80000 Federal Register / Vol. 80, No. 246 / Wednesday, December 23, 2015 / Rules and Regulations

DEPARTMENT OF THE INTERIOR II. Major Provision of the Regulatory finding that listing the African Action subspecies () as a Fish and Wildlife Service This action revises the taxonomic threatened species was warranted and classification of the Asiatic lion proposed to list the subspecies as a 50 CFR Part 17 (currently classified as P. l. persica and threatened species under the Act. We also proposed a rule under section 4(d) listed as an under of the Act to provide conservation [Docket No. FWS–R9–ES–2012–0025; 450 the Act) to P. l. leo based on a measures for the African lion. 003 0115] taxonomic change. The P. l. leo subspecies will be listed as an Summary of Changes From the RIN 1018–BA29 endangered species and the P. l. Proposed Rule melanochaita subspecies will be listed Endangered and Threatened Wildlife We fully considered comments from as a threatened species in the List of the public and the peer reviewers on the and Plants; Listing Two Lion Endangered and Threatened Wildlife in Subspecies proposed rule to determine our final title 50 of the Code of Federal listing status of lion. This final rule AGENCY: Fish and Wildlife Service, Regulations (CFR) at 50 CFR 17.11(h). incorporates changes to our proposed Interior. This action will also add a rule under rule based on the comments we received section 4(d) of the Act for P. l. ACTION: Final rule. that are discussed under Summary of melanochaita which is set forth at 50 Comments and Responses and newly SUMMARY: We, the U.S. Fish and CFR 17.40(r). available scientific and commercial Wildlife Service (Service), determine Background information that became available after endangered status for the lion the close of the comment period. We The Endangered Species Act of 1973, subspecies Panthera leo leo and accept the as recommended as amended (Act) (16 U.S.C. 1531 et threatened status for P. l. melanochaita by the International Union for seq.), is a law that was passed to prevent under the Endangered Species Act of Conservation of Nature (IUCN) Species of species by providing 1973, as amended (Act). We are also Survival Commission Cat Classification measures to help alleviate the loss of publishing a concurrent rule under Task Force: P. l. leo (Asia and western, species and their . Before a plant section 4(d) of the Act. This rule central, and northern Africa) and P. l. or species can receive the provides for conservation measures for melanochaita (southern and eastern protection provided by the Act, it must P. l. melanochaita. Africa). Here we evaluate the status of first be added to the Federal List of the lion species (P. leo), which includes DATES: This rule is effective January 22, Endangered and Threatened Wildlife or the previously unreviewed population 2016. the Federal List of Endangered and of P. l. leo in India (formerly P. l. ADDRESSES: This final rule is available Threatened Plants in part 17 of title 50 persica). Additionally, we have on the Internet at http:// of the Code of Federal Regulations incorporated new population estimates www.regulations.gov and comments and (CFR). Section 4 of the Act and its and population trends for the lion into materials received, as well as supporting implementing regulations at 50 CFR part our Species Information section. documentation used in the preparation 424 set forth the procedures for adding Based on comments by peer reviewers of this rule, will be available for public species to these lists. and others, we revised the section on inspection, by appointment, during Previous Federal Actions trophy , providing additional normal business hours at: U.S. Fish and information on the practices that experts Wildlife Service; 5275 Leesburg Pike; In a final rule published in the have identified as undermining the Falls Church, VA 22041. Federal Register on June 2, 1970 (35 FR of trophy hunting, FOR FURTHER INFORMATION CONTACT: 8491), the Asiatic lion (currently listed recommended best practices and Branch of Foreign Species, Ecological under the Act as Panthera leo persica) reforms, biological impacts of trophy Services, U.S. Fish and Wildlife Service, was listed under the Act’s precursor, the hunting on lion populations, and MS: ES, 5275 Leesburg Pike, Falls Endangered Species Conservation Act of corruption in range countries, and Church, VA 22041–3803; telephone, 1969, as an endangered species and has expanded our assessment of the level of 703–358–2171; facsimile, 703–358– remained listed as an endangered threat that trophy hunting presents to 1735. If you use a telecommunications species under the Act. the species. Additionally, we have device for the deaf (TDD), call the On March 1, 2011, we received a incorporated information on , Federal Information Relay Service petition dated the same day from the corruption, traditional use of lion parts (FIRS) at 800–877–8339. International Fund for , and products, disease, and climate the Humane Society of the United change. Under the discussion of the 4(d) SUPPLEMENTARY INFORMATION: States, Humane Society International, rule in the preamble, we further clarify Executive Summary the Born Free Foundation/Born Free factors we will consider when making USA, Defenders of Wildlife, and the an enhancement finding for importation I. Purpose of the Regulatory Action Fund for requesting that the of sport-hunted trophies of P. l. We are listing two subspecies of lion, African lion subspecies be listed as melanochaita. Panthera leo leo and P. l. melanochaita, endangered under the Act. The petition Based on the information we received under the Endangered Species Act of identified itself as such and included and our assessment of that information, 1973, as amended (Act). We are listing the information as required by 50 CFR we have altered our finding. Some of the the P. l. leo subspecies as an endangered 424.14(a). On November 27, 2012, we information we received indicated species and the P. l. melanochaita published a ‘‘positive’’ 90-day finding threats may be worse than previously subspecies as a threatened species (77 FR 70727) indicating that we would indicated. Due to significant differences under the Act. We are also finalizing a initiate a status review of the African in the impacts of threats within the rule under section 4(d) of the Act that lion. species, we found that P. l. leo and P. will provide for conservation measures On October 29, 2014 (79 FR 64472) l. melanochaita qualify for different for P. l. melanochaita. we published in the Federal Register a statuses under the Act.

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Species Information and, consequently, by several mtDNA analyses should be backed up international organizations and by studies on nuclear DNA (nDNA, Taxonomy governing bodies. As a result, this is the inherited from both parents) and The lion (Panthera leo) was first classification on which the conservation morphological traits before assigning described by Linnaeus (1758, in Haas et of the species is largely based. However, taxonomic importance to them (Barnett al. 2005, p. 1), who gave it the name results of recent genetic research call et al. 2014, pp. 1, 8). Felis leo. It was later placed in the genus into question this classification. Recently, Mazak (2010, entire) Panthera (Pocock 1930, in Haas et al. In recent years, several genetic studies examined morphological characteristics 2005, p. 1). Although the classification have provided evidence of an of 255 skulls of wild and found of the modern lion as P. leo is accepted evolutionary division within lions in within the scientific community, there Africa (see Barnett et al. 2014, p. 6; considerable variation throughout the was a lack of consensus regarding lion Dubach et al. 2013, p. 746; Bertola et al. species’ range, with variation being intraspecific taxonomy (Mazak 2010, p. 2011 (entire); Antunes et al. 2008 greater within populations than between 194; Barnett et al. 2006b, p. 2120). (entire); Barnett et al. 2006a, pp. 511– them. However, according to Dubach et Based on morphology, traditional 512). These studies include analysis of al. (2013, p. 742), the genetic distinction classifications recognize anywhere from DNA samples from all major regions of of lions in southern and eastern Africa zero subspecies (classifying lions as one the species’ range, though some regions from those elsewhere in the species’ monotypic species) up to nine are sparsely represented. A major range is confirmed by results of studies subspecies (Mazak 2010, p. 194, citing genetic subdivision among lions occurs by Antunes et al. (2008, entire) which, several sources). The most widely in Africa, with lions in southern and in addition to analysis of mtDNA, also referenced of the morphology-based eastern Africa being distinct from and included analysis of nDNA sequence taxonomies is an eight-subspecies (six more diverse than lions elsewhere and microsatellite variation. extant) classification provided by (western and central Africa and Asia) The recent results of genetic research Hemmer (1974, in Nowell and Jackson (Figure 1). Lions in western and central renewed the debate on lion taxonomy 1996, p. 312; Barnett et al. 2006a, p. 507; Africa (as well as now-extinct North among the experts. For this reason, the Barnett et al. 2006b, p. 2120), which is African lions) are more closely related IUCN Species Survival Commission Cat recognized by the Integrated Taxonomic to lions in India than to lions in Specialist Group commissioned a Cat Information System (ITIS) (ITIS 2013, southern and eastern Africa (Barnett et Classification Task Force from among its unpaginated). It divides the lion species al. 2014, pp. 4–8; Dubach et al. 2013, expert members to reach a consensus on into: Panthera leo persica (India); P. l. pp. 741, 746–747, 750–751; Bertola et taxonomy for the group. As we leo, commonly referred to as the Barbary al. 2011, entire). According to Dubach et explained in our proposed rule, until lion (Morocco through Tunisia, extinct); al. (2013, p. 753), current range collapse the results of the IUCN Cat P. l. senegalensis (West Africa east to and fragmentation is too recent a Classification Task Force became the Central African Republic (CAR)); phenomenon to explain the reduced available, we concluded that the P. l. azandica (northern Zaire); P. l. genetic variability in these regions. taxonomy of the species was bleyenberghi (southern Zaire and Rather, the low genetic diversity in and presumably neighboring areas of unresolved, but, as required by the Act, between western and central African we based our status review in our and ); P. l. nubica (East Africa); lion populations suggests they have a P. l. krugeri (Kalahari region east to the proposed rule on the best available shorter evolutionary history than the scientific and commercial information, Transvaal and Natal regions of South more genetically diverse lions in Africa), and P. l. melanochaita, also which was the taxonomy that was most southern and eastern Africa (Bertola et widely recognized by taxonomic called the Cape lion (Cape region of al. 2011, p. 1362). Several authors argue experts: P. leo leo (African lion) and P. , extinct) (Nowell and that the origin of these genetically leo persica (Asiatic lion) and reviewed Jackson 1996, p. 312). distinct groups may be the result of the status of the petitioned entity, the In 1987, O’Brien (1987a, entire; regional and recolonizations African lion. 1987b, entire) reported the first results during major climate (and consequently of genetic studies conducted on lion biome) fluctuations during the In June 2015, after the close of the samples from some, but not all, regions Epoch (Barnett et al. 2014, comment period on our proposed rule, of the species’ range using early genetic pp. 5–8; Bertola et al. 2011, pp. 1362– IUCN posted an updated Red List techniques. Lions in India differed from 1364). Assessment for lion. In this assessment, lions in Africa, supporting a two- These findings on lion genetic a new two-subspecies classification is subspecies classification for extant relationships are based primarily on proposed based on the recommendation lions: P. l. leo and P. l. persica, the analysis of mitochondrial DNA of the IUCN Cat Classification Task African and Asiatic lion, respectively (mtDNA), which is inherited only from Force: P. l. leo of Asia (India) and (O’Brien et al. 1987, Meester and Setzer the mother. Because lions display sex- western, central, and northern Africa, 1971, Ellerman et al. 1953, in Dubach biased dispersal, in which males leave and P. l. melanochaita for southern and 2005, p. 16). According to Dubach their natal range and females tend to eastern Africa (Bauer et al. 2015a, (2005, p. 16), most taxonomic remain in their natal range, one would unpaginated) (Figure 1), which is authorities recognize this two- expect gene flow in females to be lower supported by Barnett et al. (2014, p. 6), subspecies taxonomy. This taxonomy than in males, resulting in greater Dubach et al. (2013, p. 746), Bertola et was also recognized by the International geographic differentiation in females al. (2011, entire), Antunes et al. (2008, Union for Conservation of Nature (Mazak 2010, p. 204). Consequently, entire), and Barnett et al. (2006a, pp. (IUCN) (Bauer et al. 2012, unpaginated) some authors state that results of 511–512).

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As required by the Act, and as taxonomic change by removing the Characteristics include sharp, retractile explained in our proposed rule, we base invalid subspecies P. l. persica. This claws, a short neck, a broad face with our listing determinations on the best entity is now included in the prominent whiskers, rounded ears and a muscular body. Lions are typically a tawny available scientific and commercial assessment of the lion species (P. leo). color with black on the backs of the ears and information. We accept the taxonomy as Species Description white on the abdomen and inner legs. Males recommended by the IUCN Cat usually have a mane around the head, neck Classification Task Force, which is The lion is the second-largest extant and chest. Lions are sexually dimorphic, supported by mtDNA analysis, as well cat species (second in size only to the with males weighing about 20–27 percent as analysis of nDNA sequence and tiger) and the largest carnivore in Africa more than females. Adult males, on average, microsatellite variation: P. l. leo (Asia (Ray et al. 2005, p. 67). As with other weigh about 188 kilograms (kg) (414 pounds and western, central, and northern (lbs)) with the heaviest male on record widely distributed large cats, there is weighing 272 kg (600 lbs). Females are Africa) and P. l. melanochaita (southern considerable morphological variation smaller, weighing, on average, 126 kg (278 and eastern Africa) (Figure 1) as the best within the species as a result of sexual lbs). The male body length, not including the available scientific and commercial selection, regional environmental tail, ranges from 1.7 meters (m) to 2.5 m (5.6 information. Because this new adaptations, and gene flow (Mazak feet (ft to 8.2 ft) with a tail from 0.9 m to 1 classification for lion includes 2010, p. 194). These include, among m (3 ft to 3.2 ft) (Nowell and Jackson, 1996). subspecies whose ranges span two others, variation in size, coat color and Lions in India tend to be smaller than continents, we assessed the status of the thickness, mane color and form, and those in Africa. Adult males weigh entire lion species (P. leo). skull characteristics (Mazak 2010, p. between 160–190 kg (353–419 lb), while Currently, the Asiatic lion (P. l. 194, citing several sources; Hollister females weigh between 110–120 kg persica) is listed as an endangered 1917, in Dubach 2005, p. 15). They are (243–265 lb) (Chellam in litt. in Nowell species under the Act. Based on the new described in the Convention on and Jackson 1996, p. 37). The record taxonomic classification for lions, we International Trade in Endangered total length for a male lion in India, are revising the List of Endangered and Species of Wild Fauna and Flora including the tail, is 2.92 m (9.6 ft) Threatened Wildlife at 50 CFR 17.11(h). (CITES) Periodic Review of the Status of (Sinha 1987 in Nowell and Jackson In the Regulation Promulgation section African Lion Across Its Range (CITES 1996, p. 37). One characteristic unique of this document, we implement a 2014, p. 3) as follows: to lions in India is a longitudinal fold

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of skin that runs along the belly and death can also occur during hunting 37). Lion reproduction in India appears (O’Brien et al. 1987, p. 100). attempts on some of their larger prey. to coincide with the fawning peak of Additionally, male lions in India do not Haas et al. (2005, entire) provide a deer (Axis axis) between have as large and full a mane as those summary of information on lion, December and January or with the in Africa, allowing their ears to always including the following: rutting season of chital and peak be visible, whereas the manes of male Prides vary in size and structure, but fawning for sambar deer (Cervus lions in Africa completely hide the ears typically contain 5–9 adult females (range, 1– unicolor) between May and June. (Nowell and Jackson 1996, p. 37; 18), their dependent offspring, and a Breeding lionesses may cue into these O’Brien et al. 1987, p. 100). coalition of 2–6 immigrant males (Heinsohn times of increased availability of food and Packer 1995; Packer et al. 1991).... sources to time births for maximum Pride sizes are smallest in arid environments survival of cubs (Banerjee and Jhala Historically, the species occurred in with limited prey species (Elliott and Cowan 2012, p. 1427). Average interbirth 1977; Hanby and Bygott 1979; Ruggiero 1991; interval is estimated to be 1.37 years; all habitats in Africa, except rainforest Schaller 1972; Stander 1992b; Wright and the hyper-arid interior of the Sahara 1960).... Males reside in a pride for however, if cubs of the previous litter (Ray et al. 2005, p. 66). Today they are [approximately] 2 years before being replaced survived to independence, it could be found primarily in savannas, although by another group of males (Packer et al. higher. After territorial takeovers and there are some remnant populations in 1988).... In the absence of a pride , females mated within an other habitat types (Riggio et al. 2013, p. takeover, males generally leave their natal average 4.8 months (Banerjee and Jhala 19). According to Nowell and Jackson pride when 2–4 years old (Bertram 1975b; 2012, p. 1424). Banjeree and Jhala (2012, (1996, p. 19), optimal habitat appears to Pusey and Packer 1987). Most females are p. 1424) found that the major cause of be open and thick bush, incorporated into their natal prides (Pusey cub mortality is infanticide due to and Packer 1987; Van Orsdol et al. territorial takeovers by adult males. scrub, and grass complexes, where 1985).... A small proportion of lions is sufficient cover is provided for hunting nomadic, including young and adult males Most observed adult mortalities (54.5 and denning. The highest lion densities without a pride. Nomadic lions follow the percent) were due to natural causes and are reached in savanna woodlands migrations of prey and hunt and scavenge 43 percent were due to human causes; plains mosaics of southern and eastern cooperatively (Bertram 1975a; Bygott et al. remaining mortalities were due to Africa (Ray et al. 2005, p. 66). The 1979; Schaller 1968, 1969; Van Orsdol et al. unknown causes. 1985). species is intolerant of anthropogenic Diet and Prey (human-caused) habitat conversion, . . . Lion productivity (measured as such as farming or overgrazing by number of surviving cubs) is limited by food. Lions are opportunistic hunters and . . . Cub mortality is high in lions and is scavengers. As scavengers, lions are (Ray et al. 2005, p. 66). In linked to periods of prey scarcity and India, the lion occurs in dry deciduous dominant and can usually readily infanticide by male lions during pride displace other predators from their kills forests (Meena et al. 2014, p. 121). Moist takeovers (Packer and Pusey 1983b; Schaller mixed and mixed forest habitats are 1972; Van Orsdol et al. 1985; Whitman and (Packer 1986, Schaller 1972, in Haas et critical to lions as they seek moist shady Packer 1997). al. 2005, pp. 4–5). As hunters, they are habitats that provide shelter from the . . . Lions are mainly active at night.... known to take a variety of prey. heat and cover to hide during peak [They] usually hunt in groups; males hunt However, they are also the largest times of human activities (Jhala et al. less frequently than do females, but males are carnivore in Africa and, as a result, 2009, p. 3391). stronger and can gain access to kills made by require large prey to survive. Ray et al. females (Bertram 1975a; Scheel and Packer (2005, pp. 66–67) summarizes lion prey General Biology 1991). Prey selection is related to seasonal in Africa as follows: weather patterns and the migration of large Lions are well studied. Much in some parts of Africa (Hanby et Lions are generalists and have been information exists on habits, behavior, al. 1995).... Lions exhibit individual recorded to consume virtually every and ecology of lions in Africa. CITES preferences in prey selection within and species larger than 1 kg in their range, as well (2014, p. 3) provides a general overview between prides in the same area (Rudnai as a wide variety of larger reptiles and birds 1973b; Van Orsdol 1984). (Nowell & Jackson 1996; Sunquist & Sunquist as follows: 2002). The constraints of large physical size Lions are generalist, cooperative hunters, Lion prides in India tend to be smaller and extended social groups, however, bind with foraging preferences changing with than those in Africa; most prides in them to large-bodied prey, and their diet is season and with lion group size. Lions live India contain an average of two females, dominated by medium-large . In in groups called ‘‘prides,’’ which are ‘‘fission- with the largest having five. Coalitions fact, only a few species of large ungulates fusion’’ social units with a stable of males will defend home ranges that comprise a majority of their diet wherever membership that sometimes divide into contain one or more groups of females, they occur (Schaller 1972; Stander 1992; small groups throughout the range. Lions but unlike lions in Africa, in India male Packer et al. 1995), and they are unable to persist in areas without large-bodied prey. have no fixed breeding season. Females give lions only associate with pride females birth every 20 months if they raise their cubs The threshold of this requirement is perhaps to maturity, but the interval can be as short when mating or on a large kill (Meena represented at , as 4–6 months if their litter is lost. Gestation 2009, p. 7; Nowell and Jackson 1996, p. , where Stander (1992) showed that lasts 110 days, litter size ranges 1–4 cubs, 37). Females are approximately 4 years lions hunting in pairs met their minimum and sex ratio at birth is 1:1. At about 4 years old at first reproduction, males 5–8 requirements hunting which, at of age, females will have their first litter and years (Banerjee and Jhala 2012, p. 1424; < 50 kg, are the smallest preferred prey males will become resident in a pride. Pride Nowell and Jackson 1996, p. 37). species recorded. takeovers by male lions and subsequent Banerjee and Jhala (2012, p. 1424) found In India, the lion’s diet is comprised infanticide of cubs sired by the ousted male that mating occurred throughout the of both small and medium prey, as well lions greatly influences reproductive success. year, but mostly in winter. Gestation as vulnerable livestock (Meena et al. Lionesses defending their cubs from the victorious males are sometimes killed during lasts 110 days; births peaked in the 2011, p. 61; Singh and Gibson 2011, p. the takeover. Infanticide accounts for 27 summer (April–May). Average litter size 1753; Meena 2009, p. 8). The most percent of cub mortality. Adult mortality is is 2.5 cubs, but as many as 5 have been commonly taken species is chital, which typically caused by humans, starvation, observed (Banerjee and Jhala 2012, pp. weighs approximately 50 kg (110 lb), disease, or attacks from other lions. Injury 1424, 1427; Nowell and Jackson 1996, p. and a larger species, the sambar deer

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(Meena et al. 2011, p. 63; Nowell and Gir , 6 per 100 km2 in al. (2009, p. 3391) found the average Jackson 1996, p. 37). The smaller size of Girnar Wildlife Sanctuary, and 2 per home range of a breeding group of the prey available in India may be 100 km2 in the surrounding agro- lionesses to be 33 km2. Similarly, Meena responsible for the smaller lion group pastoral land (Banerjee and Jhala 2012, (2009, pp. 7–8) found home ranges of sizes and less interaction between male p. 1421; Banerjee et al. 2010, p. 249). females and males to be 35 km2 and 85 and female groups (Meena 2009, p. 8; Aside from human-related mortality, km2, respectively. Nowell and Jackson 1996, p. 37). prey availability is likely the primary Historically, domestic cattle also determinant of lion density in Africa Range constituted a major portion of the lion’s (Fuller and Sievert 2001, in Winterbach The historical range of the lion et al. 2012, p. 7). In areas of low natural diet (Nowell and Jackson 1996, p. 37) included most current continental prey density, or high human contact, and remains a significant portion today African countries (Chardonnet 2002, pp. lions may prey on livestock (see (Meena et al. 2011, pp. 63, 64; Singh 25–28) and extended from Greece Human-Lion Conflict). and Gibson 2011, pp. 1753–1754). The through eastern Europe, southwest Asia proportion of wild prey and domestic Movements/Home Range (the Middle East), and India (Bauer et al. livestock in a lion’s diet may vary by Availability of prey is perhaps the 2015a, unpaginated; Nowell and Jackson season and between protected areas and primary factor that determines the 1996, p. 38). Lions have undergone peripheral areas (Meena et al. 2011, pp. ranging behavior of large carnivores dramatic range retraction from this 64, 65). (Gittleman & Harvey 1982, Van Orsdol historical distribution (Ray et al. 2005, Prey availability affects the et al. 1985, Grant et al. 2005, Hayward p. 67). Extirpation of lions in Europe reproduction, recruitment, and foraging et al. 2009, in Winterbach et al. 2012, occurred almost 2,000 years ago. The behavior of lions and, as a result, p. 4). Home-range sizes of lion prides species was extirpated from southwest strongly influences lion movements, correlate with lean-season prey biomass Asia within the last 150 years and abundance, and population viability (Van Orsdol et al. 1985, in Haas et al. northern Africa in the 1940s (Bauer et (Winterbach et al. 2012, p. 7, citing 2005, p. 4) and, therefore, vary widely al. 2015a, unpaginated; Black et al. several sources). Lion densities are among habitats. Average range sizes of 2013, p. 1; Nowell and Jackson 1996, p. directly dependent on prey biomass lion prides in Africa are 26–226 km2, 38). Today, lions occur only in Asia and (Van Orsdol et al. 1985, in Packer et al. but can be considerably larger (Stander sub-Saharan Africa (Table 1). In Asia, 2013, p. 636; Hayward et al. 2007, 1992b; Van Orsdol et al. 1985; Viljoen P. l. leo only remains in the Gir Forests entire). In Africa, lion densities range 1993, in Haas et al. 2005, p. 4). In areas of India. Within sub-Saharan Africa, P. from 8–13 lions per 100 square of low or variable prey biomass, annual l. leo and P. l. melanochaita remain in kilometers (km2) in Selous range requirements for a single lion 34 range countries (35 with South Reserve and up to 18 per 100 km2 in pride can exceed 1,000 km2 (Packer et Sudan, which gained its independence protected areas of eastern Africa and al. 2013, p. 636). Funston (2011, p. 5) as a country in July 2011) and have been South Africa (Creel and Creel 1997, found the home ranges of lion prides in recently extirpated from 12 African Nowell and Jackson 1996, in Haas et al. the dune-savanna habitat of Kgalagadi range countries and potentially 2005, p. 4). In India, densities are Transfrontier Park to range from 1,762 extirpated from another 4 (Bauer et al. estimated to be 15 lions per 100 km2 in to 4,532 km2. In India, however, Jhala et 2015a, unpaginated) (Table 1).

