aided in their persistence over time. Management recommendations Research conducted thus far and field observations by state wildlife agency personnel suggest the Black Bear populations in both southeastern and southcentral Kentucky are experiencing population growth and range expansion. We suggest that favorable forest conditions, greater tolerance by humans, and expanding source populations in neighboring states are the primary factors responsible for driving this occurrence. As a result, Kentucky now appears to have two small but relatively stable populations of Black Bears, although the reproductive core remains largely confined to areas near adjacent states with 302 Northeastern Naturalist Vol. 20, No. 2 historically larger Black Bear populations. If increased and sustainable harvest levels are desired, then continued bear population growth will need to occur over the next two decades given the species slow growth rate. Based on the small population size and the current limited range of females within each Black Bear population, we recommend continuation of a highly conservative harvest limit for the immediate future in Kentucky to avoid population declines of the species that have occurred elsewhere as a result of overharvest (Eastridge and Clark 2001). Continued research and monitoring of the two major Kentucky Black Bear populations will be important for making science-based management decisions so as to ensure long-term persistence of the species in Kentucky. In addition, as the Black Bear moves farther west in the state into more populous areas, public education will be vitally important to reduce human-bear conflic t. On a landscape scale, the continued recolonization of Kentucky could be the latest westward step in the genesis of a functional Black Bear metapopulation that spans networked forests in Kentucky and links populations in the Mississippi Valley of Arkansas and Missouri to those in the central Appalachian Mountains. Land acquisition and management that establishes movement corridors, creates additional Black Bear sanctuaries, and facilitates connectivity of other Black Bear populations to those in Kentucky could have far-reaching consequences not only for regional conservation of this species, but for other game species such as Elk and White-Tailed Deer (DeVivo et al. 2011, Yarkovich et al. 2011), and regional biodiversity at-large. We therefore highly recommend that wildlife managers and scientists use this unique opportunity to continue to study this recolonzation phenomenon so as to further our ecological understanding of not only the Black Bear, but community and population dynamics of large mammals in general. Acknowledgments We would like to thank all those that contributed in a wide variety of ways to the completion of this work from funding, fieldwork, and data collection to the actual writing and editing of this paper. 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