TABLE 1—RANGE COUNTRIES OF P. l. leo AND P. l. melanochaita [Information derived from Bauer et al. 2015a, unpaginated, IUCN 2006a, IUCN 2006b, and Chardonnet 2002]

Subspecies Countries

Panthera leo leo ...... Algeria 1, Benin, Burkina Faso, Cameroon, CAR, Chad, Congo 2, Coˆte d’Ivoire 2, DRC, Egypt 1, Gabon 2, Gambia 2, Ghana 3, Guinea 3, Guinea-Bissau 2, India, Liberia, Libya 1, Mali 2, Mauritania 2, Morocco 1, Niger, Nigeria, Senegal, Sierra Leone 2, Togo 3, Tunisia 1. Panthera leo melanochaita ...... Angola, , 2, Djibouti 2, Eritrea 2, , , 2, , , Na- mibia, 3, , South Africa, Sudan/, Swaziland, , , Zambia, . 1 Lions extirpated. 2 Lions considered recently extirpated (Bauer et al. 2015a). 3 Lions considered possibly extirpated (Bauer et al. 2015a).

The confirmed lion range in western classified by Bauer et al. (2015, pastoral land (Bauer et al. 2015a, Africa (the total size of protected areas unpaginted) as possibly extinct total unpaginated; Banerjee and Jhala 2012, where lions were confirmed) is approximately 1.8 million km2 (694,984 p. 1421; Jhala et al. 2009, pp. 3384, estimated at 49,000 km2, or 1.1 percent mi2), which is over half (52 percent) of 3385; Nowell and Jackson 1996, p. 38). of the historic range (Henschel et al. the range classified as extant by the Distribution and Abundance 2014, p. 5). The most recent estimate of previous estimate conducted by Riggio the lion’s range throughout Africa et al. (2013, p. 26), which was based on The general distribution of lions in comes from Bauer et al. (2015a, estimates of savanna habitat. The lion’s Africa is summarized by Ray et al. unpaginated) who estimate the extant range in Asia is estimated to be (2005, p. 67) as follows: lion range (areas reasonably confident approximately 10,500 km2 (4,054 mi2), Currently, lions are restricted mainly to that lions persist based on recent which occurs within the Gir National protected areas and surrounding records) to be approximately 1.6 million Park and Wildlife Sanctuary (Gir conservancies or ‘game management areas,’ 2 2 with the largest populations in East and km (617,763 mi ), or 8 percent of the Protected Area), Girnar Wildlife southern Africa. Where protection is poor, historical range in Africa. The areas Sanctuary, and surrounding agro- particularly outside protected areas, range

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loss or population decreases can be There have been few efforts in the past to 88 percent) concentrated in a single significant. Declines have been most severe estimate the number of lions in Africa. Myers population. in West and Central Africa, with only small, (1975) wrote, ‘‘Since 1950, their [lion] Bauer et al. (2015a, unpaginated) isolated populations scattered chiefly numbers may well have been cut in half, attempted to correct for outdated through the Sahel. Lions in the region are perhaps to as low as 200,000 in all or even declining in some protected areas and, with sources in Riggio et al. (2013) by less.’’ Later, Myers (1986) wrote, ‘‘In light of applying regional trends (discussed the exception of southern Chad and northern evidence from all the main countries of its Central African Republic, are virtually absent range, the lion has been undergoing decline below) to 2002 population estimates for from unprotected areas (Bauer 2003). in both range and numbers, often an central, eastern, and southern Africa Estimates of lion abundance on a large accelerating decline, during the past two from Bauer and Van Der Merwe (2004) geographical scale are few in number. decades.’’ In the early 1990s, IUCN SSC Cat and Chardonnet (2002); estimates for For a variety of reasons—including low Specialist Group members made educated western Africa were taken from densities, large ranges, cryptic ‘‘guesstimates’’ of 30,000 to 100,000 for the Henschel et al. (2014) because of the coloration, nocturnal and wary habits— African Lion population (Nowell and Jackson greater precision of their estimate. lions are difficult to count (Riggio et al. 1996). Applying regional trends to Bauer and 2013, p. 31; Bauer et al. 2005, p. 6). Van Der Merwe (2004) lion populations Ferreras and Cousins (1996, entire) There are large areas of the species’ estimates, Bauer et al. (2015a, range in which no data are available on provided the first quantitatively derived unpaginated; supporting information, lion occurrence or abundance (IUCN estimate of lion abundance in Africa Table 7) estimated lions in central 2006b, pp. 12–13). Species experts using a GIS-based model calibrated with Africa to be 590, eastern Africa to be recognize that estimating the size of the information obtained from lion experts. 7,345, and southern Africa to be 10,385 lion population in Africa is an Ferreras and Cousins predicted lion (Table 2). When regional trends were ambitious task, involving many abundance in Africa in 1980 to be applied to Chardonnet (2002) lion uncertainties (Bauer et al. 2012, 75,800. Later, four additional efforts— estimates, Bauer et al. (2015, unpaginated). Estimates, particularly Chardonnet (2002), Bauer and Van Der unpaginated; supporting information, throughout Africa or broad region-wide Merwe (2004), IUCN (2006a, 2006b), Table 7) estimated lions in central estimates tend to rely to a considerable and Riggio et al. (2013)—estimated lion Africa to be 1,748, eastern Africa to be extent on expert opinion or inference population sizes ranging from 23,000 to 13,316, and in southern Africa to be (Riggio et al. 2013, p. 21; Chardonnet 40,000 (Table 2). 15,925 (Table 2). In total, Bauer et al. 2002, p. 19). Consequently, there is a Between 2006 and 2012, Henschel et (2015, unpaginated) estimate the lion large degree of uncertainty in these al. (2014, p. 2) conducted field surveys population in Africa to be between estimates. In addition, to date all efforts in protected areas within designated 18,841 and 31,394. However, the to estimate the number of lions in Africa Lion Conservation Units (LCUs) of authors found that the study by Bauer have used different methods; therefore, western Africa to confirm lion presence and Van Der Merwe (2004) was more the results of earlier estimates cannot be where evidence of occurrence was conservative and stricter on data directly compared to those of later lacking and to establish population quality; therefore they have a greater estimates to determine population estimates where lions occurred. Lions confidence in an estimate closer to trend. were absent from protected areas in 5 of 20,000 lions in Africa. Additionally, the The earliest estimates of lion the 10 countries in western Africa lion population in India was estimated abundance in Africa were educated where lions were considered to be to be 445 by Bauer et al. (2015a, guesses made during the latter half of present (Henschel et al. 2014, p. 4). unpaginated). In 2015, the Government the 20th Century. Bauer et al. (2008, Henschel et al. (2014, p. 5) estimated of Gujarat completed its latest census, unpaginated) summarize the only 400 lions remain in the entire estimating 523 lions in India (BBC 2015, information as follows: western region, with most (about 350, or unpaginated) (Table 2).

TABLE 2—ESTIMATES OF LION ABUNDANCE [Rows may not tally due to rounding]

Western Africa Central Africa Eastern Africa Southern Source (percent of (percent of (percent of Africa (percent India Total total) total) total) of total)

Ferreras & Cousins 1996 (es- ...... 75,800 (18,600 in protected timate for lion abundance in areas). 1980). Chardonnet 2002 ...... 1,163 (3 per- 2,815 (7 per- 15,744 (40 19,651 (50 ...... 39,373 cent). cent). percent). percent). Bauer & Van Der Merwe 2004 850 (4 per- 950 (4 per- 11,000 (48 10,000 (44 ...... 23,000 cent). cent). percent). percent). IUCN 2006 1 (as calculated by 1,640 (5 per- 2,410 (7 per- 17,290 (52 11,820 (37 ...... 33,160 Riggio et al. 2013). cent). cent). percent). percent). Riggio 2013 (based on esti- 480 (1 per- 2,419 (7 per- 19,972 (57 12,036 (34 ...... 34,907 mates of savanna habitat). cent). cent). percent). percent). Henschel et al. 2014 ...... 406 (n/a) ...... Bauer et al. 2015a (trends ap- ...... 590 (3 per- 7,345 (39 per- 10,385 (55 ...... 18,726 * plied to Bauer and Van Der cent). cent). percent). Merwe 2004). Bauer et al. 2015a (trends ap- ...... 1,748 (6 per- 13,316 (42 15,925 (51 ...... 31,394 * plied to Chardonnet 2002). cent). percent). percent). Bauer et al. 2015a ...... 445

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TABLE 2—ESTIMATES OF LION ABUNDANCE—Continued [Rows may not tally due to rounding]

Western Africa Central Africa Eastern Africa Southern Source (percent of (percent of (percent of Africa (percent India Total total) total) total) of total)

Government of Gujarat ...... 523 2015 **. 1 Estimates were made for individual Lion Conservation Units (defined management units), and were given as population size classes rather than specific figures. As calculated by Riggio et al. * Total includes estimate for western Africa taken from Henschel et al. (2014). ** As reported in BBC 2015, unpaginated.

As previously stated, extant lion 2013, p. 636; Funston 2011, p. 3; all fenced populations, and most were populations are limited to protected Ferreira and Funston 2010, pp. 201, reestablished in the past 20 years and areas. These populations are largely 203). quickly reached capacity (Bauer et al. isolated and many are small. P. l. leo Bauer et al. (2015a, unpaginated), 2015b, pp. 1–2). Populations in eastern (totaling approximately 1,500 lions), is using a time trend analysis of census Africa decreased overall by 59 percent divided into 15 populations in and data, determined the trend of lion (Bauer et al. 2015a, unpaginated). The around protected areas; of these, 14 are populations from 1993 to 2014. Overall, population was the only large remaining populations from a total of 38 these lion populations decreased by 43 population surveyed that did not historical occurrences in western and percent in 21 years (Table 3). However, decrease. Katavi National Park central Africa, while one occurs in India the authors found significant regional experienced complete loss of lions from (Bauer et al. 2015a, unpaginated; differences. In Asia, the single an estimated 1,118 in 1993 to zero in Henschel et al. 2015b, unpaginated; population increased by 55 percent 2014 (Bauer et al. 2015a, unpaginated, Brugie´re et al. 2015, p. 515; Henschel et (Bauer et al. 2015a, unpaginated). The supporting information Table 3; Bauer al. 2014, pp. 4–5; Jhala et al. 2009, p. population inside the protected area has et al. 2015b, p. 1). Western and central 3384). Nearly 90 percent of the lions in stabilized and expanded into Africa (combined) experienced the western Africa persist in a single surrounding agro-pastoral land (Bauer et largest decline at 66 percent (Table 3). population, the W-Arly-Pendjari (WAP) al. 2015b, p. 2; Breitenmoser et al. 2008, All populations are declining, except Complex (Henschel et al. 2014, p. 5). unpaginated). Additionally, the 2015 the population in Pendjari; populations Based on Bauer et al. (2015a, census of Gir Sanctuary and in Comoe´ and Mole are now likely unpaginated; Supporting Information, surrounding forest areas showed a 27 extinct (Bauer et al. 2015a, unpaginated, percent increase from the 2010 census Table 3) and Bauer and Van Der Merwe supporting information Table 3; Bauer (The Guardian 2015, unpaginated). In (2004, pp. 28–30), most P. l. et al. 2015b, p. 1). Furthermore, almost southern Africa, the sample populations melanochaita occur in approximately 68 all lion populations in Africa that overall increased by 8 percent (Bauer et protected areas throughout southern and historically exceeded 500 individuals, al. 2015a, unpaginated). However, one eastern Africa, with larger populations the minimum number estimated to occurring in Botswana, Kenya, Namibia, of the largest populations, Okavango, and populations of 6 unfenced reserves constitute a viable population South Africa, Tanzania, Zambia, and (according to Riggio et al. 2013, p. 32 Zimbabwe. are declining (Bauer et al. 2015a, unpaginated, supporting information and Bjo¨rklund in Riggio et al. 2013, p. Population Trends Table 3; Bauer et al. 2015b, p. 1). Fifteen 32), are declining (Bauer et al. 2015b, p. Based on the best available of the 23 sample populations in 1). information, lion range and numbers southern Africa were fenced; none Although these trends are based on 47 have clearly declined over the past experienced sharp declines and many sample populations, they comprise a several decades. However, not all lion small fenced populations are increasing substantial portion of the total populations have declined—some have (Bauer et al. 2015a, unpaginated, remaining lion populations; therefore, increased or remained stable, and some supporting information Table 3; Bauer the authors are confident in applying have been restored to areas from which et al. 2015b, p. 1). South Africa was the the observed trends to regions and the they were previously extirpated (Bauer only African country with growth in species as a whole (Bauer et al. 2015a, et al. 2015a, unpaginated; Packer et al. every population. However, these were unpaginated).

TABLE 3—REGIONAL TRENDS FOR 47 MONITORED LION POPULATIONS FROM 1993–2014 [Bauer et al. 2015a, unpaginated; supporting information Table 7].

Estimated lions in sample populations Percent Region change 1993 2014

Asia ...... 312 485 +55 Southern Africa ...... 4,887 5,265 +8 Eastern Africa ...... 3,112 1,266 -59 Western and Central Africa ...... 1,304 439 ¥66

Total ...... 9,615 7,455 ¥22%

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Using these rates of change, the protected areas where lions occur will habitat loss is partly responsible for the authors calculated that the population be under increased pressure as more decline of lions to a single population in 5 countries (Botswana, India, land is needed to satisfy the agricultural in a protected area. However, due to Namibia, South Africa, and Zimbabwe), needs of the human population. good protection and management, lions or 25 percent of the lion’s range, Inadequate management and law have dispersed to forested areas outside increased by 12 percent, while the enforcement has led to of the the protected area, extending their range population in the remaining 75 percent lion’s prey base in Africa for , from an initial 1,883 km2 to 10,500 km2 of the range decreased by 60 percent which has been critically depleted. (Johningh et al. 2007, Singh 2007, and (Bauer et al. 2015a, unpaginated), Additionally, human population growth Divyabhanusinh 2005, in Banerjee et al. resulting in a 43 percent population in Africa has led to human-lion conflict, 2010, p. 248; Singh 2007, in Jhala et al. decrease of the entire lion species particularly on the edge of protected 2009, p. 3384). Farming has been between 1993 and 2014. areas, when pastoralists invade encouraged in the area and has The growth rate estimates discussed protected areas to allow their herds to flourished. Cultivated areas have above are the best available information graze or when lions move out of created refuge areas and corridors for on global trends for lion populations, protected areas in search of prey, often lion movement (Vijayan and Pati 2001 although Bauer et al. (2015b, p. 2) preying on domestic livestock. Human- in Meena et al. 2014, p. 124). At this caution that these numbers are rough lion conflict leads to indiscriminate time, no information indicates habitat estimates. However, it is unlikely that killing of lions, primarily as a result of loss is currently threatening the lion regional declines are a product of retaliatory or preemptive actions to population in India. In Africa, however, differences in methodological protect livestock and human lives. The despite lions being mainly found in shortcomings. Sample populations are close proximity of lions to humans and protected areas, habitat loss and all monitored with at least partial domestic livestock throughout their degradation continue to be among the protection. Research sites are known to range exposes them to diseases, mainly main threats to lions (IUCN 2006a, p. be generally avoided by poachers and transmitted through livestock and 18; Ray et al. 2005, pp. 68–69). encroachers. Therefore, the estimated domestic dogs, which can impact The main cause of lion habitat loss growth rates may be less optimistic. It general fitness, reproduction, and and degradation is expansion of human is likely that unmonitored, unfenced lifespan. These are in addition to settlements and activities, particularly populations will have suffered greater diseases that naturally occur in lion due to agriculture and intensive rates of decline than reported since lack populations in Africa. Furthermore, in livestock grazing (IUCN 2006a, p. 18; of management generally means a lack some areas of Africa improper IUCN 2006b, p. 23; Ray et al. 2005, pp. of conservation effort (Bauer et al. management has resulted in reduced 68–69; Chardonnet 2002, pp. 103–106). 2015b, p. 3). lion numbers due to excessive lion From 1970 to 2000, the human The work of Packer et al. (2013a, pp. harvests from trophy hunting. population in sub-Saharan Africa 639–640) predicts future declines Subsequently, some lion populations increased by 126 percent (from 282 within a number of protected areas. are negatively impacted by infanticide million to 639 million) (United Nations Bauer et al. (2015b, p. 2) found that if following pride takeovers by new males. (UN) 2013, p. 9), while at about the regional trends remain unchanged in the Because habitat loss has resulted in same time (1975 to 2000), agriculture future, lions in western and central small, isolated populations across its area increased by 57 percent (from just Africa would likely lose a third of their range, lions face threats from stochastic over 200 million ha to almost 340 population in 5 years and half of their events, such as a disease epidemic and million ha) and natural vegetation in the population in 10 years. The population inbreeding depression. An emerging region decreased by 21 percent (Brink in eastern Africa is likely to decline by threat to lions is trade in bones and and Eva 2009, p. 507). In 2009, a third in 20 years and half in 30 years. other body parts for traditional approximately 1.2 billion ha, or 40 The Okavago population, Botswana, medicine. These causes of lion percent, of Africa’s land area was in will also likely decline by a third in 20 population declines are widespread and permanent pasture or crops, with the years (Bauer et al. 2015b, p. 2). Many likely to continue. The impacts of these vast majority (31 percent) in pasture lion populations are expected to threats are likely to be exacerbated by (UNEP 2012b, p. 68). Riggio et al. (2013, disappear within the next few decades climate change. Projected changes p. 29) estimate the original extent of such that the intensely managed indicate negative impacts to available savanna habitat in Africa to be populations in southern Africa will habitat and, therefore, the range of the approximately 13.5 million km2. Based replace savanna landscapes as sites for lion, prey availability, and the number on an analysis of land-use conversion the most successful conservation of of disease outbreaks as well as and human population densities, Riggio lions. susceptibility to those diseases. et al. (2013, p. 29) found current savanna habitat that is suitable for lions Summary of Threats Habitat Loss to be fragmented and to total about 3.4 Today, lions are mainly restricted to Habitat destruction and degradation million km2 (or 25 percent of African protected areas; however, they still face have been extensive throughout the savanna habitat). This indicates a serious threats that stem from range of the lion, resulting in local and substantial decrease in lion habitat over inadequate management of those areas regional lion population extirpations, the past 50 years and explains, in part, and increasing pressure on natural reduced lion densities, a dramatically why lions are limited to protected areas. resources to meet the needs of a growing reduced range (see Range), and small, Based on a comparison of land-use human population. Habitat loss has fragmented, and isolated lion and human population data, Riggio et been extensive throughout the range of populations that are increasingly al. (2013, p. 23) determined that a the lion, resulting in local and regional limited to protected areas (see density of 25 or more people per km2 lion population extirpations and a Distribution and Abundance) (Singh served as a proxy for the extent of land- dramatically reduced range with 2007, in Jhala et al. 2009, p. 3384; Ray use conversion that would render isolated lion populations that are et al. 2005, p. 69; Bauer and Van der habitat unsuitable for lions. Woodroffe increasingly limited to protected areas. Merwe 2004, pp. 29–30; Nowell and (2000, p. 167) analyzed the impact of As the human population increases, the Jackson 1996, pp. 20–21). In India, people on predators by relating local

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carnivore extinctions to past and pp. 98–99; Newmark 2008, pp. 322–324; Tanzania, and Uganda (UN 2015, p. 4). projected human population densities IUCN 2006b, pp. 20–22; Ogutu et al. Africa has the fastest population growth and estimated 26 people per km2 as the 2005, entire). Mesochina et al. (2010a, p. rate in the world (UN 2015, pp. 3, 9; mean human density at which lions 74) state that widespread destruction of UNEP 2012a, p. 2), and future went locally or regionally extinct. In wildlife habitat and human population growth in sub-Saharan 1960, 11.9 million km2 of the original encroachment in wildlife corridors are Africa is projected to be large and rapid 13.5 million km2 of savanna habitat had major threats to lion conservation in (UN 2013, p. 9). By 2100, Angola, fewer than 25 people per km2; however, Tanzania and consider loss of suitable Burundi, DRC, Malawi, Mali, Niger, in 2000 that number decreased to 9.7 habitat as a top threat to lion survival in Somalia, Uganda, Tanzania and Zambia 2 million km (Riggio et al. 2013, p. 29). the country. The Kenya Wildlife Service are projected to increase by at least five- Expansion of human settlements, indicates that habitat loss due to land- fold (UN 2015, p. 9). agriculture, and/or livestock grazing are use changes and human encroachment reported as occurring in or on the into previously wild areas is having a By 2050, the UN projects the human periphery of several areas identified by major impact on lion range size in population of Tanzania to almost triple Riggio et al. (2013, suppl. 1) as lion Kenya (Kenya’s National Large its 2010 population, reaching a density strongholds (viable populations) and Carnivore Task Force 2010, p. 21). of 137 people per km2, whereas Kenya’s potential strongholds (IUCN 2006a, p. In southern Africa, the extent of population is projected to more than 16; IUCN 2006b, pp. 20–22), and are current habitat destruction and double, reaching a density of 167 people particularly a threat in western, central, degradation appears to vary widely. For per km2 (Table 4). Human population and eastern Africa and some parts of example, according to the Zambia growth, and resulting pressures exerted southern Africa. Expansion of Wildlife Authority (2009 pp. 4–5), on habitat, are also expected to vary agriculture and livestock grazing are unplanned human settlement and other widely in the southern region. reported in or around two of the larger land-use activities in game management Population increases from 2010 to 2050 populations of P. l. leo in Africa, WAP areas are a major threat to the long-term are projected to range from about 23 Complex and a Chad-CAR population survival of the lion in Zambia. They percent (South Africa) to well over 200 (Heschel et al. 2014, pp. 5–6; Houessou note that conversion of natural habitat percent (Zambia), with 2050 densities in et al. 2013, entire; Chardonnet et al. in game management areas for cropping the region ranging from 5 people per 2010, pp. 24–26; IUCN 2008, pp. 8, 28– and grazing of livestock has led to km2 (Botswana and Namibia) to 432 29); management in portions of both is habitat destruction and indicate that people per km2 (Uganda) (Table 4). The reported as weak (Heschel et al. 2014, elimination of tsetse flies and human populations of most other pp. 5–6; IUCN 2008, p. 8). Eastern subsequent increase in pastoralist current and recent lion range countries Africa contains approximately 40 activities in game management areas are also expected to have very high percent of all the lions in Africa (Table places the lion under renewed direct growth rates (Table 4). The country- 2). Seven of the seventeen major P. l. conflict with humans. On the other wide human population densities melanochaita populations identified by hand, according to Funston (2008, pp. provided here (and in Table 4) are not Riggio et al. occur in eastern Africa; six 123–126), in several areas of southern of which occur in Tanzania and Kenya. Africa where lions were recently directly comparable to the density Between 1990 and 2010, Kenya’s human extirpated, lions are reestablishing as a thresholds determined by Riggio et al. population grew from 23 million (40/ result of, among other factors, adequate (discussed above) due to the differences km2) to 41 million (70/km2), whereas protection of habitat and prey. in scale at which they were made. Tanzania’s grew from 25 million (27/ Projections of future growth in human However, country-wide population km2) to 45 million (48/km2) (UN 2013, populations, areas converted to densities relate the number of humans pp. 421, 798). Not unexpectedly, agriculture, and livestock numbers to land area and, consequently, are expansion of agriculture and livestock suggest suitable lion habitat will indicative of the level of pressure that grazing is occurring in these countries continue to decrease across its range will exist to convert land to uses that (Brink et al. 2014, entire; UNEP 2009, p. into the foreseeable future. Between will meet the needs of the human 91; Mesochina et al. 2010a, p. 74), 2015 and 2050, half of the world’s population. This situation is including in or around these major population growth is expected to occur particularly the case given that much of populations (Ogutu et al. 2011, entire; in 9 countries, 6 of which are within the sub-Saharan Africa is rural and locals Mesochina et al. 2010a, pp. 71–74, 76; lion’s range (India, Nigeria, Democratic depend on agriculture for their Packer et al. 2010, pp. 8–9; UNEP 2009, Republic of the Congo (DRC), Ethiopia, livelihood.

TABLE 4—HUMAN POPULATION PROJECTIONS IN COUNTRIES CONTAINING THE 47 SAMPLE LION POPULATIONS USED BY BAUER ET AL. (2015), EXCEPT COˆ TE D’IVOIRE AND GHANA WHERE LIONS ARE CONSIDERED EXTIRPATED [Population data is from UN 2013]

UN Population estimate, in thousands 2) Subspecies Country (people/km 1950 2010 2050 2100

P. l. leo ...... India ...... 376,325 1,205,625 1,620,051 1,546,833 (114) (367) (493) (471) Benin ...... 2,255 9,510 22,137 32,944 (20) (84) (197) (293) Burkino Faso ...... 4,284 15,540 40,932 75,274 (16) (57) (149) (275) Cameroon ...... 4,467 20,624 48,599 82,393 (9) (43) (102) (173) Nigeria ...... 37,860 159,708 440,355 913,834

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TABLE 4—HUMAN POPULATION PROJECTIONS IN COUNTRIES CONTAINING THE 47 SAMPLE LION POPULATIONS USED BY BAUER ET AL. (2015), EXCEPT COˆ TE D’IVOIRE AND GHANA WHERE LIONS ARE CONSIDERED EXTIRPATED—Continued [Population data is from UN 2013]

UN Population estimate, in thousands 2) Subspecies Country (people/km 1950 2010 2050 2100

(41) (173) (477) (989) Senegal ...... 2,477 12,951 32,933 58,180 (13) (66) (167) (296) P. l. melanochaita ...... Kenya ...... 6,077 40,909 97,173 160,423 (10) (70) (167) (276) Tanzania ...... 7,650 44,973 129,417 275,624 (8) (48) (137) (292) Botswana ...... 413 1,969 2,780 3,025 (1) (3) (5) (5) Mozambique ...... 6,442 23,967 59,929 112,018 (8) (30) (75) (140) Namibia ...... 485 2,179 3,744 4,263 (1) (3) (5) (5) South Africa ...... 13,683 51,452 63,405 64,135 (11) (42) (52) (53) Uganda ...... 5,158 33,987 104,078 204,596 (21) (141) (432) (849) Zambia ...... 2,372 13,217 44,206 124,302 (3) (18) (59) (165) Zimbabwe ...... 2,747 13,077 26,254 32,608 (7) (33) (67) (83)

Although urbanization is increasing survival (see Loss of Prey Base) and Nations Environment Programme in sub-Saharan Africa, the majority of increased human-lion conflict resulting (UNEP) 2012a, p. 3; Chardonnet et al. the population is rural (UN 2014, p. 20). in lion mortality (see Human-Lion 2010a, p. 19; International Assessment About 60–70 percent of the sub-Saharan Conflict)—two of the main factors that of Agricultural Knowledge, Science and population relies on agriculture and influence the distribution and Technology for Development (IAASTD) livestock for their livelihood (UNEP population viability of large carnivores 2009, pp. 3–4, 8; United Nations 2006, pp. 82, 100, 106; IAASTD 2009, p. such as lions (Winterbach et al. 2014, p. Economic Commission for Africa 2008, 2). Much of the agriculture and 1; Riggio et al. 2013, p. 18). Ray et al. pp. 3–5). The result of this process is livestock-raising is at subsistence level (2005, p. 69) note that, although lions accelerated transformation of natural (IAASTD 2009, pp. 8, 28). As a result, have a wide tolerance for habitats, they landscapes at the expense of wilderness a large portion of the growing are generally incompatible with humans that sustains species such as lions and population will depend directly on and human-caused habitat alteration their prey (Chardonnet et al. 2010a, p. expansion of agriculture and livestock and loss; they are the least successful 19). grazing to survive. Between 2010 and large African carnivore outside Urbanization is also increasing in 2050, the population of sub-Saharan conservation areas (Woodroffe 2001, in India, but like sub-Saharan Africa, the Africa is projected to more than double Winterbach et al. 2012, p. 6). Further majority of the population is rural (UN to more than 2 billion (from 831 million fragmentation and isolation of lion 2014, p. 22; Swain et al. 2012, p. 1). In to 2.1 billion) (UN 2013, p. 9). During habitat and populations can also impact the State of Gujarat, 70 percent of all about this same time period (2005 to dispersal and genetic viability (see workers are rural based, with almost 52 2050), the area of cultivated land is Deleterious Effects Due to Small percent being cultivators and projected to increase by 51 million ha Population Sizes). agricultural laborers (Swain et al. 2012, (approximately 21 percent) Large carnivores with low potential p. 1). Suitable lion habitat within the (Alexandratos and Bruinsma 2012, p. for cohabitation with humans have a Gir Protected Area appears to be secure; 107). However, this figure does not high risk of local extinction. In order to however, habitat outside this area that is include range land, and the majority of survive, they require larger contiguous vital for dispersal may experience agricultural land in Africa is devoted to habitats with fewer negative human increasing pressure in the future. grazing (UNEP 2012b, p. 68). The impacts than do more resilient species Dispersal corridors and resource-rich number of livestock (cattle, sheep, and (Winterbach et al. 2012, p. 5). As human habitats outside the protected area are goats) in sub-Saharan Africa is projected populations continue to rise in sub- important to avoid inbreeding to increase about 73 percent, from 688 Saharan Africa, the amount of land depression and extirpation of the lion million to 1.2 billion, by 2050 required to meet the needs of those population from stochastic events. Due (Alexandratos and Bruinsma 2012, p. populations is constantly increasing to the population growth of lions in 133). (Brink et al. 2014, entire; Brink and Eva India, there is increased movement, Expansion of human settlements and 2009, entire; Eva et al. 2006, p. 4), a dispersal, and establishment of lion in activities into lion habitat renders the problem accentuated by slow rates of natural habitats outside the protected habitat unsuitable for lions primarily technological progress in food area. Twenty-five percent of the lion because it results in reduced availability production and land degradation from population is found in Girnar Wildlife of the wild prey that lions depend on for both overuse and natural causes (United Sanctuary, coastal areas, and natural

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habitats along the Shetrunji River to be a concern for the lion population sustenance, leading to increased northeast of Gir (Meena 2014, p. 27). as conservation initiatives have ensured presence of hunters in protected areas Additionally, the size of the Gir availability of ample prey (Banerjee et and higher than average declines in Protected Area implies that dispersing al. 2010, p. 249; Khan et al. 1996 and wildlife (Brashares et al. 2004, pp. lions will inevitably cross the protected Singh and Kamboj 1996 in Meena 2010, 1180–1181). area boundaries (Meena 2010, p. 212). p. 209; Jhala et al. 2009, p. 3384). The The sale of bushmeat is an important When lions move, they must cross semi-nomadic pastoral communities livelihood in Africa (Chardonnet et al. heavily populated human settlements that inhabit the Gir Forests are primarily 2010, p. 27; Mesochina et al. 2010a, p. and agricultural fields (Meena 2010, p. vegetarian (Banerjee et al. 2013, p. 2); 38; Abwe and Morgan 2008, p. 26; 209). Traditional land uses are quickly therefore, there is no great demand for Bennett et al. 2007, p. 885; Fa et al. changing in the region due to limestone bushmeat. However, in most African 2006, p. 507). The little meat produced mine and infrastructure development countries, large carnivores such as lions from domestic livestock is unaffordable (Banerjee et al. 2010, p. 250). are under serious threat through for common people (Bouche´ et al. 2010, Additionally, tourist activities (safaris to decreased prey abundance (Bauer et al. p. 1001). Bushmeat hunting is rarely see the lions and religious pilgrimages 2014, p. 97) due to unsustainable and practiced solely for subsistence. It to visit temples located within and on increasingly commercialized bushmeat supplies meat for local consumption the border of protected areas) can have hunting in and around protected areas and trade, urban markets, and even detrimental impacts to wildlife if not (Bauer et al. 2015a, unpaginated; international markets (Lindsey et al. carefully planned. For example, Henschel et al. 2015, unpaginated; 2013b, pp. 86–87). Outlets for the sale construction of a road has been Henschel et al. 2014, p. 5; Lindsey et al. of bushmeat have arisen in some areas, proposed to circle the outside of the 2013b, p. 84; Lindsey and Bento 2012, and full-time commercial bushmeat whole Gir Protected Area System pp. 1–2, 61; Scholte 2011, p. 7; Bouche´ traders occur in most southern and (Meena 2014, p. 28). Altering this et al. 2010, pp. 1000, 1001; Cragie et al. eastern African countries (Lindsey et al. habitat would result in land-use 2010, p. 2227; Brashares et al. 2004, p. 2013b, p. 86). Significant distribution of changes, promoting rapid development 1181; Fischer and Linsenmair 2001, pp. bushmeat to Europe and the United and urbanization and thereby 132, 133). States, where it is sold at elevated disconnecting corridors for lion Humans in Africa rely on protein prices, drives increasing movement (Meena 2014, p. 28; Banerjee obtained from bushmeat, resulting in commercialization of trade, a greater et al. 2010, p. 250). Furthermore, direct competition for prey between number of hunters, adoption of more humans and lions, and commercial crossing these areas renders lions more efficient hunting methods, and an poaching of wildlife is becoming a vulnerable to disease transmission (See unprecedented pressure on wildlife significant threat to many species, Disease below) and conflict with populations (Stiles 2011 and Barnett including those that lions rely upon for humans (see Human-Lion Conflict 2000 in Lindsey et al. 2013b, p. 88). food. Subsistence hunting was below). Because lions are social and Many illegal hunters are poor (Barnett traditionally carried out with the use of territorial, they need adequate space to 2000 in Lindsey et al. 2013b, p. 88; spears, which had minimal impact to survive. Lack of adequate habitat will Lindsey and Bento 2012, p. 37; Scholte have a bearing on the lion’s ecology, wildlife populations. Spears have since 2011, p. 7). Bushmeat trade can provide behavior, and population structure been replaced by automatic weaponry a quick income to purchase other food (Meena 2014, p. 28). (Chardonnet et al. 2010, p. 27) and Growing human populations have snares, which are most commonly used and essentials (Lindsey et al. 2013b, p. been associated with declines in large (Lindsey et al. 2013b, p. 83). These 82; Lindsey and Bento 2012, p. 62). carnivore populations all over the methods allow for poaching of large Hunters are wealthier than non-hunters world, and high human density is numbers of animals for the bushmeat (Knapp 2007 in Lindsey et al. 2013b, p. strongly associated with local trade, particularly snares, which are 86) and enjoy elevated social status. extirpation of large carnivores (Linnell cheap, difficult to detect, and This growing demand and the et al. 2001, Woodroffe 2001, in unselective as they can kill nontarget availability of modern weapons have led Woodroffe and Frank 2005, p. 91; animals ranging from rodents to to many African wildlife species being Woodroffe 2000, entire). Chardonnet et (Lindsey et al. 2013b, p. 83). hunted at unsustainable levels and the al. (2002, p.103) indicate that the The human population in a majority lion prey base becoming depleted in distribution maps of lion of African countries within the range of many areas (Hoppe-Dominik et al. 2011, subpopulations tend to confirm a direct the lion has quadrupled since the 1960s p. 452; Chardonnet et al. 2010, pp. 6, inverse correlation of lion density and (Riggio et al. 2013, p. 29; IUCN 2009, p. 13–14, 27; Packer et al. 2010, p. 8; Frank numbers with human activity and 15), increasing the demand for et al. 2006, p. 12). Because wildlife has presence. Further, Packer et al. (2013a, bushmeat. Bushmeat contributes been depleted in non-protected areas, entire) found that lions in unfenced significantly to food security, and is illegal bushmeat hunters are reserves are highly sensitive to human often the most important source of increasingly focusing efforts on population densities in surrounding protein in rural areas (Nasi et al. 2008 protected areas (Lindsey et al. 2013b, p. communities. in Lindsey et al. 2013b, p. 82). It 84). Weak management effectiveness comprises between 6 percent (southern and inadequate law enforcement have Loss of Prey Base Africa) and 55 percent (CAR) of a facilitated poaching for bushmeat in One of the most important human’s diet within the lion’s range in protected areas and resulted in a requirements for carnivore survival, Africa (Chardonnet et al. 2005, p. 9; widespread decrease in large mammal including lion, is prey availability, as it IUCN 2006b, p. 19). In western Africa, populations, including lion prey, in affects reproduction, recruitment, and bushmeat is a secondary source of these areas (Henschel et al. 2015b, foraging behavior and, therefore, also protein, with fish being the primary unpaginated; Henschel et al. 2014, pp. impacts lion movement, abundance, and source. However, when widespread loss 5, 7; Lindsey et al. 2013b, pp. 84, 88; population viability (Winterbach et al. of jobs and income occurs due to poor Lindsey and Bento 2012, p. 61; Scholte 2012, p. 7, citing several sources). In fish harvests, bushmeat becomes an 2011, p. 7; Bouche´ et al. 2010, pp. 99, India, prey abundance does not appear important source of income and 1001; Brashares et al. 2004 in Craigie et

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al. 2010, p. 2227; Fischer and where human population growth rates and Bento 2012, p. 63). Given the Linsenmair 2001, p. 134). are high, encroachment and habitat loss widespread and significant decrease in Significant decreases in prey occurs, and people are dependent on lion prey throughout its range in Africa, abundance have occurred in protected bushmeat. Protected areas in Ethiopia, it is apparent that enforcement of laws areas throughout Africa (Lindsey et al. Mozambique, Tanzania, and Zambia are and regulations is not adequate. 2013b, pp. 84, 85; Scholte 2011, pp. 2, increasingly settled (Lindsey et al. Additionally, weak management of 8; Craigie et al. 2010, p. 2225); Botswana 2013b, pp. 87, 88; Lindsey and Bento protected areas has caused declining (Bauer et al. 2014, pp. 101, 103); CAR 2012, p. 64; Scholte 2011, p. 7). Hunting prey populations (Henschel et al. 2015, (Bouche´ et al. 2010, pp. 99, 1000; Roulet is more prevalent close to borders and unpaginated; Henschel et al. 2014, pp. 2004 in Bouche´ et al. 2010, p. 1002); near human settlements as the longer 5–6; Craigie et al. 2010, entire). Chad (Potgieter et al. 2009 in Bouche´ et the distance, the more time, effort, and The human population in the al. 2010, p. 1002); Coˆte d’Ivoire (Fischer cost is needed to find and transport developing world is projected to and Linsenmair 2001, p. 134); DRC meat; the chances of detection are also increase rapidly, suggesting human (Martin and Hillman-Smith 1999 in increased with distance (Lindsey et al. pressure on protected areas will also Bouche´ et al. 2010, pp. 1001–1002); 2013b, pp. 84, 88; Brashares et al. 2001, increase (Lindsey et al. 2013b, p. 84; Ghana (Brashares et al. 2004, p. 1182); p. 2475). Additionally, communities Brashares et al. 2001, p. 2475). Without Kenya (Western et al. 2009, pp. 2, 3, 4); often retain livestock as assets and rely intervention, wildlife resources will be Mozambique (Lindsey and Bento 2012, on bushmeat for daily protein needs lost in many areas with severe p. 63); Sudan (UNEP 2006 in Bouche´ et (Barnett 2000 in Lindsey et al. 2013b, p. ecological impacts (Lindsey et al. 2013b, al. 2010, p. 1001); Zambia (Simasiku et 88). Furthermore, many communities p. 84). Because lion densities closely al. 2008 in Lindsey et al. 2013b, p. 84); lack the rights over land and in most mirror prey densities, we can expect and Zimbabwe (Zimbabwe Parks and cases in Botswana, Tanzania, Zambia, that lion populations will also be lost in Authority 2015, p. and Zimbabwe, the government retains Africa. 9). Bouche´ et al. (2010, p. 1001) found a significant portion of revenue from Human-Lion Conflict that large wilderness areas spanning the wildlife; therefore, those that bear the The lion population in and around boundaries of Chad, CAR, DRC, and costs of wildlife do not receive benefits, the Gir Protected Area, India, lives Sudan suffered depleted wildlife and bushmeat hunting is the only way among and is surrounded by many abundance. Lindsey et al. (2013b, p. 84) to benefit from wildlife (Lindsey et al. pastoral and forest settlements (Banerjee concluded that the case studies 2013b, p. 88). represented only a tiny fraction of the and Jhala 2012, p. 1421; Singh and areas in savannas that are severely Throughout the African range Gibson 2011 in Banerjee and Jhala 2012, impacted by bushmeat hunting. Craigie countries, hunting of wildlife is p. 1421; Banerjee et al. 2010, p. 249; et al. (2010, p. 2226) stated their study regulated by various laws and Singh 2007 in Jhala et al. 2009, p. 3385). might underestimate the extent of regulations and harvests are controlled The lion population of Gir has increased decline that has occurred in Africa’s through permitting systems and quotas and dispersed into the large agro- protected areas because data came from (Lindsey et al. 2013b, pp. 82–83). In pastoral area adjacent to the protected sites with resources to carry out long- many countries, the use of snares, area. Only 10 percent of lions in India term monitoring programs and poison, and automatic weapons, among occur in the human-free portion of Gir increased management may be other methods, is prohibited. Single- National Park (Banerjee et al. 2013, p. associated with greater capacity to shot firearms, muzzle-loading firearms, 8). Conflict there, like in Africa, arises address threats. shot guns, and bows and arrows are from predation of livestock and Low lion population densities have legal under certain circumstances when associated threats to security of pastoral been found to correspond with low prey permitted, and in some cases specific livelihoods (Karanth and Chellam in densities (Van Orsdol et al. 1985, calibers and bow strengths are given Banerjee et al. 2013, p. 1). The lion’s Hayward et al. 2007 in Bauer et al. depending on the species being hunted diet there includes livestock (Banerjee et 2015a, unpaginated; Bauer et al. 2014, (Lindsey et al. 2013b, p. 82). Hunting al. 2013, p. 6; Meena et al. 2011, pp. 63– p. 103; Bauer et al. 2010, p. 363). laws also specify hunting seasons and 65). Between 2001 and 2010 the number Regional trends in lion populations, as prohibit hunting in certain protected of villages reporting depredation of discussed above, mirror regional trends areas, hunting certain species, and livestock increased (Meena et al. 2014, in populations in western, hunting young or pregnant animals. pp. 122–123). Additionally, Meena eastern, and southern Africa between Therefore, bushmeat hunting is illegal (2012, p. 36) found that in all Forest 1970 and 2005 (Bauer et al. 2015a, in most situations due to violations of Divisions, except Gir West, annual unpaginated; Henschel et al. 2015, one or more of these restrictions livestock predation increased more than unpaginated). Overall, Craigie et al. (Lindsey et al. 2013b, p. 83). However, 100 percent in 5 years. However, despite (2010, p. 2225) found a 59 percent penalties for violations are inadequate the lion’s close occupation with human decline in large mammal populations. and do not inhibit illegal bushmeat settlements and increased predation on Regional differences in herbivore hunting. Penalties typically comprise livestock, human-lion conflict and population abundance were also warnings, community service, or fines associated retaliatory killing was not detected. While population sizes in that are often lower than the value of the found to be a major source of lion southern Africa increased by 24 percent, meat, or the hunter is not penalized at mortality (Pathak et al. 2002 in Banerjee they declined by 52 percent and 85 all. Many governments lack the will and and Jhala 2012, p. 1427), mainly due to percent in eastern and western Africa, most state wildlife agencies lack the low economic losses via certain respectively (Craigie et al. 2010, p. resources or expertise to effectively husbandry practices and a 2225). enforce laws (Lindsey et al. 2013b, compensation scheme (Meena et al. Continent-wide decreases in prey p.88). Some government officials and 2014, pp. 123, 124; Banerjee et al. 2013, abundance in African protected areas police are known to purchase bushmeat, pp. 6–7, 8), cultural ethics (Raval 1991 are driven by human population growth despite it coming from an illegal source, in Banerjee et al. 2013, p. 2; Banerjee et (Craigie et al. 2010, p. 2225), especially which further contributes to ineffective al. 2013, p. 8), and strict legal along the boundaries of protected areas regulation of illegal hunting (Lindsey enforcement (Banerjee et al. 2013, p. 8).

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Although some lions have been killed Begg 2010, p. 2; Hazzah et al. 2009, p. Attacks on Livestock in Africa (Meena 2008 and Meena et al. 2007 in 2428; Moghari 2009, p. 36; Kissui 2008, The most significant cause of human- Meena 2010, p. 211), the lion p. 422; Hazzah 2006, pp. 15, 23, 25). lion conflict is livestock depredation. In population remained stable between Human-lion conflict stems from addition to bushmeat trade, the demand 2001 and 2010 (Meena et al. 2014, p. human population growth and the for food to meet increasing needs of a 123). resulting overlap of humans and growing population has been met by Although human-lion conflict is not wildlife habitat, with associated intensified agriculture and livestock currently considered a threat to the lion practices (Chardonnet et al. 2010, p. 19). population in India due to tolerance of livestock encroachment and decreasing availability of prey (Hoppe-Dominik et As natural habitats are converted to lion presence by the pastoralist agricultural or pastoral land, the lion’s community (Banerjee et al. 2013, pp. 1– al. 2011, p. 452; Chardonnet et al. 2010, pp. 6, 13–14; Frank et al. 2006, p. 12; natural prey base is further reduced 2, 8; Pathak et al. 2002 in Banerjee and (Chardonnet et al. 2010, p. 27; Jhala 2012, p. 1427), human-caused Hazzah 2006, pp. 14, 15). Lion Gebresenbet et al. 2009, p. 9). As a result mortality is likely to increase in the populations are increasingly restricted of prey species becoming depleted in future due to increased human-lion to protected areas due to human many areas, lions seek out livestock conflict and will be a major threat to the expansion and associated expansion of (and in some cases, humans) for food persistence of the lion population livestock husbandry and agricultural (Zimbabwe Parks and Wildlife (Banerjee and Jhala 2012, p. 1428). activities. Despite being within Management Authority 2015, p. 9; Similar to the observed transition in the protected areas, lions, due to their large Burkina Faso 2014, p. 20; Hoppe- Maasai community in eastern Africa, home range, often range beyond Dominik et al. 2011, p. 452; Chardonnet traditional value systems of pastoralists protected area borders where they are et al. 2010, pp. 6, 13–14, 27; in India are rapidly changing under the exposed to and impacted by people influence of globalization and free living on adjacent land. Therefore, most Gebresenbet et al. 2009, p. 9; Moghari 2009, pp. 78, 83; Frank et al. 2006, p. markets. The younger generation is conflict occurs at protected area 12; Hazzah 2006, pp. 17–18; Patterson et becoming less tolerant to even small boundaries (Henschel 2015, pers. al. 2004, pp. 507, 514). Therefore, lion monetary losses. These changes in comm.; Woodroffe and Ginsberg 1998, attacks occur at the highest frequency in attitudes will likely result in less p. 2126). It is along these borders that areas where natural prey abundance is tolerance of livestock loss to lions villages are often established and lowest (Packer et al. 2010, p. 9; Frank (Banerjee et al. 2013, p. 8). An indefinite human encroachment occurs due to et al. 2006, pp. 9, 12; Patterson et al. increase in humans and livestock within conversion of natural habitats for Gir Forests would upset the current 2004, p. 507). agriculture and grazing livestock, which Pastoralists allow increasing numbers balance by altering forest composition increases the chance of human-lion or population dynamics of prey species of livestock to graze in and adjacent to encounters (Sogbohossou et al. 2011, protected areas, and villagers farm up to and would be detrimental to pp. 51, 62; Chardonnet et al. 2010, p. 23; conservation (Banerjee et al. 2013, p. 8). the boundaries of protected areas, Mesochina et al. 2010a, p. 39; subjecting livestock and humans to Furthermore, with an expanding lion Mesochina et al. 2010b, p. 33; Moghari population that disperses and uses lions and increasing the risk of 2009, p. 14). Furthermore, cattle herders predation and the number of livestock habitat in agro-pastoral areas densely enter the protected areas, and lions populated with human villages, there is lost to predation (Brugie´re et al. 2015, p. move beyond the borders of protected 514; Bauer et al. 2014, p. 98; Burkina an increased potential for human-lion areas in search of food, increasing conflict (Meena 2010 and Singh 2007 in Faso 2014, pp. 19–22; Hazzah 2013, p. interactions between humans and lions 1; Chardonnet et al. 2010, pp. 11–12; Meena et al. 2014, pp. 120, 121). Due to and the risk of human-lion conflict high human density and demand for Uganda Wildlife Authority 2010, p. 27; (Burkina Faso 2014, pp. 19–20, 21; Moghari 2009, pp. 1, 90). Additionally, land, most human-free protected areas Hazzah et al. 2013, p. 1; Republic of in India, and elsewhere, are too small to poor husbandry practices and grazing of Namibia 2013, p. 13; Bauer et al. 2010, hold viable populations of large livestock within or adjacent to protected p. 365; Chardonnet et al. 2010, pp. 11– carnivores for the long term (Narain et areas increase exposure of livestock to 12; Mesochina et al. 2010a, p. 39; al. 2005 and Karanth 2003 in Banerjee lions and increase livestock loss Mesochina et al. 2010b, p. 33; Packer et et al. 2013, p. 8). (Uganda Wildlife Authority 2010, p. 27; Human-lion conflict and associated al. 2010, pp. 2, 6; Gebresenbet et al. Woodroffe and Frank 2005 in Moghari retaliatory killing of lions has played a 2009, p. 9; Moghari 2009, pp. 1, 14, 25, 2009, p. 35; Hazzah and Dolrenry 2007, major role in the reduction of lion 26, 78; Kissui 2008, p. 422; Hazzah pp. 22–23). Furthermore, conversion of populations throughout Africa (Lion 2006, p. 2). Hunting zones are thought rangeland to agricultural use has Guardians 2013, p. 1; Lion Guardians to serve as buffers; however, these areas blocked several migratory routes for 2011, p. 2; Hazzah and Dolrenry 2007, are not adequate as a low density of Tanzania’s wildebeest and p. 21; Frank et al. 2006, p. 1; Patterson competitors in these areas may attract populations, which likely forces lions to et al. 2004, p. 508) and is a threat to wildlife, including lions, which further rely more on livestock (Packer et al. remaining lion populations (Bauer et al. disperse into villages, causing conflicts 2010, p. 9). Because most protected 2010, p. 363; Hazzah et al. 2009, p. (Sogbohossou et al. 2011, p. 51). Lion areas are too small to support a lion’s 2428; Moghari 2009, p. 31; Kissui 2008, attacks can have various impacts on large home range, adjacent dispersal p. 422; Frank et al. 2006, pp. 1, 3, 10; those communities that coexist with areas are often used for supplementary Ray et al. 2005 in Hazzah 2006, p. 2; conflict-causing animals, generating food, putting them in greater contact IUCN 2006b, p. 18). Conflict between resentment towards them. When lions with livestock and humans (Kissui humans and wildlife has been linked to in Africa cause or are perceived to cause 2009, p. 422; Moghari 2009, p. 27). population declines, reduction in range, damage to livestock, property, or Conditions worsen as livestock numbers impacts to small population people, the response is generally to kill and area under cultivation increase, demographics, and even species them (Dickman 2013, pp. 378–379; leading to overgrazing, further habitat extinctions (Dickman 2013, p. 377; Moghari 2009, p. 25; Frank et al. 2006, destruction, and greater depredation Sogbohossou et al. 2011, p. 61; Begg and p. 1). rates (Gebresenbet et al. 2009, p. 9;

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Hazzah 2006, p. 61; Frank et al. 2005, to catch, and lions may turn to labor and, in some cases, the herd being Ntiati 2002, Mishra 1997, Meriggi and livestock. Migratory prey species such too large for the enclosures (Bauer et al. Lovari 1996, Rao 1996, Mech et al. 1988 as zebra and wildebeest will move to 2010, p. 365). in Hazzah 2006, p. 18). other areas for forage and replenished Attacks on Humans in Africa The use of fences to subdivide water sources, leaving lions to turn to rangeland interferes with traditional wet livestock as an alternate food source. Although lions generally avoid and dry season grazing schedules for Migratory prey may also move outside people, they will occasionally prey on livestock and wildlife (Hazzah 2006, pp. of protected areas. Opportunities for humans, causing serious injury or death 58–59). Restricting wildlife movement livestock predation on communal land (Dickman 2013, pp. 380, 384; reduces wild prey and, when combined increase when lions follow migratory Chardonnet et al. 2010, pp. 11, 12, 13; with an increase in livestock numbers, prey out of protected areas Moghari 2009, pp. 14, 49, 26, 88; Bauer increases the rate of human-lion conflict (Sogbohossou et al. 2011, p. 50; Packer et al. 2001 in Moghari 2009, pp. 31, 78, (Hazzah 2006, pp. 59, 61). Although et al. 2010, p. 9; Kissui 2008, p. 427; 84; Frank et al. 2006, p. 1; Hazzah 2006, well-built bomas (a livestock enclosure) Patterson et al. 2004, p. 514; Frank et al. pp. 14, 17; Patterson et al. 2004, p. 507). can effectively constrain cattle and keep 2006, p. 6). Attacks on humans appear to be more predators out (Frank et al. 2006, p. 8), Traditional livestock husbandry frequent in southern and eastern Africa they are traditionally built to keep practices are effective at reducing and rare in western and central Africa livestock confined, but do not offer depredation of livestock by lions (Bauer et al. 2010, p. 363; Chardonnet et effective protection from predators (Chardonnet et al. 2010, p. 35; Moghari al. 2010, pp. 12, 13; Mesochina et al. (Moghari 2009, p. 35). In the absence of 2009, p. 35; Frank et al. 2006, p. 2; 2010a, pp. 29–30; Frank et al. 2006, pp. reliable methods for protecting Hazzah 2006, p. 22). These practices 1, 10), although attacks on humans have livestock, some amount of depredation include livestock being closely herded been reported in Burkina Faso (Burkina can be expected, and some lions can by men and dogs during the day and Faso 2014, pp. 19, 22). Environmental become habitual livestock killers (Frank being brought into bomas at night with factors such as vegetative cover, habitat, et al. 2006, p. 9). people living in huts around them climate, seasonality, and prey Rates of livestock depredation vary (Frank et al. 2006, p. 4). However, availability may affect the rate of attacks with regional rainfall that correlate with traditional practices are being replaced on humans. A certain amount of prey availability, including changes in by less diligent husbandry practices, vegetative cover is crucial for lion’s herding strategies, movement of prey, which is increasing conflict (Woodroffe hunting success; however, in some and movement of lions (Lion Guardians and Frank 2005 in Moghari 2009, p. 35; cases, the vegetative cover may make it 2011, p. 6; Moghari 2009, p. 32; Hazzah Frank et al. 2006, pp. 2, 10; Hazzah and more difficult to catch prey, leading to 2006, pp. 17, 18; Patterson et al. 2004, Dolrenry 2007, p. 23). In Botswana, more attacks on humans. Additionally, p. 514). For example, in some parts of livestock are often left to wander dense cover near settlements allows Zimbabwe, Kenya, and Tanzania, outside bomas at night (Frank et al. lions to hide or stalk humans at a close livestock losses occur during the dry 2006, p. 5). In Kenya and Tanzania, distance (Mesochina et al. 2010a, p. 39; season. During this time, herders travel social changes are altering traditional Moghari 2009, p. 85; Frank et al. 2006, farther for forage and water, they use Maasai pastoral livelihoods, reducing p. 12). temporary bomas that are typically dependency on livestock, and reducing Provoked attacks on humans are weak, they are unfamiliar with traditional livestock care and usually associated with someone carnivore movements in these new management, leaving livestock more approaching a lion too closely or trying areas, and livestock are weak due to vulnerable to predation (Chardonnet et to injure or kill it and stealing a lion’s disease, which makes them more al. 2010, p. 35; Hazzah and Dolrenry prey for bushmeat (Chardonnet et al. vulnerable to predator attacks by lions 2007, pp. 22–23). Young Maasai boys 2010, p. 14; Uganda Wildlife Authority (Hazzah 2006, p. 17). Additionally, traditionally guarded herds at night; 2010, p. 27). Unprovoked attacks are herders are dependent on resources however, increased access to schools usually associated with old, sick, or within protected areas, and livestock has left herds unattended to wander into injured lions that turn to humans as may be left to wander for days or weeks predator areas at night (Chardonnet et easy prey. Additionally, there are risks during a prolonged drought to find al. 2010, p. 35). of unprovoked attacks associated with forage, increasing opportunities for In the Pendjari area of Benin, certain human activities. These attacks on livestock by lions traditional enclosures are low with few activities include walking alone at (Sogbohossou et al. 2011, p. 44; branches. These structures and the lack night, sleeping outside, and surprising a Chardonnet et al. 2010, p. 24; Frank et of enclosures encourage livestock lion, particularly if it has cubs (Begg and al. 2006, p. 6). In Benin, other parts of predation (Butler 2000, Mazzolli et al. Begg 2010, pp. 3, 21; Chardonnet et al. Kenya, the Maasai Steppe region of 2002, and Wang and Macdonald 2006 in 2010, pp. 14, 15; Mesochina et al. Tanzania, and Queen Elizabeth National Sogbohossou et al. 2011, p. 51). 2010a, pp. 38, 39; Mesochina et al. Park, Uganda, livestock losses were Surveillance of a main pasture area 2010b, p. 32; Uganda Wildlife Authority greater during or following the rainy south of Waza National Park in 2010, p. 27; Moghari 2009, p. 85; Frank season (Sogbohossou et al. 2011, p. 49; Cameroon and improved enclosures et al. 2006, pp. 11, 12). The most Moghari 2009, p. 88; Kissui 2008, pp. around Waza National Park and common context for attacks on humans 427, 428; Frank et al. 2006, p. 6; Pendjari National Park, Benin, led to a occurs during harvest, due to prey Patterson et al. 2004, pp. 510, 514). significant decrease in depredation dispersal during the wet season, bush Weakened prey and readily available (Bauer et al. 2010, p. 365). However, pig attraction to crops, and because carcasses provide easy meals during people do not invest much into humans are particularly vulnerable in times of drought, and wild herbivores improving enclosures even though they makeshift tents while protecting crops tend to concentrate near available water appear to be economically efficient, (Frank et al. 2006, p. 12). sources, making them easier to prey on ecologically effective, and culturally and leading to fewer livestock attacks. acceptable. Even enclosures that were Retaliatory Killing of Lions in Africa However, when rains return, the built as part of a conservation project Livestock provide an economic value abundant grass makes wild prey harder were not used full time due to lack of to humans, particularly those in extreme

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poverty who rely solely on livestock for unpaginated; Tanzania 2015, p. 13; negative impacts on lion populations their protein source and livelihood. Republic of Namibia 2013, pp. 12, 13– (Frank et al. 2006, p. 9). When lions have no economic value to 14; Begg and Begg 2010, p. 15; Studies have shown that lion local communities and they kill or are Chardonnet et al. 2010, pp. 41–42; populations are declining in areas perceived to kill livestock, the economic Packer et al. 2010, pp. 9–10; Uganda where pastoralism persists and the impact can be significant (Bauer et al. Wildlife Authority 2010, pp. 13, 42; presence of mobile pastoralists are a 2015a, unpaginated; Hazzah et al. 2014, Gebrensenbet et al. 2009, p. 7; Hazzah good indicator of lion extinction p. 852; Chardonnet et al. 2010, p. 12; et al. 2009, p. 2429; Moghari 2009, pp. (Brugie´re et al. 2015, p. 519; Hazzah et Mesochina et al. 2010a, p. 38; 52, 89, 91; Ikanda 2008, pp. 5–6; Hazzah al. 2009, p. 2428). Within protected Mesochina et al. 2010b, p. 33; and Dolrenry 2007, p. 21; Frank et al. areas, human–wildlife conflict is likely Gebresenbet et al. 2009, p. 9; Moghari 2006, pp. 2–4, 7; Hazzah 2006, p. 52; under-reported because cattle herders 2009, pp. 4, 25, 49; Kissui 2008, pp. 423, IUCN 2006b, p. 15). Retaliatory killings are within the protected areas illegally 429; Hazzah 2006, p. 24; IUCN 2006a, have been reported as a significant and, therefore, unlikely to report it (Chardonnet et al. 2010, p. 14; pp. 23, 24; IUCN 2006b. pp. 18–19; threat to lion populations in protected Mesochina et al. 2010b, p. 34). For Frank et al. 2006, p. 3). Subsequently, areas of western and central Africa example, Etosha National Park and those lions that reside on the edge and (Tumenta et al. 2009 and Henschel et al. Caprivi Game Park have the highest outside of protected areas, where there 2010 in Sogbohossou et al. 2011, p. rates of lions killed per 100 km2, yet it is an increased risk of exposure to 100), Botswana (Bauer et al. 2014, pp. humans and livestock, are subject to may be that just under half of the lions 98, 103), Botswana and South Africa that are killed are reported (Republic of retaliatory killing across Africa. (Kgaladi Transfrontier Park; Funston Boundary transgression leads to lions Namibia 2013, p. 14). Although we do 2011, p. 1), Cameroon (Delongh et al. not have information on human–lion predating on livestock, and in turn, be 2009 and Tumenta et al. 2010 in subject to pre-emptive or retaliatory conflict from all lion range countries, it Sogbohossou et al. 2011, p. 60), Kenya is reasonable to conclude that lions are killing (Bauer et al. 2014, pp. 98, 103; (Patterson et al. 2004, Kolowski and Funston 2011, pp. 1, 3, 5, 6–7); being killed as a result of conflict in all Holekamp 2006, and Hazzah et al. 2009 however, this type of killing of lions major African range countries, due to in Sogbohossou et al. 2011, p. 60), also occurs within protected areas their depredation on livestock (Frank et Tanzania (Tanzania 2015, p. 13; Kissui (Henschel et al. 2015, unpaginated; al. 2006, p. 4). 2008 in Sogbohossou et al. 2011, p. 60), Zimbabwe Parks and Wildlife and Zimbabwe (Zimbabwe Parks and Factors That Drive Retaliation in Africa Management Authority 2015, p. 10; Wildlife Management Authority 2015, p. Several anthropogenic factors drive Burkina Faso 2014, pp. 19, 21, 22; 10). the level of resentment towards lions Tumenta et al. 2009 and Henschel et al. In areas of high conflict, identifying and the extent of retaliatory killing 2010 in Sogbohossou et al. 2011, p. 100; (Dickman 2013, pp. 379, 385), including Moghari 2009, p. 49). Furthermore, the responsible animal is often difficult, and a token animal may be killed the extent of the loss caused by the lions killing of lions outside of protected and the wealth and security of the instead (Hazzah 2006, p. 25), leaving the areas may disrupt movement of lions to people affected (Dickman 2013, p. 381; problem lion to continue to attack and other areas that could contribute to the Mesochina et al. 2010b, p. 54; Moghari the potential for additional retaliatory viability of larger resident populations 2009, pp. 14, 25; Hazzah 2006, p. 81). killings. In Tanzania, game officers kill (White 2015, pers. comm.). This Depending on alternative assets or numerous lions each year in retaliation occurrence greatly impacts already- incomes, the economic impact of lions dwindling lion populations. Even if for attacks (Frank et al. 2006, p. 12). killing livestock can be significant. mortality occurs outside of protected Whereas shooting or spearing target Domestic livestock can provide manure, areas, population dynamics inside specific problem animals, poisoning is milk, and meat, and are the basis of protected areas are negatively impacted. indiscriminate and is known to remove many family incomes, savings, and When lions outside of protected areas entire prides at once (Frank et al. 2006, social standing; losses can amount to a are removed, either through retaliatory pp. 2, 10, Living with Lions no date, large proportion of a subsistence killings or trophy hunting, territorial unpaginated). In the absence of reliable herder’s annual income. These losses gaps that are left are filled by lions from methods for protecting livestock, rural are generally uncompensated, closer to the core of the protected area, people often turn to indiscriminant reinforcing negative community exposing more lions to human–lion methods, like poisoning, to control attitudes toward lions and causing conflict along the borders of the livestock depredation. Poisoning is an retaliation (Dickman 2013, pp. 380, 381; protected area and creating a population easy method for lethal control since it Chardonnet et al. 2010, pp. 11, 12, 18, sink (Brugie´re et al. 2015, p. 514; is readily available, and reinforcing 29; Hazzah et al. 2009, p. 2428; Moghari Sogbohossou 2014, p. 3; Loveridge et al. bomas or more carefully tending 2009, pp. 14, 25, 27, 36; Kissui 2008, pp. 2007, pp. 552, 555; Woodroffe and livestock requires time and effort. The 422–423). Furthermore, a common Ginsberg 1998, p. 2162). use of Furadan, a widely available and perception among local communities is The availability of guns and poison cheap agricultural pesticide, is that lions are conserved at the cost of makes killing suspected predators particularly lethal to wildlife and is community safety and uncompensated cheaper and easier than other control increasingly being used to kill predators financial losses. When the people who methods, such as reinforcing bomas in small pastoralist areas of Kenya and suffer significant costs from wildlife feel (Hazzah et al. 2009, p. 2429; Moghari Tanzania. Livestock carcasses are that the wildlife’s needs are being put 2009, p. 35; Frank et al. 2006, p. 14; doused with the poison, killing before their own needs, their frustration Hazzah 2006, p. 3). Spearing, shooting, predators and scavengers that feed on can lead to retaliatory killings (Dickman trapping, and poisoning of lions, as them (Frank et al. 2006, pp. 2, 10, Living 2013, p. 382). Additionally, government either a preventive measure or in with Lions no date, unpaginated). officials and local tour and hunting retaliation for livestock and human Poisoning of bush pig carcasses to kill operators experience economic gain attacks, occurs regularly (Brugie´re et al. lions is not uncommon after attacks on from lions, whereas the communities 2015, p. 519; Bauer et al. 2015a, humans. These practices have serious bear the costs in livestock losses

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(Hazzah et al. 2014, p. 852). This Hazzah 2006, p. 99). In some areas of et al. 2011, entire; Packer et al. 2011, situation further contributes to negative Africa, locals believe in ‘‘spirit lions,’’ a entire; Loveridge et al. 2007, entire). attitudes toward lion conservation lion whose body is overtaken by evil to Depending on how trophy hunting is programs (Moghari 2009, p. 37). kill rivals or their livestock (West 2001 regulated and managed, trophy hunting Lions are particularly vulnerable to in Dickman 2013, pp. 381–382). Because can be a tool for conservation, but may retributive killing because they are often people believe spirit lions are created by also have negative impacts on lions driven by a perceived level of lion their enemies, the number of perceived (Bauer et al. 2015a, unpaginated; predation on livestock rather than actual spirit lions, and killing of these lions, Lindsey et al. 2013a, p. 1; Whitman et levels of conflict. In some locations, increases during times of social tension al. 2004, pp. 176–177; Loveridge et al. other predators (e.g., baboons (Papio (Dickman 2013, p. 382.) 2007, p. 548). ursinus), spotted hyenas (Crocuta Cultural beliefs can also have a In response to growing international crocuta), and (Panthera positive impact on lions. An association recognition of reduced population pardus)) as well as disease are with a totem is an important component numbers, many countries began responsible for the majority of livestock of certain cultures and could explain implementing moratoriums banning the losses and human casualties, yet it is why retaliatory killing is uncommon in sport hunting of lions. In this document lions that are sought and killed more some areas despite negative perceptions. we use the terms moratorium and ban often. In the Pendjari Biosphere Reserve, However, the positive impact may not interchangeably. A ban or moratorium Sogbohossou et al. (2011, p. 74) found continue as cultural beliefs dwindle due can be permanent, long term, or that just one case of a nonlethal attack to urbanization and modernization temporary, and can occur in countries on a human in a decade and mere (Sogbohossou et al. 2011, pp. 73, 75). that have hunting quotas in place (e.g., rumors of attacks in other regions was Social tensions within tribes and Botswana and Zambia). Having both a enough to cause people to perceive lions between local communities and other moratorium and a quota in place at the as a threat. Negative perceptions of lions communities, the government, park same time means that, although the may be based on an over-estimated officials, or tourists can lead to conflict country may have a hunting quota, the number of lions in a community or and retributive killing of lions (Dickman country has halted authorization of protected area and an over-estimated 2013, p. 382; Hazzah 2006, p. 75). trophy hunting pursuant to that quota number of human–lion conflicts Locals often report that wildlife until some later date or until some (Dickman 2013, p. 380; Begg and Begg authorities do not react effectively when further action is taken, as prescribed by 2010, p. 20; Chardonnet et al. 2010, pp. chronic livestock raiders are reported that country. 12, 21–22; Hazzah et al. 2009, p. 2436; (Frank et al. 2006, p. 9). Significant Trophy hunting is currently banned Maclennan et al. 2009 in Hazzah et al. numbers of lions have been killed when in 12 range countries: Angola, 2009, p. 2429; Moghari 2009, pp. 77–78, promised benefits were not received or Botswana, Congo, Gabon, Ghana, India, 107, 150; Holmern et al. 2007 in adequate compensation was not Kenya, Malawi, Mauritania, Niger, Moghari 2009, p. 34; Butler 2001 in provided for livestock and human losses Nigeria, and Rwanda (CITES 2014, p. Moghari 2009, p. 34; Kissui 2008, pp. (Dickman 2013, p. 383; Hazzah 2006, p. 14; Meena 2014, p. 26; Lindsey et al. 426, 428, 429; Hazzah 2006, pp. 18–19, 45). 2013a, entire; Lindsey 2013, pers. comm.; Jackson 2013, pp. 7–8). In 1977, 83–85, 96, 98, 107, 111; Patterson et al. Trophy Hunting 2004, pp. 514, 515). One cause for the Kenya banned all sport hunting (Elliot disproportionate blame put on lions is Lions are a key species in sport and Mwangi 1998, p. 3). Botswana that the lion is a highly visible species. hunting, or trophy hunting, as they are banned between 2001 and It is a large-bodied species that lives in considered one of the ‘‘big five’’ African 2004, and then again from 2008 to the groups and has cultural significance. species (lion, , , rhino, present (Davison et al. 2011, p. 114). Because of its physical presence, there and cape buffalo) touted to be the most Benin imposed a 2-year moratorium, is often a hyper-awareness of the challenging to hunt due to their and CAR a 3-year moratorium, in the potential risk for lion attacks and lions nimbleness, speed, and behavioral early 2000s (Lindsey et al. 2013a, p. 4). may be blamed simply because they unpredictability (Lindsey et al. 2012a, In January of 2013, Zambia placed a have been seen in an area (Dickman p. 2). However, with the documented moratorium on sport hunting in 19 game 2013, pp. 380–381). decline in lion population numbers management areas. While a few other Cultural beliefs and traditions can throughout Africa, sport hunting of game management areas and private have a negative impact on lions. lions for trophies has become a highly game ranches in Zambia remain open Because cattle are of great cultural complex issue. for sport hunting for other species, the significance to Maasai, their loss can Trophy hunting is carried out in a nationwide moratorium on sport impose social or cultural costs and number of range countries and is hunting of cats remains in place (White incite greater resentment and higher considered an important management 2015, pers. comm.; ABC News 2014, levels of retributive killing (Dickman tool for conserving land and providing unpaginated; Flocken 2013, 2013, p. 384; Kissui 2008, p. 429; financial resources for lion unpaginated). Trophy hunting is Hazzah 2006, p. 99). Cultural beliefs conservation. However, management restricted to problem or dangerous still motivate ritual lion hunts for young programs are not always sufficient to animals in Ethiopia and Uganda Maasai warriors. Despite being deter unsustainable off takes (harvests), (Lindsey 2008, p. 42). In our proposed outlawed, this practice persists due to which has occurred in many areas rule, we had conflicting information community secrecy. However, it is (Lindsey et al. 2013a, pp. 8–9; Packer et regarding whether Cameroon had or has easily disguised as retaliatory killings al. 2006 in Bauer et al. 2015a, a lion hunting moratorium (CITES 2014, for livestock predation. The prohibition unpaginated). Documented declines in p. 14; Lindsey 2013, pers. comm.; of ritual lion hunts provides a greater lion populations of Africa are a result, Jackson 2013, p. 8). During the public incentive for participating in retaliatory in part, of mismanaged trophy hunting comment period, a peer reviewer hunts (Hazzah et al. 2014, p. 852; Packer (Rosenblatt et al. 2014, entire; confirmed that Cameroon has not put a et al. 2010, p. 10; Moghari 2009, pp. 13– Sogbohossou et al. 2014, entire; Becker moratorium in place for lions, either in 14, 28; Ikanda 2008, pp. 5, 6; Kissui et al. 2013, entire; Lindsey et al. 2013a, the past or present (Bauer 2015, pers. 2008, p. 423; Frank et al. 2006, p. 10; entire; Packer et al. 2013, p. 636; Croes comm.). Additionally, Zimbabwe has

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suspended trophy hunting in the being allocated to locations with only country with a lion quota less than Gonarezhou area (Conservation Force reportedly higher levels of human–lion the recommended 1 lion per 2,000 km2. 2015, pers. comm.). conflict (Lindsey et al. 2013b, p. 4). It should be noted that although quotas As of May 2014, approximately 18 Generally, the conservation principle are currently set higher than countries in Africa allowed legal behind scientifically based quotas is to recommended, the actual offtake for hunting of lions for trophies: Benin, limit total offtake of the species to either each of the countries overall has been Burkina Faso, CAR, DRC, Ethiopia, Coˆte equal or slightly lower than the growth consistently lower than the set quota d’Ivoire, Mali, Mozambique, Namibia, rate of the target specimens (e.g., males (Table 5). However, in Burkina Faso, Senegal, Somalia, RSA, Sudan, vs. female), such that damage to the Zambia, Namibia, and Zimbabwe, the Tanzania, Togo, Uganda, Zambia integrity and sustainability of that actual harvests are greater than Packer’s (nationwide moratorium on sport population is prevented. Scientifically recommended offtake (Lindsey et al. hunting of cats is currently in place), based quotas do not apply solely to 2013a, p. 8). For instance, five countries and Zimbabwe. However, in 2013 lion sport hunting, but set the limits for total maintain quotas to allow for 5–31 lion trophy hunting was documented to offtake for a particular timeframe; other trophies to be taken per year: Benin (5), occur in only 8 countries, specifically potential offtake includes problem- Burkina Faso (20), Cameroon (30), CAR Benin, Burkina Faso, CAR, animal control (to reduce human– (31), and Namibia (15). Only Mozambique, Namibia, South Africa wildlife conflict), translocation (to Mozambique currently has a quota (RSA), Tanzania, and Zimbabwe expand conservation), (reducing lower than the recommendation of (Lindsey 2013, pers. comm.). Four population pressures), and local hunting Packer et al. (2001, p. 1651). In 2013, the (for protein/meat or employment) (WWF countries, Burundi, Guinea Bissau, quota was set at 42–60 lions, which Lesotho, and Swaziland, provide no 1997, pp. 8–10). For quotas to be translates to 1 lion per 2,400km2 (or 0.8 legal protection for lions (CITES 2014, sustainable, scientists and policy lions per 2,000km2). Between 2011 and p. 14). makers must evaluate a multitude of 2012, Tanzania maintained the highest Where trophy hunting occurs, quotas factors including the species’ biological quota for lions at 315 (Lindsey et al. are set by the government for the factors (i.e., reproductive rate, gender 2013a, p. 6). purpose of limiting the actual number of ratios, age, and behavior), as well as lions killed (offtake) during a given community and client objectives (WWF Several countries have begun to timeframe. A scientifically based quota 1997, pp. 14–19). reduce their quotas as they have begun is the maximum number of a given Creel and Creel (1997, p. 83, executive implementing recommendations as species that can be removed from a summary) suggest that, for a quota to be outlined by Lindsey et al. (2013a, pp. 8– specific population without damaging considered sustainable for lions, it 9), Hunter et al. (2013, unpaginated), the biological integrity and should be limited to no more than 5 and Packer et al. (2011, p. 151) (Bauer sustainability of that population (World percent of the population. Distinct from 2015, pers. comm.; Henschel 2015, pers. Wildlife Fund (WWF) 1997, p. 9). Two the quota, Packer et al. (2011, p. 151) comm.; White 2015, pers. comm.; primary concerns have been raised by recommend actual lion offtake should Tanzania 2015, pers. comm. Zimbabwe the scientific and international not exceed more than 1 lion per 2,000 2015, pers. comm.). In 2011, community with regard to current lion km2 (Bauer 2015, pers. comm.; Henschel Zimbabwe’s quota was set at 101 lions; quotas. These are that (1) existing quotas 2015, pers. comm.; Packer et al. 2015, in 2014, it was reduced to 50 lions are set above sustainable levels, and (2) per comm.; Creel and Creel 1997, p. 83, following the implementation of age the data used for setting quotas is executive summary). However, most restrictions (Henschel 2015, pers. inconsistent and not scientifically based range countries have their quotas set comm.). Following pressure from the (Hunter et al. 2013, unpaginated; well above these recommendations European Union to ban lion trophies if Lindsey et al. 2006, p. 284) (see (Bauer 2015, pers. comm.; Henschel their quota remained higher than the 1 Potential Impacts of Trophy Hunting). 2015, pers. comm.; Packer 2015, pers. lion per 2,000 km2 recommendation, For example, recent quotas do not comm.). Specifically, Lindsey et al. Burkina Faso proposed to reduce the set appear to address safeguards for (2013a, p. 8) found that of the nine quota of 20 lions in the 2014/2015 sustainability nor has a systematic countries allowing trophy hunting of season to 6 in the 2015/2016 season approach been established for setting lions in 2013 (including data from (Henschel 2015, pers. comm.). South lion quotas (Hunter et al. 2013, p. 2; Zambia prior to the moratorium in Africa has not set a quota for the take Lindsey et al. 2013b, p. 8). Additionally, 2013), eight have quotas set higher than of wild lions since 99 percent of the it has been noted that previous quotas current recommendations by Packer et trophy-hunted lions are reportedly not in Namibia, Mozambique, and al. (2011, p. 151) and five have quotas of wild origin but captive born (Hunter Zimbabwe may have been influenced by set to more than double Packer’s et al. 2013, p. 2; RSA 2013, pp. 5, 7) human–lion conflict, with higher quotas recommendations. Mozambique is the (Table 5).

TABLE 5—ANNUAL TROPHY QUOTAS AND OFFTAKE BY COUNTRY (APPROXIMATE) AS OF 2013*

Annual lion Annual Country trophy quotas Year(s) of data offtakes Year(s) of data

Panthera leo leo Benin ...... 5.0±0 2007–2009 2.0±0.4 2007–2009 Burkina Faso ...... 20.0±0 2006–2009 13.3±1.45 2006–2009 Cameroon ...... 29.2±2 2006–2010 6.9±1.0 2006–2010 CAR ...... 31 2009 13.7±6.9 2008–2011 Panthera leo melanochaita ...... Mozambique ...... 42–60 2013 19.2±7.3 2008–2011 Namibia ...... 14.5 2010 14.0±3.2 2008–2011 Tanzania ...... 315 2011–2012 85 2011–2012 Zambia (moratorium) 1 ...... 74(50 2) 2012 47 2012

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TABLE 5—ANNUAL TROPHY QUOTAS AND OFFTAKE BY COUNTRY (APPROXIMATE) AS OF 2013*—Continued

Annual lion Annual Country trophy quotas Year(s) of data offtakes Year(s) of data

Zimbabwe ...... 101(503) 2011 42.5±7.5 2008–2011 * Source: Lindsey et al. 2013a. p.6. 1 Zambia enacted a moratorium on sport hunting in 19 game management units. Sport hunting remained open in other game management units and on some private game ranches. Sport hunting of all cats is currently banned throughout Zambia (White 2015, pers. comm.). 2 Approximate average quota for Zambia in the few years prior to the moratorium placed on cat hunting in 2013. (White 2015, pers. comm.). 3 In 2014, Zimbabwe reduced its quota to 50 due to implementation of age restrictions (Henschel 2015, pers. comm., citing Lindsey pers. comm.)

Potential Benefits of Trophy Hunting aside a combined 420,000 km2 of capita lower than the poverty level communal land, 188,000 km2 of (Barnett and Patterson 2005, p. iii). Proponents and most lion experts 2 support trophy hunting as a commercial land, and 420,089 km of These combined challenges highlight 2 the need for innovative solutions. conservation tool for the lion if it is state land totaling over 1,028,000 km for sport hunting purposes (Barnett and Conservationists and range countries practiced in a sustainable and Patterson 2005, p. iii). recognize the value of the wildlife scientifically based manner (Henschel As a species with a considerable range sector; if managed sustainably, there is 2015, pers. comm.; Hunter 2011, entire; (up to 1,000 km2) (Packer et al. 2013, p. potential to contribute to rural economic van der Merwe 2013, entire; Hunter et 636; Haas et al. 2005, p. 4), suitable development while simultaneously al. 2013, entire) because it can provide: habitat is important to the survival of protecting the unique ecological habitats (1) Incentives for the conservation of the species, and the marked decline in and species contained therein large tracts of prime habitat, and (2) suitable habitat is a significant threat to (Chardonnet et al. 2010, p. 33; Kiss funding for park and reserve the species (see Habitat Loss). The land 1990, pp. 1, 5–15). management, anti-poaching activities, currently designated in Africa for use in For species such as the lion to persist, and security activities. sport hunting has helped to reduce, but local communities must benefit from or As habitat loss has been identified as not eliminate, the impact of habitat loss receive a percentage of funds generated one of the primary threats to lion on the lion. from tourism such as wildlife viewing, populations, it is notable that the total If trophy hunting is part of a photography, or trophy hunting (White amount of land set aside for hunting scientifically based management 2013, p. 21; Martin 2012, p. 57; Kiss throughout Africa, although not program, it may provide direct [editor] 1990, pp. 1, 5–15). The ameliorating threats to habitat loss, economic benefits to the local economic value of a species, such as exceeds the total area of the national communities and may potentially create lion, can encourage range countries to parks, accounting for approximately half incentives for local communities to develop management and conservation of the amount of viable habitat currently conserve lions, reduce the pressure on programs that involve local available to lions (Chardonnet et al. lion habitat, and reduce retaliatory communities and which would 2010, p. 34; Packer et al. 2006, pp. 9– killing, primarily because lions are ultimately discourage indiscriminate 10). For example, in Tanzania, 25–33 viewed as having value. Conversely, killings by local communities (Groom percent of the total area, covering over lack of incentives could cause declines 2013, pp. 3, 5; Hazzah et al. 2013, p. 1; 247,000 km2 and encompassing 190 in lion populations because lions are White 2013, p. 21; Martin 2012, p. 49). hunting units, has been set aside for viewed as lacking value and they kill If local communities see no benefit of sport hunting purposes; this has livestock, which are of great value to lions being present in their communal resulted in an area 5.1 times greater than communities (see Human–lion Conflict). areas, sustainable use of lions becomes Tanzania’s fully protected and gazetted Over the last few decades, less competitive with other land-use parks (Jackson 2013, p. 6; Barnett and conservationists and range countries options, such as grazing and livestock Patterson 2005, p. 61). Tanzania also has have realized the integral role local management, and local communities land set aside for sport hunting in the communities play in the conservation of become unwilling and unable to manage form of safari areas, communal land, lions and their habitat; when their wildlife heritage (Barnett and and privately owned properties that communities benefit from a species, Patterson 2005, p. iii). When the value make up 23.9 percent of the total land they have incentive to protect it. of lions in areas outside national parks base (Barnett and Patterson 2005, pp. Therefore, using wildlife as a source of is diminished, those areas are likely to 76–77). income for rural populations has be converted to forms of land use less In Botswana, despite the current ban increasingly been employed throughout suitable for lions, such as agriculture, on lion hunting, the country currently the lion’s range countries in Africa. livestock pastures, or areas of resource has over 128,000 km2 of gazetted Many of these countries are classified as extraction, making lions even more wildlife management areas and ‘‘developing’’ nations; specifically, vulnerable to expanding human controlled hunting areas set aside for seven of the ten countries (we include settlement (Van der Merwe 2013, p. 2). hunting purposes, which equates to 22.1 Cameroon here) where trophy hunting is Community conservancies that benefit percent of the country’s total area. This permitted have 27–64 percent of their from trophy hunting have specifically amount is in addition to 111,000 km2 human populations living in severe been formed as a way to protect wildlife (or 19.1 percent of the country’s total poverty (United Nations Development and habitat. As an example, in Namibia, area) set aside as habitat in the form of Programme’s (UNDP) 2014, 160,000 km2 of community national parks, game reserves, and forest unpaginated; Barnett and Patterson conservancies were established in part reserves (Barnett and Patterson 2005, p. 2005, p. iii). These countries often have due to revenue from trophy hunting. 7). In 2000, five countries in southern high population growth, high These conservancies benefit the local Africa (Botswana, Namibia, South unemployment, limited industry, and a communities, which in turn protect lion Africa, Tanzania, and Zimbabwe) set Gross Domestic Product (GDP) per habitat. In 2012, the Save´ Valley

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Conservancy (Zimbabwe) ‘‘provided see Potential Impacts of Trophy currently, efforts are being made in over $100,000 USD worth of support to Hunting). many range countries to incorporate adjacent villages or farmers in the Many range countries rely heavily on incentives at the local level (Barnett and resettled areas. Assistance included tourism (predominantly and Patterson 2005, p. vi). drilling boreholes, maintaining safari hunting) to provide funding for In summary, if part of a scientifically boreholes, dredging of dams, building wildlife management (IUCN 2006a, p. based management program (including a clinics and schools, assisting with 24). Additionally, revenue generated scientifically based quota), trophy repairs, maintenance and materials for from these industries provides jobs, hunting of lions can provide direct schools, education initiatives, school such as game guards, cooks, drivers, and benefits to the species and its habitat, field trips, provision of computer security personnel and often brings in both at the national and local levels. equipment in schools, and craft revenue for local microbusinesses that Trophy hunting and the revenue programs’’ (Groom 2013, p. 5). sell art, jewelry, and other crafts. generated from trophy hunting are tools Connecting conservation to community Revenue generated from scientifically that range countries can use to facilitate benefits can provide a value for wildlife, based management programs can be maintaining habitat to sustain large including lions, where there was used to build and maintain fences, ungulates and other lion prey, previously resentment or indifference, provide security personnel with protecting habitat for lions, supporting helping to instill a sense of importance weapons and vehicles, provide the management of lion habitat, and for lion conservation. Additionally an resources for anti-poaching activities, protecting both lions and their prey base estimated 125,000 kg of game meat is and provides resources for habitat through anti-poaching efforts. While provided annually to rural communities acquisition and management scientifically based trophy hunting by trophy hunters in Zambia at an (Chardonnet et al. 2010, pp. 33–34; alone will not address all of the issues estimated value of $250,000 USD per Newmark 2008, p. 321). For example, that are contributing to the declined year, which is considerable for rural trophy hunting revenue in the Save´ status of the lion, it can provide benefits locations where severe poverty and Valley Conservancy in Zimbabwe has to the species. malnutrition exists (White 2013, p. 21), enabled $150,000–$250,000 USD to be Potential Impacts of Trophy Hunting further providing a value for wildlife, invested in anti-poaching activities, including lions. As stated above, local including the removal of wire-snares An issue critical to the conservation communities can benefit from the (Groom 2013, p. 5). Revenue from of lions is sustainable management of trophy hunting industry by additional trophy hunting can also increase the trophy hunting by lion range countries. employment opportunities and revenue ability of many African countries to Lion experts agree that, if trophy generated for local microbusinesses. manage wildlife populations both hunting is well regulated and managed, within and adjacent to reserves; many of Many range countries have recognized it can be a tool for conservation (Bauer these hunting areas are geographically et al. 2015a, unpaginated; Lindsey et al. the need to incorporate incentives and linked to national parks and reserves, local community benefits into their 2013a, p. 1; Whitman et al. 2004, pp. providing wildlife corridors and buffer 176–177; Loveridge et al. 2007, p. 548). trophy hunting regulations, land zones (Chardonnet et al. 2010, p. 34; management policies, and lion However, problems with the current Newmark 2008, p. 321). management of lion hunting increase conservation action plans (Lindsey et al. Depending on the country in which a 2013a, pp. 2–3; Zambia Wildlife the likelihood of negative impacts on hunter visits, there may be several the species (note that because 99 Authority 2009, p. 10; Windhoek 2008, different fees associated with trophy p. 18; IUCN 2006a, pp. 22, 24; IUCN percent of hunted lions in South Africa hunts, including game fees, observer are captive-bred, we exclude them from 2006b, pp. 23, 28; Zimbabwe Parks and fees, conservation fees, permit fees, Wildlife Management Authority 2006, this discussion) (Hunter et al. 2013, p. trophy handling fees, and government 2). Lindsey et al. (2013a, pp. 8–9) and unpaginated). Of the ten countries payments in terms of taxes, as well as where lion trophy hunting currently Hunter et al. (2013, p. 2) identified six safari operator fees (Barnett and key practices undermining sustainable occurs (we are including Cameroon and Patterson 2005, p. 71). In the late 1990s, South Africa here), seven have management of lions: Tanzania reported annual revenue of • developed National Poverty Reduction $29.9 million USD from all trophy Arbitrary establishment of quotas and Strategies in partnership with the hunting, South Africa $28.4 million excessive harvest International Monetary Fund (for a • lack of age-restriction implementation USD, Zimbabwe $23.9 million USD, • complete list, see http://www.imf.org/ Botswana $12.6 million USD, and fixed quotas external/np/prsp/prsp.aspx). Each of • hunting of females Namibia $11.5 million USD; the revenue • these countries has incorporated generated solely from lion hunting was lack of minimum hunt lengths in sustainable natural resource some countries not broken out (Barnett and Patterson • development as a priority and discussed 2005, p. iv). According to Groom (2013, general problems associated with benefit distribution and management to p. 4), a 21-day lion hunt in Save´ Valley management of trophy hunting rural communities (Benin 2000, Conservancy, Zimbabwe, may be sold As discussed above, one of the unpaginated; Burkina Faso 2000, for approximately $2,500 USD per day, primary practices experts identify as unpaginated; CAR 2000, p. 45; United with an additional trophy fee of undermining sustainable trophy hunting Republic of Tanzania 2000, pp. 13, 21; approximately $10,000 USD. Between is the use of non-scientific information Zambia 2000, unpaginated). Although 2005 and 2011, lion hunting in Save´ underlying the development of quotas we acknowledge the steps many Valley Conservancy provided an (Lindsey et al. 2013a, p. 8). The best countries have taken to address local estimated net income (based on 26 available monitoring data should be community incentives, most of the lions) of approximately $1,365,000 USD used to set quotas if they are to be countries are currently not transparent in per-night charges and roughly scientifically based and sustainable. about the benefits provided to local $260,000 USD in trophy fees (Groom However, monitoring data are often communities, and due to the high 2013, p. 4). In the past, government and lacking (Barnett and Patterson 2005, p. revenue potential, are subject to private landowners were the primary 102). A limited number of independent, corruption (Packer 2015, pers. comm.; beneficiaries of the revenue gained; scientific population counts of lions

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have occurred across their range, levels to ensure sustainability while for each client and submitted to especially in hunting concessions other methods are being developed and different government departments’’ (LionAid 2014a, pers. comm.; Packer refined. According to Hunter et al. (Barnett and Patterson 2005, p. 100). 2015, pers. comm.; Packer et al. 2011, p. (2013, p. 5), ‘‘such caps provide a short- Additionally, governmental bodies have 143). While some existing quota term means of reducing the risk of sometimes failed to analyze data and allocations have been derived from negative population impacts while more provide feedback to operators; experts information provided by hunting robust methods are being implemented. agree this failure undermines the concession operators, it has been noted Areas that are smaller than 1,000 km2 purpose of the system and encourages that many hunting concession operators should be granted the equivalent noncompliance. have not allowed independent fraction of 0.5 lions per year: For In the absence of reliable population population studies to take place, example, an area of 200 km2 would be estimates, age restriction on trophy possibly as a result of illegal activity allocated 0.1 lions per year, or one tag harvests can ensure sustainability and corruption (LionAid 2014a, pers. every ten years. Such a system would (Lindsey et al. 2013a, p. 8; Packer et al. comm.; Packer 2015, pers. comm.). Lion reduce the extent to which hunting in 2006, pp. 6–8). Whitman et al. (2004, experts also describe an over-reliance on small concessions adjacent to protected pp. 176–177) found that if offtake is subjective opinions, including input areas affects protected populations, as in restricted to males older than 6 years of from concession operators, in the Zambia and Zimbabwe.’’ age, trophy hunting will likely have process of developing quotas (Lindsey et Species experts also recommend, as minimal impact on the pride’s social al. 2013a, p. 8). As a result, information part of reforming trophy hunting, structure and young. By removing only underlying current quotas in much of adoption by range countries of an males 6 years of age or older, younger the species’ range has been inconsistent, adaptive quota management system that males remain in residence long enough biased, and/or lacking. It is difficult to would allow for quotas to fluctuate to rear a cohort of cubs (allowing their predict with accuracy what level of annually based on the population trends genes to enter the gene pool; increasing offtake would be appropriate to ensure of the species. An adaptive quota the overall genetic diversity); a quota is sustainable for a given management system would not only recruitment of these cubs ensures lion population without accurate prevent over-harvesting of lions, but population growth and therefore, information on the size of the resource would also prevent excessively sustainability. Simulations indicate that (LionAid 2014a, pers. comm.; Barnett conservative quotas (Hunter et al. 2013, populations with quotas of more than and Patterson 2005, p. 102). Therefore, p. 5). two male lions of minimum eligible age quotas not scientifically based are often Recognizing the inconsistencies in the of 3–4 years were more likely to too high to maintain sustainability and process of setting a quota and the experience extinction events than overharvest occurs. information on which they are based, populations with hunting restricted to a Lions are particularly vulnerable to range countries and conservationists minimum eligible age of 5–6-year-old excessive harvests due to impacts have been working to establish a set of males (Whitman et al. 2004, p. 176). associated with the removal of males best practices in order to create a more Additionally, full implementation and (Hunter et al. 2013, p. 2). As stated consistent, scientifically based approach enforcement of this age-based strategy before, except in Mozambique, quotas to determining quotas. The could potentially cause the need for are higher than the recommended recommended best practices include: (1) quotas to become irrelevant or maximum harvest of 1 lion per 2,000 establishing processes and procedures eliminated entirely. Age restrictions will km2. Additionally, mean actual harvests that are clearly outlined, transparent, naturally restrict offtake to a limited are higher than the recommended 1 lion and accountable; (2) establishing number of individuals that meet the age per 2,000 km2 offtake in Burkina Faso, processes and procedures that are CITES criteria (Loveridge et al. 2007, p. 549; Zambia, Namibia, and Zimbabwe compliant; (3) demonstrating Whitman et al. 2004, p. 177). (Lindsey et al. 2013, p. 8). Multiple management capacity; (4) standardizing Implementing this approach in the researchers have documented declines information sources; (5) establishing field involves conducting an age in lion populations across the range of monitoring systems for critical data; (6) assessment of male lions using the species as a result of mismanaged recording and analyzing trophy hunting identification techniques, such as mane trophy hunting. Specifically, negative data; (7) conducting data collection and development, facial markings, nose impacts to lions from excessive offtakes analysis for each hunting block and pigmentation, and tooth-aging to have been documented in Benin concession; and (8) establishing a establish the relative age of the target (Sogbohossou et al. 2014, entire), primary body who will approve quotas lion. Tooth wear on incisors, yellowing Cameroon (Croes et al. 2011, entire), (Burnett and Patterson 2005, p. 103). and chipping of teeth, coupled with Tanzania (Packer 2011, entire), Zambia Each country that allows trophy scars, head size, mane length and color, (Rosenblatt et al. 2014, p. entire; Becker hunting has some data collection system and thinning hair on the face, as well as et al. 2013, entire), and Zimbabwe in place; most countries have a central other factors can be an indicator of (Groom et al. 2014, entire; Davidson et wildlife authority that requires advanced age in lions (Whitman and al. 2011, entire; Loveridge et al. 2007, operators to submit data collection Packer 2006, entire). entire). Additionally, the effects of over- forms or questionnaires providing Whitman et al. (2004, p. 176) harvesting can extend into adjacent details of each of their hunts. However, postulated that ‘‘the most reliable index national parks where hunting does not according to the authors, these in the Serengeti/Ngorongoro lions is the occur (Packer et al. 2013, p. 636). guidelines have not been followed extent of dark pigmentation in the tip of Most experts consider the throughout much of the range countries, the nose, which becomes increasingly recommendation by Packer et al. (2011, which has led to a variety of compliance freckled with age. Individual variation p. 151) to limit offtake to no more than issues. Some systems have been overly in nose coloration is sufficiently low 1 lion per 2,000 km2 throughout its complex and cumbersome. ‘‘In 2000, that age can be estimated up to 8–9 range (or 1 per 1,000 km2 in areas with Zimbabwe, for example, had nine years. The noses of 5-yr-old males are 50 high density of lions) to be the best different forms, which contain [percent] black so the rule of thumb available science and recommend each essentially the same information, that would be to restrict all trophy hunting country impose a quota cap at those had to be completed by safari operators to males with noses that are more than

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half black.’’ Although this varies Niassa National Reserve), Tanzania, and Packer and Pusey 1984, and Pusey and individually and regionally, Zimbabwe have committed to Packer 1994) demonstrated that cub recommended best practices could be implementing the age-based strategy mortality increases when a new male regionally tailored. Packer et al. (2006, (White 2013, p. 14; Davidson et al. 2011, joins a pride. Infanticide is a common p. 7) note that males in South Africa p. 114; Whitman et al. 2004, p. 176), practice among many species, including require an additional 1–2 years to only two have fully implemented it lions (Hausfater et al. 1984, pp. 31, 145, become competitive with other males, (Henschel 2015, pers. comm.). Thus far, 173, 487). Removing a younger male and suggest a 7-year minimum might be Mozambique and Zimbabwe have lion allows another male of the pride to judicious for some regions. Therefore, implemented this strategy and shown a take over and kill the former patriarch’s there is concurrence by species experts reduction in total offtake (Henschel cubs; offspring younger than 2 years of that national or regional guidelines 2015, pers. comm.). They also appear to age are generally unable to defend should be developed to accompany be transparent in their implementation. themselves and may be killed or forced those produced in Tanzania and Zambia Tanzania has implemented age to disperse from the pride prematurely, (Lindsey et al. 2013a, p. 8; Packer and restrictions and shown reductions in which also often leads to death (Elliot Whitman 2006, entire). offtake; however, there is concern et al. 2014, p. 1054; Packer 2001, p. 829; According to Lindsey et al. (2013a, p. related to transparency (in terms of Pusey and Packer 1984, p. 279). This 8), some operators were uncertain of trophy quality data) and the scientific behavior is believed to be advantageous their ability to age lions; however, based objectivity of the evaluating body has to the incoming male as it increases and on research conducted in Niassa been questioned. Benin and Burkina accelerates the opportunity for the new National Reserve, Mozambique, hunters Faso committed to implementing age male to sire a cohort of cubs. When can be taught to age lions effectively. restrictions in 2014; their progress is females give birth to cubs, the female While experts agree it may be difficult currently pending. Lastly, Mozambique, generally does not return to estrus until to determine the exact age of a lion, excluding Niassa National Reserve and the cubs are around 18–24 months old broader categories based on age have Cameroon have not yet instituted or (Pusey and Packer 1984, p. 281). been developed to assist officials. For committed to the strategy (Henschel Following the loss of her cubs, however, example, Tanzania officials have ‘‘aging 2015, pers. comm.). Lack of a female will return to estrus rather sessions’’ wherein each concession implementation of age-based strategies quickly; females will resume mating operator is required to bring in the may undermine the successful use of within days or weeks, thus increasing skulls of their trophies for examination. trophy hunting as a sustainable the likelihood that the new male will Each skull is then classified as conservation strategy. have the chance to sire the next cohort. ‘‘acceptable’’ (6+ years old), ‘‘accepted Additionally, experts believe that Pusey and Packer (1984, p. 279) with penalties’’ (4–5 years old), and importing countries should have the calculated that infant fatality during ‘‘not accepted with deterrent penalties’’ ability to ascertain that the imported male takeovers accounted for 27 percent (<4 years) (Tanzania 2015, pp. 23–24). trophies originated from hunting of all cub fatalities under the age of 12 Tanzania reports that this step is concessions that fully comply with best months. required prior to any issuance of a practices. According to Lindsey et al. Further, when an adult male lion in CITES export permit. (2007, p. 3; Lindsey et al. 2006, pp. 285, a pride is killed, surviving males who Species experts place high emphasis 288), there is a market in the United form the pride’s coalition are vulnerable on the requirement for both enforcement States for conservation-based hunting. to takeover by other male coalitions, and and transparency in the strategy. A fully ‘‘In a survey of prospective clients 45– this often results in injury or death of transparent quota allocation system 99 percent were unwilling to hunt the remaining males (Davidson et al. would be one in which a quota under various scenarios if conservation 2011, p. 115). allocation system is based on scientific objectives would be compromised, and Recently, Elliot (2014, p. 1054) data received from all hunting areas and 86 percent were more willing to postulated that the impacts of male concession units annually, and would purchase a hunt if local communities takeovers due to trophy hunting may be require trophies to be independently would benefit’’ (Lindsey et al. 2007, p. more severe than previously recognized. evaluated, data on the trophies (e.g. age, 3). Experts agree that a fully transparent Specifically, when a pride male is sex, origin) be available nationally and system would allow hunters to choose removed and a new male takes over, internationally, and quotas based upon operators who have demonstrated a subadults may be forced to disperse data obtained from the previous hunting commitment to conservation principles; from the pride. These males are then at season (Henschel pers. comm. 2015). this system could provide incentives for a disadvantage as they are often Lion experts recommend age-based operators to comply with the inexperienced and physically smaller strategies be incorporated into lion recommended best practices. which may prevent them from being management action plans (Hunter et al. Harvesting of males that are too young able to compete with older males for 2013, pp. 4–5; Lindsey et al. 2013a, p. can have devastating impacts to the territory. In the study, Elliot found 100 8). Although the 6-year method has population. If male lions are harvested percent fatality for all males who potential to reduce the rate of too young (even as old as 3 years of age), dispersed earlier than 31 months old. infanticide in lion populations used for combined with quotas that are too high, The study concluded that dispersal of trophy hunting (Hunter et al. 2013, p. 4– the population will be driven to subadults is highly related to the 5; Lindsey et al. 2013a, p. 8), the issue extinction as female populations presence of incoming males, resulting in of incorporating this strategy into each collapse as they eventually are unable to a type of delayed infanticide, as many country’s conservation strategy and/or mate (Whitman et al. 2004, p. 176). of the subadults do not survive the action plan, and following up with Additionally, excessive trophy hunting dispersal. This effect may be amplified implementation, enforcement, and and taking of males under a certain age in populations that have a high offtake transparency has yet to be observed in cause male replacements and increased rate. Therefore, the author concluded many of the lion’s range countries infanticide rates (when males kill young that age restriction and reducing offtake (Henschel 2015, pers. comm.). While lion cubs sired by other males) could reduce takeover rates by new several countries, including Benin, (Whitman et al. 2004, p. 175). Packer males, allowing subadults a longer Burkina Faso, Mozambique (only in (2001, p. 829, citing Bertram 1975, period to mature prior to dispersal and

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thus, reducing the number of subadult et al. 2006, p. 7; Whitman 2004, p. 177). place, thereby, hunters only pay for deaths (Elliot et al. 2014, p. 1055). If concession operators selectively animals hunted. Other range countries A lack of mature males dispersing remove males in a manner that continue to have fixed quotas in place reduces the genetic viability of promotes healthy population growth, and charge a percentage of the quota populations and may contribute to local the lion population could yield more regardless of success (CAR charges 50 population extinctions (See Deleterious males in the long term (Davidson et al. percent; Namibia 100 percent; Tanzania Effects Due to Small Population Sizes). 2011, p. 114; Packer et al. 2006, p. 7; 40 percent; Zambia 60 percent; Selective offtake of large males may also Whitman et al. 2004, p. 176). Zimbabwe 30 percent). This approach modify the genetic evolution of lions. Hunter et al. (2013, pp. 2, 5) and facilitates harvesting of trophies even if Allendorf and Hard (2009, p. 9987) and Lindsey et al. (2013a, p. 9) identified a sufficiently old lion is not found Loveridge et al. (2007, p. 553) consider hunting of female lions to be another (Hunter et al. 2013, p. 6). Therefore, the genetic and evolutionary role of aspect of trophy hunting that is harmful harvested lions are often of lower selective hunting on wildlife to lion populations. Specifically, quality, younger, and less desirable populations. As individuals who females are the most productive portion male lions, as operators and hunters, display certain characteristics (such as of a population; if females are removed who had already paid the trophy fee, largest size) are more likely to be from a pride, there is inherent risk that had no incentive to be selective. harvested, this type of selective removal dependent cubs will die and the overall Abolishing fixed-quota fees and only will bring about genetic change in future breeding success of the pride will be allowing optional quotas will encourage generations. Specifically, removing the reduced. Packer et al. (2001 in Packer et and reward operators who are selective males with the most desirable traits al. 2006, pp. 5, 7) report that ‘‘large and follow age restrictions (Lindsey et from a population ultimately affects prides out-compete smaller prides and al. 2013a, p. 9; Packer et al. 2006, pp. upcoming generations as those per capita reproduction is lowest in 5, 9). individuals are no longer contributing to prides of only 1–2 females.’’ Lindsey et To ensure hunters have adequate time the gene pool. ‘‘For example, the al. (2013a, pp. 2, 4, 9) indicate that a to be selective in trophies harvested, frequency of elephants (Loxodonta loss of a female increases a pride’s and to ensure the revenue earning africana) without tusks increased from vulnerability to territory loss. As a potential is maximized, experts 10 percent to 38 percent in South result, removing females has injurious recommend that a minimum stipulated Luangwa National Park, Zambia, effects on the overall success of the hunt length be set at 21 days. However, apparently brought about by poaching of population and, ultimately, the number many countries either have no limits on elephants for their ’’ (Jachmann et of harvestable males. length of hunting safaris or have too Lindsey et al. (2013a, pp. 2, 4, 9) al. 1995 in Allendorf and Hard 2009, p. short a minimum length (Lindsey et al. indicate that quotas are currently 9987). This comparison relates to lion as 2013a, p. 9). Currently, there are no set the removal of the largest males available for female lions in some lengths for hunting safaris in consequentially results in females locations within Namibia, and between Mozambique, Namibia, Zambia, and breeding with less desirable males and 1990 and 2011, in Zimbabwe (Packer et Zimbabwe. Burkino Faso has a thus, perpetuating the production of less al. 2006, p. 4). Between 1998 and 2004, minimum requirement of 12 days, and desirable individuals. Selective offtake Zimbabwe maintained a mean quota of Benin and Cameroon require 12 to 14 based on gender also has the potential 0.3 ± 0.1/100 km2 for female lions; days. Tanzania has a minimum length of to skew sex ratios and impact breeding during the same period, actual offtake 21 days while CAR varies from 12 to 21 success, as has been the case for lions was lower at 0.08 ± 0.1/100 km2, or a days (Lindsey et al. 2013a, pp. 2–3). (Allendorf and Hard 2009, p. 9991; mean of 30.6 percent of the quota Loveridge et al. 2007, p. 553). The actually harvested (Loveridge et al. Several other problems with current authors state that in order to maintain 2007, p. 551). Zimbabwe discontinued management of lion trophy hunting are the highest yield and viability of the issuing quotas for females in 2011. likely to worsen negative impacts most desirable males, one option is to be Female hunting is not allowed associated with hunting of lions and less selective (Allendorf and Hard 2009, elsewhere within the range of the undermine conservation incentives. p. 9991). Specifically as related to lions, species (Lindsey et al. 2013a, p. 2). Corruption, allocation of hunting this would mean implementing age Species experts recommend that the concessions, and lack of benefits and restrictions so that the more desirable trophy hunting of females be prohibited, recognition of the role communities males are not harvested prior to unless the management plan is play in conservation have been successful reproduction. specifically to control the size of the identified (Lindsey et al. 2013a, pp. 2– Whitman et al. (2004, pp. 175–177) lion population (Hunter et al. 2013, p. 3, 9). found that if offtake is restricted to 5; Lindsey et al. 2013a, p. 9). Corruption is widespread within the males 6 years of age or older, the Another deficiency in current trophy range of the lion (Transparency impacts of trophy hunting are likely to hunting management is the use of fixed International 2014, unpaginated). All be minimal on the prides social quotas. There are two primary types of but one lion range country (Botswana) structure and reproduction. Therefore, quotas, ‘‘fixed’’ and ‘‘optional.’’ Trophy scored below 50 (out of 100) on experts recommend that a 6-year age fees for fixed quotas require the Transparency International’s 2014 restriction should be implemented for payment of a portion (40–100 percent) Corruption Perception Index (CPI), all hunting concessions throughout the of the lion trophy fee, regardless of which measures perceived levels of lion’s range. whether the hunt is successful, whereas public sector corruption based on expert Species experts have suggested an optional quotas are paid by operators opinion and is based on a scale of 0 additional mechanism that could help only when the lion is shot. Until 1999, (highly corrupt) to 100 (very clean). reduce infanticide. In concessions male lions were typically on fixed Approximately half of the current lion where operators can distinguish quotas, whereas female lions were range countries—including Tanzania between resident and solitary under optional quotas. According to and Kenya, where more than half of all individuals, removal of the nomadic Lindsey et al. (2013a, pp. 2–3), wild lions occur—are among the most males may reduce the likelihood of a Mozambique, Benin, Burkino Faso, and corrupt countries in the world, ranking possible conflict and take-over (Packer Cameroon all have optional quotas in in the lower 30 percent of 174 countries

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assessed (Transparency International the success of national strategies to public auctions for 3-year periods, while 2014, unpaginated). conserve these species (Smith et al. community conservancies lease for a 5- Corruption is particularly prevalent in 2003, p. 69). Corruption can reduce the year period via a closed-tender process. areas with extreme poverty effectiveness of conservation programs Zimbabwe holds a public auction for (Transparency International 2014, by reducing the funding, law state safari areas, with the option to unpaginated; Michler 2013, pp. 1–3; enforcement, and political support extend 5 years based on performance. Kimati 2012, p. 1; Garnett et al. 2011, p. available for conservation, and also by Communal Areas Management 1; IUCN 2009, p. 89; Leader-Williams et acting as an incentive for the Programme for Indigenous Resources al. 2009, pp. 296–298; Kideghesho 2008, overexploitation of resources (Garnett et (CAMPFIRE) areas are leased on 3–10 pp. 16–17). Certain circumstances tend al. 2011, p. 1, citing several sources; year-period using a closed-tender to promote corruption, such as Smith and Walpole 2005, p. 252). Given process (Lindsey et al. 2013a, pp. 2–3). opportunity for financial gain, weak rule the financial gains to be made from lion The chief complaint regarding this of law, abnormal concentrations of trophy hunting, and the high level of system is that concession areas are power in one individual or institution, corruption in many lion range countries leased to operators without regard for no counter-balancing mechanisms in (Packer 2015, pers. comm.; the operators’ track record in place among different government Transparency International 2014, conservation. Zimbabwe is the only agencies, and reliance on discretionary unpaginated), it is reasonable to country that renews based on operator powers for allocation of permits, conclude that corruption and the performance (Lindsey et al. 2013a, pp. licenses, or activities (Smith et al. 2015, inability to control it are having 2, 9). Lindsey et al. (2007, p. 2) found p. 953; Nelson 2009, unpaginated; Luo negative impacts on decisions made that various countries have problems 2005 in Smith et al. 2015, p. 953). about lion management in many areas of with their allocation process, ‘‘with the Corruption manifests itself in several the species’ range and on lion effect that they are sometimes sold too ways, such as embezzling of public populations, and undermine steps to cheaply, allocated for periods too short funds, fraud, demanding or accepting reform hunting of lions. The impacts to promote responsible custodianship, bribes to overlook illegal activities, highlight the importance of and occasionally given to unlicensed interference in decisions to implement transparency within the hunting operators. . .. In several countries large conservation measures, and offering industry and independent verification citizen quotas are provided to urban patronage, nepotism, and political of processes such as quota setting, residents at low prices, reducing influence (Vargas-Hernandez 2013 in trophy monitoring, and concession revenues from trophy hunting and Smith et al. 2015, p. 953; Garnett et al. allocation (Lindsey et al. 2013a, p. 9). reducing incentives for communities to 2011, p. 1; Leader-Williams et al. 2009, In recent years, leadership in several conserve wildlife.’’ Experts believe that p. 301; Kaufmann 1997 in Leader- African lion range countries has taken basing the ability to renew a concession Williams et al. 2009, p. 297). With steps to address corruption, or activities lease on operators’ past performance respect to lion management, it may that facilitate corruption, associated records could be an incentive for include, for example: Infringement of with wildlife management. For example, operators to comply with best practices. hunting regulations in the field; in 2013, the Tourism Minister of Zambia Thus, experts recommend concession acceptance of bribes to overlook illegal banned hunting in 19 game management allocation should base concession lease activities such as poaching; interference areas for 1 year due to allegations of renewals on operator performance in or mismanagement in monitoring and corruption and malpractice among the regard to best practices compliance. setting of hunting quotas and in issuing hunting companies and various As discussed under Human–lion of licenses; misappropriation of hunting government departments. Some game Conflict, the risk of retaliatory killing is fees; allocation of hunting blocks based management areas and privately owned elevated in many cases due to the fact on patronage and nepotism or to game ranches were not included in the that communities living in close persons presumably considered to be of ban, but lion hunting appears to be proximity to lion populations often bear financial or other strategic importance; prohibited throughout the country the cost of that proximity (e.g., loss of and allocation of hunting blocks at less (Michler 2013, pp. 1–3). Whether recent valuable livestock due to lion than competitive prices (see Leader- reforms taken by various lion range depredation), but receive little of the Williams et al. 2009, pp. 301–305; countries will reduce the effect of benefits generated by the presence of Nelson 2009, unpaginated). corruption on lion management and, lion in the trophy hunting and Peh and Dori (2010, pp. 336–337) therefore, lion populations is as yet ecotourism industries (Lindsey et al. show that global indices of corruption unknown. 2013a, p. 9). Trophy hunting can and governance are highly correlated Most concessions in the African range generate millions of dollars in annual with those of environmental of the lion use a closed-tender process revenue (see Potential Benefits of performance—countries with high for land management. A closed-tender Trophy Hunting). levels of corruption have lower levels of system is the process of selling a In the past, government and private environmental performance. Further, product by inviting a specific group of land owners were the primary Smith et al. (2003, entire) found strong potential buyers to provide a written beneficiaries of the revenue gained; associations between changes (declines) offer by a specified date. In the case of currently efforts are being made in many in elephant and numbers and a hunting concession, the owner of the range countries to incorporate governance scores. Governance scores, property thus sells a lease on a property incentives at the local level (Barnett and which were based largely on for a given length of time. Countries that Patterson 2005, p. vi). Many range Transparency International’s CPI, use this process for state-owned lands countries are now recognizing the need explained observed changes in numbers include Benin (lease is for 5 years); to incorporate incentives and local of elephants and better Burkina Faso (20 years); Cameroon (10 community benefits into their trophy than per capita GDP, Human years, renewable); CAR (10 years hunting regulations, land management Development Index scores, and human (renewable); Mozambique (10+ years); policies, and lion conservation action population density. These results Tanzania (5 years); and Zambia (10–15 plans. Most countries that allow lion suggest that political corruption may years based on status of wildlife). In trophy hunting have developed National play a significant role in determining Namibia, state concessions lease land by Poverty Reduction Strategies and

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discussed benefit distribution and evidenced by the fact that 99 percent of 2). Researchers have documented management to rural communities (see lion trophies from South Africa are of declines in populations across the range Potential Benefits of Trophy Hunting). captive origin. Lindsey et al. (2012, p. of the species that were a direct result Although positive steps are being taken 21) warn that future efforts to control of mismanaged trophy hunting to address local community incentives, hunting of captive-bred lions could (Rosenblatt et al. 2014, p. entire; most of the countries are currently not potentially increase the demand for Sogbohossou et al. 2014, entire; Becker transparent about the benefits provided wild lion trophies and result in et al. 2013, entire; Lindsey et al. 2013, to local communities, and due to the excessive harvests. However, we also entire; Croes et al. 2011, entire; Packer high revenue potential are subject to note that trade in bones of captive lions 2011, entire; Loveridge et al. 2007, corruption. could stimulate harvest of wild lions to entire). Six management weaknesses supply a growing bone trade (Lindsey et have been identified in the current Captive Lions al. 2012, p. 20). Hunting of captive lions management of lion hunting. These In analyzing threats to a species, we could also potentially undermine the weaknesses include: (1) A lack of focus our analysis on threats acting price of wild hunts and reduce scientifically based quota that results in upon wild specimens within the native incentives for conservation of wild lions excessive harvests; (2) a lack of range of the species, because the goal of in other African countries (Lindsey et al. enforcement in age restrictions, which the Act is survival and recovery of the 2012, p. 12). leads to unsustainable harvests, species within its native ecosystem. We Limited research has been conducted increased rates of infanticide, and do not separately analyze ‘‘threats’’ to on the use of captive-raised lions for population declines; (3) hunting of captive-held specimens because the reintroduction purposes. Existing female lions in Namibia, which statutory five factors under section 4 (16 research has generally found that decreases reproduction success, thereby U.S.C. 1533) are not well-suited to captive-raised lions are not as able to decreasing males available for trophy consideration of specimens in captivity, successfully adapt to conditions out of hunting; (4) the use of fixed quotas, and captive-held specimens are not captivity and therefore, the success rate which encourages hunters to be eligible for separate consideration for is much reduced compared to the use of unselective in their take of a trophy (i.e., listing. However, we do consider the wild-caught lions. Although some they will kill younger, less desirable extent to which specimens held in potential exists that the captive-lion males); (5) a lack of minimum hunt captivity create, contribute to, reduce, or industry in South Africa may benefit lengths or minimum lengths that are too remove threats to the species. some local wild populations, additional short to allow hunters the time needed In 2009, approximately 3,600 captive- research would be needed to verify this to be more selective in their take of held lions were managed for trophy claim. As a result, we do not believe trophies; and (6) general problems hunting across 174 breeding facilities in that the captive-lion industry currently associated with management of trophy South Africa ((Lindsey et al. 2012, p. 18, contributes to, reduces, or removes hunting, including corruption, citing Taijaard 2009; Barnett et al. threats to the species. allocation of concessions, and lack of 2006a, p. 513). The captive-breeding Summary of Trophy Hunting benefits to communities and recognition industry often publicizes captive of the important role they play in breeding and reintroduction of captive- If trophy hunting of lions is part of a conservation. born species into the wild as a potential scientifically based management Most P. l. leo populations are solution to the decrease in wild lion program, it can provide considerable extremely small, isolated, and rapidly populations. However, lions raised in benefits to the species by reducing or declining. Of the 18 countries captivity often develop a variety of removing incentives to kill lions in documented to allow lion trophy issues that make them unsuitable for retaliation for livestock losses, and by hunting, 8 are in the range of P. l. leo. reintroduction. Captive lions in general reducing the conversion of lion habitat However, we note that due to the lack are not suitable for reintroduction due to agriculture. Trophy hunting, if of lions in some of these countries, it is to their uncertain genetic origins managed well and with local unlikely that all of these countries could (Barnett et al. 2006a, p. 513; Hunter et communities in mind, can bring in conduct lion trophy hunts. A study al. 2012, p. 3), potential maladaptive needed revenue, jobs, and a much- found that quotas in Benin and Burkina behaviors, and higher failure risk needed protein source to impoverished Faso are too high for sustainability, compared to translocated individuals local communities, demonstrating the although Burkina Faso has proposed to (Hunter et al. 2012, pp. 2–3). Research value of lions (Groom 2013, pp. 1–3; reduce their quota in the 2015–2016 has indicated that restoration efforts Lindsey et al. 2006, pp. 283, 289). In season (Henschel 2015, pers. comm.; using wild-caught individuals have a addition, the amount of habitat that has Lindsey et al. 2013a, p. 6). Actual much higher rate of success than those been set aside by range countries harvests in Burkina Faso were also using captive-raised individuals for a specifically for trophy hunting has found to be higher than the level large variety of species (Hunter et al. greatly increased the range and habitat recommended by Packer et al. (2011, p. 2012, p. 21). Currently, reintroduction of lions and their prey base, which 151). Additionally, Benin and Burkina efforts of captive-raised lions have not contrasts the overall ongoing rate of Faso have committed to implementing been shown to address the underlying habitat destruction occurring in Africa. an age-based strategy, but have yet to causes of populations’ declines The total amount of land set aside for implement it. As a result, species throughout the species range. trophy hunting throughout Africa experts agree that there is no level of We note that while the captive-lion exceeds the total area of the national offtake that would be sustainable for P. industry may not be contributing to the parks, providing half the amount of l. leo populations in their current conservation of the species in the wild viable lion habitat (Chardonnet et al. condition (Bauer 2015, pers. comm.; via reintroduction, the captive-lion 2010, p. 34; Packer et al. 2006, pp. 9– Henschel et al. 2014, entire; Henschel et industry in South Africa may reduce the 10). al. 2010, entire). pressures of trophy hunting on the wild The main problem with mismanaged Of the 18 countries documented to populations in South Africa (Hargreaves trophy hunting stems from excessive allow lion trophy hunting, 10 are in the 2010b in Lindsey et al. 2012, p. 12; harvests and impacts associated with range of P. l. melanochaita. However, Lindsey et al. 2012, p. 19), which is removal of males (Hunter et al. 2013, p. we note that, like the situation with P.

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l. leo, due to a lack of lion populations purposes that are not detrimental to the generally valid for 6 months, and a in some of these countries, it is likely survival of the species in the wild and trophy could in theory be exported the that fewer countries could conduct lion that the specimen will not be used for year after it was hunted). The second trophy hunts. A study found that primarily commercial purposes. For live limitation to interpreting this Namibia, Tanzania, Zambia, and specimens, a finding must also be made information is that, although many Zimbabwe all had quotas higher than that the recipient is suitably equipped to permits may indicate that an animal is the recommended level for house and care for the specimens of wild origin (source code ‘‘W’’), these sustainability; however, Zimbabwe has (CITES Article III(3)). Export permits are permits may be incorrectly coded. This reduced their quota. Mozambique issued only if findings are made that the is true for South Africa, where during (Niassa National Reserve) is the only specimen was legally acquired and the the period of 2000 to 2009, animals that location found to have a quota below export is not detrimental to the survival were captive born and released into the recommended level. Age-based of the species in the wild, and that a private reserve systems were assigned strategies have been implemented and living specimen will be so prepared and an incorrect source code of ‘‘W.’’ South shown to reduce offtakes in shipped as to minimize the risk of Africa has since requested their Mozambique (only in Niassa National injury, damage to health, or cruel provincial authorities to use the correct Reserve, excludes the rest of the treatment, and that the CITES source code for ‘‘captive bred’’ in order country), Tanzania, and Zimbabwe. Management Authority of the exporting to correctly reflect the source of sport- Furthermore, Zimbabwe and Niassa country is satisfied that an import hunted lion trophies; however, some National Reserve are the only two permit has been granted for the provinces are not complying (RSA 2013, locations that have fully implemented specimen (CITES Article III(2)). pp. 8–9). Based on South African trade an age-based strategy with transparency, CITES Appendix II includes species data, the bulk of lion exports and their an element experts say is critical to a that are less vulnerable to extinction parts and products (including trophies) quota allocation system. Several other than species listed in Appendix I, and are from captive-born lions (RSA 2013, countries have made commitments to ‘‘although not necessarily now p. 7). implement the age-restrictions strategy threatened with extinction, may become Tanzania, with one of the largest lion but their progress is pending. In South so unless trade in specimens of such populations (Hamunyela et al. 2013, pp. Africa, 99 percent of the lion trophies species is subject to strict regulation in 29, 283; Riggio et al. 2013, p. 32; Ikanda are captive bred, and, therefore, were order to avoid utilization incompatible 2008, p. 4; Baldus 2004, pp. 5, 6), was not the result of removing lions from the with their survival.’’ Species listed in the largest exporter of wild-origin lion wild. Appendix II of CITES may be trophies, but their exports have Unless reforms are made to the commercially traded, subject to several decreased significantly since 2008. In current management of trophy hunting, restrictions. 2008, approximately 138 trophies were we expect the declines specifically Although each country has its own exported from Tanzania; in 2010, 128 documented from excessive offtakes in method of regulating trophy hunting, were exported; in 2011, 55 were Benin, Cameroon, Tanzania, Zambia, international trade of lion trophies must exported; in 2012, 62 were exported (it and Zimbabwe to continue. adhere to CITES. International trade of should be noted that in 2012 Tanzania Furthermore, we expect excessive lion parts and products (including established an annual quota to limit harvests to further contribute to declines trophies) are reported by both the trophy hunting to no more than 50 in the species across its African range. exporting and importing countries and tracked by the United Nations animals (Jackson 2013, p. 7); and in Import/Export of Lion Trophies Environment Programme World 2013, 11 were exported (UNEP–WCMC The lion species (Panthera leo) is Conservation Monitoring Centre 2014, unpaginated). Again, it should be listed in Appendix II of CITES; (UNEP–WCMC). noted that there may be discrepancies however, the former Asiatic lion (P. l. According to the UNEP–WCMC between the annual quota and the actual persica) is listed in Appendix I. CITES CITES Trade Database, between 2005 number of trophies exported in a given is an international agreement through and 2012, exports of lion trophies year (see http://www.cites.org/common/ which member countries work together demonstrated a decreasing trend, if resources/TradeDatabaseGuide.pdf for to protect against over-exploitation of exports of captive-born lions from South additional information). Regardless, the animal and plant species found in Africa are excluded (UNEP–WCMC numbers of lion trophies exported by international trade. Parties regulate and 2014, unpaginated). UNEP–WCMC Tanzania according to the UNEP– monitor international trade in CITES- indicates that 521 lion trophies were WCMC CITES Trade Database suggest a listed species—that is, their import, exported (excluding South Africa) in decreasing trend. export, and reexport, and introduction 2005 and 303 were reported (excluding Additionally, some trophies are from the sea—through a system of South Africa) in 2012. exported from source countries under permits and certificates. CITES lists It should be noted that there are the ‘‘skins’’ category. According to the species in one of three appendices— limitations to interpreting the above most recent data available, the United Appendix I, II, or III. reported information. The 2004 guide to States imported skins of wild origin An Appendix-I listing includes using the CITES Trade Database from four African countries in 2013; 9 species threatened with extinction indicates that the outputs produced by from Mozambique, 5 from Tanzania, 2 whose trade is permitted only under the CITES Trade Database can be easily from South Africa, and 22 from exceptional circumstances, which misinterpreted if one is not familiar Zimbabwe. The purpose code for these generally precludes commercial trade. with it (CITES 2004b, p. 5). The number imports was ‘‘Trophy Hunt,’’ except for The import of specimens (both live and of ‘‘trophies’’ reported does not the two skins from South Africa which dead, as well as parts and products) of necessarily equate to the number of were coded as ‘‘Commercial.’’ an Appendix-I species generally lions hunted. Additionally, the number For 2013, the most recent year for requires the issuance of both an import of trophies reported for a given year in which complete CITES trade data are and export permit under CITES. Import the trade report does not equate directly available, U.S. CITES Annual Report permits are issued only if findings are to the number of animals hunted in that trade data indicate that the United made that the import would be for given year (CITES export permits are States allowed the direct import of lion

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trophies from seven African countries, CITES 2014, p. 12), the trade in lion increasingly used as a replacement for as follows: skins is most likely one of the biggest tiger bone. Thus, the lion bone trade Botswana = 1 trophy (originated from threats to lion survival in western Africa could potentially follow the same Mozambique) due to the rarity of lions in the region, course as the tiger bone trade: Become Burkino Faso = 3 trophies the extent of the trade, and the high lucrative, spur considerable demand Mozambique = 5 trophies price of lion skins. from suppliers of the black market, Namibia = 9 trophies In southern and eastern Africa, trade result in extensive poaching of wild South Africa = 545 trophies (the in lion parts, particularly lion bone, to individuals, and have significant majority of which are reported to be Asia is generally considered a severe impacts to wild populations. of captive-born origin; additionally 2 potential threat to the species (Bauer Certain aspects of the current lion captive trophies originated in South 2015, pers. comm.). According to CITES bone trade suggest that the potential for Africa, imported to Canada, and then (2014, p. 14), there is ‘‘clear scope for the trade to impact wild lion imported into the ) the international trade in lion body populations may be high. For example, Tanzania = 3 trophies parts for [traditional Chinese medicine evidence suggests that demand from Zambia = 17 trophies and traditional African medicine] to Asia for lion bone is increasing rapidly. Zimbabwe = 44 trophies grow uncontrollably, as it has done for Based on Williams (2015, pp. ix–x, 46), other big cats.’’ during 1982–2000, only nine lion Based on CITES trade data, lion Lion bones are used as a substitute for skeletons were exported from trophy exports have decreased tiger (Panthera tigris) bone in traditional worldwide sources, destined primarily throughout most of the lion’s range, Asian medicine and in Asian luxury to Europe. CITES permit records show including Tanzania, which has one of products (Williams et al. 2015, pp. 2–3, only three exported from South Africa the largest lion populations. South 5; Graham–Rowe 2011, pp. s101–s102). prior to 2008, destined for Denmark. In Africa is the only country where exports Lion bones are difficult to distinguish 2008, South Africa began issuing CITES have increased because most of these from tiger bones (Williams et al. 2015, permits for the export of skeletons of trophies are of captive origin. pp. 8, 102; Wildlife Protection Society captive-bred lions to Asia. These Traditional Use of Lion Parts and of India 2007, unpaginated), and are exports currently appear to come Products sold into Asian markets as tiger bone primarily from South Africa’s captive- fakes (Williams et al. 2015, pp. 2–3, 62, bred lion hunting industry as a Lion parts and products are used in citing several sources). Tiger bone is byproduct of trophy hunting. The many African countries as medicine, highly valued in Asia, primarily in number of lion skeletons for which nutrition, talismans, and decorations, China and Vietnam, and there is South Africa issued permits for export and in traditional ceremonies and considerable demand for it (Williams et to Asia (China, Viet Nam, Thailand and rituals (CITES 2014, p. 7; Burton et al. al. 2015, p. 1; Gratwicke et al. 2008, pp. Lao PDR) increased tenfold from 2008 to 2010, p. 4). CITES (2014, p. 8) reports 2–5; Graham-Rowe 2011, pp. s101– 2011, from about 50 to about 573 that many African countries, including s102). Consequently, tiger bones are one skeletons, respectively, representing a Somalia, Nigeria, Burkina Faso, Kenya, of the most lucrative products on the total of 1,160 skeletons or about 10.8 and Cameroon, maintain local markets illegal wildlife market (Haken 2011, in metric tons (11.9 US tons) of lion bone in lion products. Parts used include Williams et al. 2015, p. 1)—the retail in 4 years (Williams 2015, pp. ix–x, 46). skin, teeth, claws, fat, whiskers, bone, price of raw tiger bone can reach Further, according to the Government of bile, testicles, meat, and tails. In $1,250–3,750 USD per kilogram (Nowell Kenya (2015, p. 3), the declared exports addition, lion bone is also used in Asia and Ling 2007, p. 23). of bones, skulls, and skeletons derived as a substitute for tiger bone in Tigers are categorized by IUCN as from wild lions also show an increasing traditional Asian medicine (Williams et endangered (Goodrich 2015, p. 2). trend through the period 2003–2012, al. 2015, pp. 2, 62). Globally, the tiger population has with total declared specimens in 2012 While quantitative data is lacking, declined from what is believed to have more than ten times those in 2003. With according to a peer reviewer (Bauer been 100,000 at the turn of the 19th respect to meeting demand for lion 2015, pers. comm.), trade in lion parts century (Jackson 1993, in Nijman and bone, Lindsey et al. (2012, p. 20) state and products is very common within Shepherd 2015, p. 1) to an estimated that there are likely to be large numbers western and central Africa. Responses to 5,000–7,000 in 1998, to 3,159 tigers in of lion bones available for export from the CITES periodic review consultation 2014 (Goodrich 2015, p. 7; Seidensticker game farms, from lionesses and non- process support this claim: Trade in lion et al. 1999, in Goodrich et al. 2015, p. trophy males, and as byproducts from skins and partial skins is described as 7). Poaching for the illegal trade in tiger animals shot as trophies. In addition, ‘‘frequent’’ in street markets in Abidjan, parts, especially bone has become a Williams et al. (2015, p. 41) report that Coˆte d’Ivoire; lion skins and canines are major driver in the species’ decline there may be between 1,400 and 6,200 described as ‘‘easily found’’ in the (Goodrich et al. 2015, p. 9; Williams et lion skeletons from past trophy hunts on markets of Dakar, Senegal; and the scale al. 2015, p. 1; Nowell and Ling 2007, p. South African game farms that could of domestic trade in illegal lion v). While wild tiger populations are potentially be used to supply demand products is described as ‘‘massive’’ in declining, the demand for tiger parts in for lion bone. However, considering the Nigeria (CITES 2014, pp. 5–6). Further, Asia is increasing (Williams et al. 2015, sharp and continuing increases in in the central African country of p. 5; United Nations Office on Drugs demand from Asia for lion bone, there Cameroon, the estimated value of a and Crime 2013, p. 81; United Nations is potential for demand to surpass the single lion carcass exceeds the trophy Office on Drugs and Crime 2010, pp. 10, availability of legally obtained lion bone fee, and at a lion conservation 17; Nowell and Ling 2007, p. 4). This and, consequently, result in poaching of conference the Government of increasing demand for tiger parts has wild lions to meet demand. Cameroon identified trade in lion skins led to the rise of tiger farms, where live In addition, recent evidence strongly as a major cause of the decline in lion captive bred tigers appear to be utilized suggests live lions are being used to populations in western and central to supply the bone trade within China supply the lion bone trade (Williams et Africa (LAGA pers. comm., in CITES (Denyer 2015, unpaginated). With tigers al. 2015, pp. ix, 2–3, 42–44). In August 2014, p. 12). According to Henschel (in difficult to obtain, lion bone may be 2006 a live Asiatic lion was observed in

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a market in Mong La, Myanmar (Oswell, Lastly, evidence suggests incentive to meaning they are constantly present, but 2010, p. 12). The town, known for poach wild lions for the bone trade may often do not cause disease. Others are incidents of wildlife trafficking, is less currently exist. According to Williams epidemic and cause a sudden severe than 2km from the Chinese border. Up et al. (2015, p. x), the 2013 price paid outbreak with the potential to cause to 2006/2007, Williams et al. (2015, p. to South African game farmers and high mortality (Craft 2008, pp. 5, 6). The x, Table 11, Figure 24) noted: landowners for lion bones was $1,260– association between disease, age, ‘‘The combined quantity of live lions and 2,100 USD per skeleton. In many lion nutritional health and other factors that lion parts and derivatives exported to East– range states this exceeds per capita GDP could predispose a lion to morbidity Southeast Asia from South Africa was (gross domestic product) (World Bank and, eventually, mortality is complex. It minimal in the broader global trade. From 2015, unpaginated). Thus, the current is often difficult to determine whether 2008, however, the quantities exported price paid for lion bone appears to mortality was due to a single factor or increased almost six-fold from the previous provide incentive in some countries to a combination. Lions could be infected year. Not only did the number of live lions poach wild lions. with and become debilitated by a exported to East–Southeast Asia reach record While the lion bone trade appears to disease, but the actual cause of death levels from this time, but also the first currently be based primarily in South could be other factors, such as fighting permits to export lion skeletons were issued. The demand for lion parts and derivatives Africa’s captive-bred lion hunting with other lions or large predators appears to have coincided with the industry, the trade appears to be having (LionAid 2014a, p. 4). strengthened conservation measures adopted little or no impact on wild lion Feline calicivirus, feline herpesvirus, in 2006–2007 to protect tigers and Asian big populations in South Africa at this feline parvovirus, feline coronavirus, cats. Accordingly, tiger parts were time—lion populations in South Africa and feline leukemia virus are endemic increasingly substituted with lion parts are stable or increasing and there is little viruses known to occur in lions of obtained from Africa. The trade in lion parts poaching of wild lions in the country , Ngorongoro and derivatives to Lao PDR dominates the (Funston and Levendal 2014, pp. 1, 26; Crater, Lake Manyara National Park, exports. Since 1998, but especially after Williams et al. 2015, pp. 79–80). , and Etosha 2007, China, Viet Nam, Lao PDR, Myanmar and Thailand have imported increasing However, the impact of the lion bone National Park (but not all viruses are amounts of live lions, lion bodies and bones trade on lion populations outside South known in all parks). However, these from South Africa.’’ Africa is unknown, and most wild lions diseases are not known to affect lion occur outside South Africa (see survival (Hunter et al. 2012, p. 2; Craft Evidence also indicates ‘‘well Distribution and Abundance). Based on 2008, p. 6; Hofmann-Lehmann 1996, pp. established’’ links between South the effect of the tiger bone trade on tiger 559, 561). Africa’s legal lion bone trade and the populations, if current conditions—for Lions within Kruger National Park Xaysavang Network, an international example, rapidly increasing demand and Hluhluwe-iMfolozi Park, South wildlife trafficking syndicate that is also and involvement of an international Africa, and Serengeti National Park, involved in the illicit rhino trade crime syndicate—continue unchanged, Tanzania, are known to be infected with in South Africa (Williams et al. 2015, then there is considerable potential for Mycobacterium bovis, a pathogen that pp. 7–10, 59; Environmental extensive poaching of wild lions to causes bovine tuberculosis (bTB). This Investigative Agency 2014, p. 13; U.S. occur in order to meet demand. pathogen is not endemic to African Department of State 2013, unpaginated). wildlife and was likely introduced from The U.S. Department of State has issued Disease cattle imported from Europe. M. bovis is a $1 million reward for information Wild lions are known to be infected transmitted to ungulates, such as leading to the dismantling of this with various pathogens (Hunter et al. (Syncerus caffer) and network. According to the U.S. 2012, p. 2; Craft 2008, p. 6; Michel et al. wildebeest (Connochaetes taurinus), Department of State, the Xaysavang 2006, p. 92; Hofmann-Lehmann et al. from domestic cattle located on the Network facilitates the killing of 1996, pp. 559–561). However, periphery of the parks (Maas et al. 2012, endangered species in Africa and information on the extent of infections p. 4206; Keet et al. 2009, pp. 4, 11; elsewhere and smuggles them to Laos and impacts of diseases on lion Renwick et al. 2007, p. 532; Michel et for export to other Asian countries (U.S. populations is limited. We found one al. 2006, pp. 92, 93; Cleaveland et al. Department of State 2013, unpaginated). study documenting disease in a single 2005, pp. 446, 449, 450). Spillover of the During 2008–2011, the vast majority wild lion in India that died from disease from buffalo to other lion prey (85%) of the permits issued by South trypanosomiasis in 2007; analysis of species, such as kudu (Tragelaphus Africa to export lion skeletons or tissue samples also detected peste des strepsiceros) and warthog carcasses were issued for exports to petits virus (PPRV), which is (Phacochoerus africanus), has also been Laos (Williams et al. 2015, pp. x, 46) not known to cause disease in documented (Keet et al. 2009, pp. 4, 11; and, for the only 2 years for which data carnivores (LionAid 2013, unpaginated; Renwick et al. 2007, p. 535; Cleaveland were available (2009 and 2010), over Balamurugan et al. 2012, pp. 203, 205). et al. 2005, p. 450). Because the lion’s half of the consignments destined for Information on the presence of disease primary prey are infected with bTB, Laos were listed as imported by Vixay and impacts to lions come from a few they are frequently exposed to large Keosavang, believed by the U.S. long-term studies that have been amounts of infected tissue and are at Department of State to be the leader of conducted in Africa, including risk of infection (Keet et al. 2009, pp. 4, the Xaysavang network (U.S. Serengeti National Park, Ngorongoro 6; Renwick et al. 2007, pp. 532, 536; Department of State 2013, unpaginated; Crater, and Kruger National Park. Michel et al. 2006, p. 93; Cleaveland et Williams et al. 2015, pp.8–10). The As a result of human population al. 2005, pp. 450, 451). Furthermore, involvement of the Xaysavang Network expansion into lion habitat, lions are predators prey on weak animals and in South Africa’s lion bone trade increasingly exposed to diseases from scavenge on carcasses, increasing their indicates there are well-established domestic animals (IUCN 2006b, p. 26). likelihood of being exposed to M. bovis avenues for laundering of illegally Because lions are a top predator, they (Renwick et al. 2007, p. 536; Michel et obtained lion bones, such as those are at a particularly high risk of al. 2006, p. 93). Transmission may also obtained from poached wild lions, into exposure to pathogens (Keet et al. 2009, occur among lions via scratching and the legal trade. p. 11). Some pathogens are endemic, biting (Keet et al. 2009, p. 7; Renwick

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et al. 2007, pp. 532–533). M. bovis is a infection (Renwick et al. 2007, p. 533). CDV events have been harmless. pathogen that causes the infected The impact of bTB on lions is largely However, in 1994 and 2001, CDV animal to remain infectious and, unknown. Researchers suggest that bTB epidemics in the Serengeti National therefore, a source of infection, until it may lower breeding success, reduce Park/ National Reserve and dies (Renwick et al. 2007, p. 531). Miller resiliency, and be a mortality factor Ngorongoro Crater, respectively, et al. (2014, pp. 495, 496) found based on data that indicate survival is resulted in unusually high mortality respiratory shedding of viable M. bovis shortened in infected lions, with death rates (Hunter et al. 2012, p. 2; Craft in living lions, meaning that lions could ranging between 2 and 5 years after 2008, p. 14; Munson et al. 2008, pp. 1, transmit bTB and serve as maintenance infection (Maas et al. 2012, p. 4212; 2; Cleaveland et al. 2007, pp. 613, 618; hosts. Renwick et al. 2007, p. 536; Keet, Roelke-Parker et al. 1996, pp. 441, 443). The social behavior of buffalo and unpublished data in Michel et al. 2006, These outbreaks coincided with climate lions allows M. bovis to spread to larger p. 93; Cleaveland et al. 2005, pp. 450, extremes that resulted in a higher areas and facilitates the transmission 451). In addition to clinical effects of number of Babesia, a tick-borne within and between prides. Drought bTB that may lead to mortality, this parasite, infections (Munson et al. 2008, conditions may also encourage the disease has also led to social changes pp. 2, 5). Babesia is common in lions, spread of this pathogen as herds must with lower lion survival and breeding but typically at low levels with no move into new areas in search of forage, success with more frequent male measurable impacts on their health potentially putting them in contact with coalition turnover and, consequently, (Craft 2008, p. 14; Munson et al. 2008, new, uninfected herds (Keet et al. 2009, higher infanticide (Keet, unpublished p. 3). However, droughts in 1993 and pp. 4, 6; Renwick et al. 2007, p. 533; data in Michel et al. 2006, p. 93). 2000 in Serengeti National Park/Maasai Michel et al. 2006, p. 93). In Kruger Research has shown adverse effects to Mara National Reserve and Ngorongoro National Park, bTB was introduced in lion individuals and subpopulations, Crater, respectively, led to large-scale the southeastern corner of the park but effects at the species population starvation and widespread die-offs of between 1950 and 1960. It gradually level are developing slowly (Michel et buffalo. This situation combined with made a northern progress and reached al. 2006, p. 97). Studies have shown that resumption of rains and fire suppression the park’s northern boundary in 2006. In impacts of bTB on lion numbers vary in Ngorongoro Crater favored 2009, the disease was found in buffalo between populations. For example, 30 propagation of ticks, vectors of Babesia, across the river boundary in Zimbabwe percent of the inbred populations in leading to unusually high tick burdens. (Keet et al. 2009, pp. 6, 11; Renwick et Hluhluwe-iMfolozi Park died due to a The compromised health of buffalo al. 2007, pp. 532, 533; Michel et al. combination of bTB and malnutrition allowed lions to feed on an inordinate 2006, pp. 92, 96, 98). A study from (Hunter et al. 2012, p. 3). However, number of tick-infested prey (Craft 2008, Kruger National Park indicated that bTB despite bTB infection and a high p. 14; Munson et al. 2008, pp. 2, 4, 5). spreads quickly through lion prevalence in prey species, the lion Exposure to either CDV or Babesia populations; in an area with high herd population in Kruger National Park has singly is not typically associated with a prevalence of M. bovis, 90 percent of remained stable (Ferreira and Funston compromise in health or an increase in lions became infected (Cleaveland et al. 2010, p. 201). mortality (Craft 2008, p. 14; Munson et 2005, p. 451). In time it will likely Epidemics of canine distemper virus al. 2008, pp. 1, 2, 3). However, the spread to Mozambique (Keet et al. 2009, (CDV) are known to have occurred in Babesia infections were exacerbated by p. 6). In Serengeti National Park, the Serengeti-Mara Ecosystem, an area the immunosuppressive effects of CDV infection may be widespread due to the that encompasses the Serengeti National and led to the unusually high mortality large, migratory wildebeest population Park, Ngorongoro Conservation Area, rates (Craft 2008, p. 14; Munson et al. that ranges throughout the Serengeti and Maasai Mara National Reserve 2008, p. 5). The Serengeti National Park/ ecosystem, including Maasai Mara (Craft 2008, pp. 13–14; Cleaveland et al. Maasai Mara National Reserve lion National Reserve (Cleaveland et al. 2007, pp. 613, 616, 618). CDV is a population lost 30 percent of its 2005, p. 450). Although an eradication common pathogen in the large population (approximately 1,000 lions), program has been implemented for population of domestic dogs (Canis but has recovered to its pre-epidemic cattle in South Africa, once an infection lupus familiaris) around the Serengeti- population levels (Craft 2008, pp. v, 14, is established in a free-ranging Mara Ecosystem, which are believed to 41; Munson et al. 2008, p. 1; Cleaveland maintenance host, like buffalo, it is be the source of CDV in lions et al. 2007, pp. 613, 617; Roelke-Parker unlikely to be eradicated (Keet et al. (Cleaveland et al. 2007, pp. 613, 617). et al. 1996, p. 444). Thirty-four percent 2009, p. 11; Renwick et al. 2007, pp. CDV is assumed to be transferred to of the Ngorongoro Crater lion 537, 538; Michel et al. 2006, p. 96). In lions by the sharing of food sources with population was killed, but frequent fact, modeling has predicted that spotted hyenas (Crocuta crocuta) or outbreaks of disease have prevented this prevalence could reach as high as 90 jackals (Canis spp.) that become population from recovering back to its percent over the next 25 years, with infected by consuming the infected carrying capacity (Craft 2008, p. 14; similar consequences for predators carcasses of domestic dogs (Craft et al. Munson et al. 2008, pp. 1, 2; Cleaveland (Renwick et al. 2007, p. 535). 2009, p. 1783; Craft 2008, p. 13). Viana et al. 2007, p. 617). The difference in Clinical signs of bTB in lions include et al. (2015, pp. 1466, 1467) recently recovery is likely due to the highly emaciation, respiratory complications, discovered that domestic dogs are not inbred nature of the Ngorongoro Crater swollen lymph nodes, draining sinuses, the sole source of CDV in the Serengeti, lion population, compared to the ataxia, and lameness (Keet et al. 2009, but rather there is likely a larger, Serengeti population, and its greater p. 13; Renwick et al. 2007, pp. 533, 534; multihost community of wildlife that susceptibility to parasitic and viral Cleaveland et al. 2005, p. 450), although contribute to outbreaks. Lions may also infections (Hunter et al. 2012, p. 2; some lions may be subclinically transmit CDV among themselves via Munson et al. 2008, p. 5; Brown et al. infected but remain asymptomatic until sharing food, fights, and mating (Craft et 1994, pp. 5953–5954). they experience another bTB infection, al. 2009, pp. 1778, 1783; Craft 2008, pp. Feline immunodeficiency virus (FIV) suffer from poor nutrition or advancing 13, 18, 71). is an endemic pathogen in many lion age, or become super-infected with CDV generally lacks clinical signs or populations of southern and eastern other diseases that may exacerbate the measurable mortality in lions, and most Africa (Maas et al. 2012, p. 4206; Adams

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et al. 2011, p. 173; Pecon-Slattery et al. The role of disease in determining effective immune response against an 2008, p. 2; Hofmann-Lehmann et al. survival and reproductive potential in emerging pathogen (O’Brien et al. 2006, 1996, pp. 555, 558; Brown et al. 1994, lions is almost completely unknown. It p. 255). For example, the inbred p. 5966). FIV is believed to have been is often difficult to determine whether populations in Hluhluwe-iMfolozi Park present in lions since the late Pliocene mortality was due to a single or lost 30 percent of lions due to a (O’Brien et al. 2012, p. 243; Troyer et al. combination of factors. Lions could be combination of bTB and malnutrition 2011, p. 2; Roelke et al. 2009, p. 3; infected with and become debilitated by (Hunter et al. 2012, p. 3). The Pecon-Slattery et al. 2008, p. 8). There a disease, but the cause of death could Ngorongoro Crater lions have not are 6 subtypes of FIV, A through F, each ultimately be due to other factors recovered to pre-outbreak numbers due with a distinct geographic area of (LionAid 2014a, pp. 4–5). Available to their inbred nature and greater endemnicity (Adams et al. 2011, p. 174; studies do not indicate that infection susceptibility to parasitic and viral Troyer et al. 2011, p. 2; Roelke et al. with a single disease is causing infections (Hunter et al. 2012, p. 2; 2009, p. 3; Pecon-Slattery et al. 2008, p. detrimental impacts to lions at the Munson et al. 2008, p. 5; Brown et al. 4; O’Brien et al. 2006, p. 262) and species level, although general body 1994, pp. 5953–5954). Additionally, differing levels of virulency (LionAid condition, health, and lifespan may be disease outbreaks can lead to extirpation 2014b, unpaginated). The social nature compromised and result in negative in small, isolated populations (Gilpin of lions allows for viral transmission impacts at the individual or population and Soule 1986 and Paul-Murphy et al. within and between prides through level. 1994 in Harvell et al. 2002). Although saliva when biting (Maas et al. 2012, p. Co-infections, however, could have we found no information indicating 4210; Pecon-Slattery et al. 2008, p. 5; synergistic effects that lead to greater presence of disease in the Indian Brown et al. 1994, p. 5953). Prevalence impacts on lions than a single infection. population, the small, isolated nature of FIV often approaches 100 percent of Lions impacted by the 1994 CDV makes the population more vulnerable adults in infected lion populations, outbreak in Serengeti National Park/ to disease outbreaks and could have a including the few remaining Maasai Mara National Reserve may have detrimental impact on the population populations in Botswana, South Africa, been more susceptible to CDV due to (Banerjee and Jhala 2012, p. 1427; and Tanzania, (LionAid 2014b, depleted immunity caused by FIV Meena 2010, p. 209; Johnsingh et al. unpaginated; O’Brien et al. 2012, p. 243; (O’Brien et al. 2006, p. 263). Troyer et 2007, p. 93). This principle also applies Troyer et al. 2011, p. 2; Roelke et al. al. (2011, pp. 5–6) found that survival to the small, isolated populations 2009, p. 3; O’Brien et al. 2006, p. 262; during the CDV/Babesia outbreak in throughout Africa. Hofmann-Lehmann et al. 1996, p. 559). Serengeti National Park/Maasai Mara Although disease is known in several National Reserve was significantly less populations, the impacts are known in FIV causes immune deficiencies that for lions infected with FIV A and/or C only a few populations where disease allow for opportunistic infections in the than FIV B. This finding suggests that has been frequently studied. Precise host (Roelke et al. 2009, p. 1; Brown et FIV A and C may predispose carriers to estimates of lions lost to disease are al. 1994, p. 5,953). With an impaired CDV pathogenesis and may increase the lacking, due to the difficulty in immune system, lions may not have an risk of mortality (O’Brien et al. 2012, p. detection. However, disease appears to appropriate and effective immune 243). Impacts of co-infections of FIV be a secondary factor influencing the response to various pathogens to which with FCV, FPV, FHV, and FCoV on decline of lions when co-infections they are consistently exposed (LionAid individual lions are negligible and do occur or when disease is combined with 2014a, p. 6). There may also be not endanger the lion population, at other factors, including environmental unrecognized immunological least in the absence of other aggravating changes, reduced prey density, and consequences (Roelke et al. 2006, p. cofactors (Hofmann-Lehmann et al. inbreeding depression. Diseases weaken 234) and adverse clinical and 1996, p. 561). individuals and allow them to succumb pathological outcomes (Roelke et al. Pathogen–pathogen interactions may to other diseases or factors. Although 2009, p. 1). Chronic effects of FIV are become more important when lions are disease does not appear to be a major important to long-term survival and under additional stress (e.g., increased driver in the status of the lion, differ according to subtype (Troyer et al. parasite load or low prey density) (Maas populations can suffer significant losses; 2011, p. 6). Studies have indicated that et al. 2012, p. 4212). Certain some may recover to pre-outbreak lions may exhibit signs of opportunistic environmental conditions may levels, others may not. Given the small infection associated with AIDS, such as exacerbate the effects of an otherwise and declining lion populations that swollen lymph nodes, gingivitis, tongue innocuous infection. For example, as remain, any loss of individuals from the papillomas, dehydration, poor coat discussed above, CDV and Babesia populations could be detrimental. condition, and abnormal red blood cell infections generally have no measurable The risk of disease may increase with parameters, and in some cases death impacts on lion health, but climatic time due to loss of genetic variation (Troyer et al. 2011, p. 2; Roelke et al. conditions increased exposure of lions associated with continued 2009, pp. 2, 3–6). Lions in Botswana to Babesia infections, which were fragmentation of populations, whether and Tanzania have demonstrated exacerbated by the immunosuppressive by habitat loss or fencing of habitat, and multiple clinical features of chronic effects of CDV and led to unusually high increased proximity to humans and immune depletion similar to HIV and mortality rates (Craft 2008, p. 14; domestic livestock that may expose domestic cat AIDS (Troyer et al. 2011, Munson et al. 2008, p. 5). Some lions lions to new diseases (IUCN 2006b, pp. pp. 2–3). However, there is no evidence infected with bTB may remain 19, 26). Additionally, changes in climate that FIV itself poses a threat to wild asymptomatic until conditions change may increase disease outbreaks in prey populations (Frank et al. 2006, p. 1); FIV and they suffer from poor nutrition due species, as well as lions (See Climate does not appear to be impacting lions in to low prey density, advancing age, or Change). Climate change could Kruger National Park (Maas et al. 2012, become super-infected with other potentially increase the likelihood of p. 4212), and no evidence of AIDS-like diseases that may exacerbate the lethal co-infections (The Heinz Center illnesses or decreased lifespan has been infection (Renwick et al. 2007, p. 533). 2012, p. 12), similar to the co-infections found in FIV lion populations in the Species with reduced genetic of CDV and Babesia in Serengeti Serengeti (O’Brien et al. 2006, p. 263). variation may be less able to mount an National Park/Maasai Mara National

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Reserve and Ngorongoro Crater lions by Riggio et al. (2013) may be smaller 1994, p. 5748). Loss of fecundity leads following drought events. for P. l. leo as pride sizes are generally to a decrease in population size, fewer smaller than those for P. l. melanochaita prides in a population, and increased Deleterious Effects Due to Small (Riggio et al. 2013, p. 32; Meena 2009, inbreeding which contributes to a Population Sizes p. 7; Nowell and Jackson 1996, p. 37). decline in the population and increases The risk of extinction is related to the Male dispersal also plays an the risk of extinction (Bjo¨rklund 2003, moment when a declining population important role in determining the level p. 521). Additionally, lack of genetic becomes a small population and is often of inbreeding in lion populations. Even variation can impact the ability of lions estimated using minimum viable if only a fraction of males do not to withstand stochastic events. For population (MVP) sizes (Traill et al. disperse, inbreeding rapidly increases example, the inbred populations in 2010, p. 28). The viability of a lion with each generation (approximately 5 Hluhluwe-iMfolozi Park were unable to population is complex, but it partly years) (Bjo¨rklund 2003, pp. 518, 520). mount an effective immune response depends on the number of prides and Even when migration rates of males is and lost 30 percent of lions due to a ability of males to disperse and interact as high as 95 or 99 percent, the combination of bTB and malnutrition with other prides, which affects likelihood of inbreeding is clearly (Hunter et al. 2012, p. 3). Additionally, exchange of genetic material (Bjo¨rklund higher than if 100 percent of males the lions of Ngorongoro Crater never 2003, p. 518). Without genetic exchange, disperse. Using a 95 percent dispersal recovered to pre-outbreak numbers due or variation, individual fitness is rate, the probability of inbreeding its inbred nature and greater reduced and species are less able to reached 57 percent and 20 percent for susceptibility to parasitic and viral adapt to environmental changes and 10 and 100 prides within 30 generations infections (Hunter et al. 2012, p. 2; stress, increasing the risk of extinction (150 years) (Bjo¨rklund 2003, pp. 518– Munson et al. 2008, p. 5; Brown et al. (Bijlsma and Loeschcke 2012, pp. 117, 519). One example is the lion 1994, pp. 5953–5954). Reductions in 119; Segelbacher et al. 2010, p. 2; Traill population in Ngorongoro Crater. New genetic variations may also limit the et al. 2010, p. 31; Bjo¨rklund 2003, p. males rarely migrate into the population lion’s ability to evolve responses to 515). due to physical barriers, and inbreeding climate change (The Heinz Center 2012, Bjo¨rklund (2003, p. 520) found that has been shown to occur (Packer et al. p. 12). the most important determining factors 1991b in Bjo¨rklund 2003, p. 521). The The lion population in India is one of for the level of inbreeding in lions is the fewer number of males present to the few populations that are increasing number of prides and male dispersal. contribute genes to the next generation, (Bauer et al. 2015a, unpaginated; BBC The MVP for lions has not been formally the more inbred the population will be 2015, unpaginated; The Guardian 2015, established and agreed upon by species (Riggio et al. 2013, p. 32). Therefore, not unpaginated; Banerjee and Jhala 2012, experts (Riggio et al. 2011, p. 5; CITES only does dispersal impact inbreeding, p. 1427) and could be considered a 2004a, p. 2; Bjo¨rklund 2003, p. 521); so does the loss of male lions due to stronghold according to the criteria set however, it has been suggested that to excessive trophy hunting and by Riggio et al. (2013, p. 22). Despite conserve genetic diversity, populations infanticide (see Trophy Hunting). being genetically less diverse, Banerjee of at least 50 prides, but preferably 100 Because the number of prides and and Jhala (2012, pp. 1424–1425) found prides (250 to 500 individuals), with no male dispersal are the most important no evidence of depressed demographic limits to dispersal, are necessary (Bauer factors for maintaining viability, parameters in the lions of India. et al. 2008 in Riggio et al. 2013, p. 32; sufficient areas are needed to support at However, intense management, Bjo¨rklund 2003, pp. 515, 518). least 50 prides, but preferably 100 including healthcare interventions, may Bjo¨rklund (2003, p. 518) found that prides, and allow unrestricted male interfere with natural selection inbreeding decreased rapidly with the dispersal (Bjo¨rklund 2003, p. 521). processes by ensuring the survival of number of prides. For example, if there Unfortunately, few lion populations unfit lions which facilitates the are less than 10 prides the likelihood of meet these criteria as almost all lion propagation of deleterious genes in the genetic effects due to inbreeding populations in Africa that historically population (Banerjee and Jahala 2012, p. increased from 0 in the beginning to 26– exceeded 500 individuals are declining, 1427). This population is also running 45 percent after 30 generations, whereas and few protected areas are large out of area to expand. Being a small, if 100 prides are present, the likelihood enough to support viable populations isolated population and less genetically is only 5 percent assuming no migration (Bauer et al. 2015a, unpaginated; Bauer diverse, it is more vulnerable to the loss into the population (Bjo¨rklund 2003, p. et al. 2015b, p. 1; Bauer et al. 2008, of any individuals due to environmental 515). Additionally, it appears that unpaginated; Riggio 2011, p. 5; Hazzah and stochastic events, and more prone inbreeding rapidly increases when the 2006, p. 2; Bauer and Van Der Merwe to local extinction events (Banerjee and number of prides falls below 50 2004, pp. 28–30; Bjo¨rklund 2003, p. Jhala 2012, p. 1428; Meena 2010, p. 209; (Bjo¨rklund 2003, p. 518, Figure 2). 521). Even within large areas, Johnsingh et al. 2007, p. 93; Thuiller et Riggio et al. (2013, pp. 20, 22) used the inbreeding will increase if dispersal is al. 2006, pp. 434–435). threshold described by Bjo¨rklund (2003) limited, (Bjo¨rklund 2003, pp. 521–522). The establishment of another free- to define, in part, lion strongholds. Furthermore, research indicates that ranging population geographically Stronghold populations of lions were there is a general lack of gene flow in separate from Gir would reduce the risk considered to be those that meet the most lion conservation units (Dubach et of extinction of this population due to necessary requirements for long-term al. 2013, pp. 749, 750; Bertola et al. stochastic events (e.g., disease outbreaks viability and were defined, in part, as 2011, p. 1364; Chardonnet et al. 2009, or floods). In the early 1990s, a second containing at least 500 individuals (100 p. 54). population was proposed at Kuno prides). Potential strongholds were Small populations (e.g., fewer than 50 Wildlife Sanctuary in Madhya Pradesh described, broadly, as areas where lions) can persist in the wild for some State (Johnsingh et al. 2007, p. 93). immediate interventions might create a time; however, the lack of dispersal and However, the Government of Gujarat has viable population and were defined, in genetic variation can negatively impact refused to allow any lions from Gir to part, as populations that contained at the reproductive fitness of lions in these be transferred to the Kuno Wildlife least 250 lions. However, the threshold populations and local extirpation is Sanctuary, despite a ruling by India’s described by Bjo¨rklund (2003) and used likely (Traill et al. 2010, p. 30; O’Brien Supreme Court (The Economic Times

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2015, unpaginated; Duerr 2014, management budgets. Cost estimates for in western Africa, compared to central, unpaginated; Meena 2014, p. 29). maintaining lion populations in eastern, and southern Africa. Countries protected areas range from an annual in the former or current western Africa Regulatory Mechanisms budget of $500 USD per km2 in smaller lion range are among the 50 poorest Regulatory mechanisms in place to fenced reserves to $2,000 USD per km2 countries in the world, and six are provide protections to African lions for unfenced reserves (Packer et al. classified as least developed countries. vary substantially throughout Africa. 2013, p. 640). This includes but is not These countries will likely be unable to The lion species (Panthera leo) is listed limited to costs associated with generate the resources required to in Appendix II of CITES; however, the permanent and temporary staff, fencing secure their remaining lion populations former Asiatic lion (P. l. persica) is installation and maintenance (fences (Henschel et al. 2014, pp. 7–8). listed in Appendix I. With the exception can cost $3,000 USD per km to install), Investment from the international of South Sudan, all of the lion range infrastructure maintenance, anti- community is needed to increase states are Parties to CITES. According to poaching activities such as surveillance management effectiveness of these the draft CITES Periodic Review of the and snare/trap removal, wildlife protected areas (Henschel et al. 2015, Status of African Lions (CITES 2014, pp. restocking fees (both for lions killed by unpaginated). 14–15) outside of CITES, lions have no illegal poaching/snares as well as other In India, most lions occur within five legal protections in four countries: trophy species killed by lions on the designated protected areas: Gir National Burundi, Guinea Bissau, Lesotho, and reserves), community outreach, and Park and Gir Wildlife Sanctuary (Gir Swaziland. However, CITES 2014 (p. 15) compensation for loss of livestock in Protected Area) and Pania, Mitiyala, and states that most of the southern and surrounding communities. However, Girnar sanctuaries (Bauer et al. 2015a, eastern lion range states have regulatory many management areas lack adequate unpaginated; Banerjee and Jhala 2012, mechanisms in place to protect lions. funding (Packer et al. 2013, p. 640; p. 1421; Singh and Gibson 2011, p. We found that most of the range states Groom 2013, pp. 4–5; Barnett and 1754; Jhala et al. 2009, pp. 3384, 3385; have national environmental legislation Patterson 2005, p. 82). Nowell and Jackson 1996, p. 38). Under to establish national parks and Of 12 protected areas assessed in India’s Wild Life Protection Act of 1972 conservation areas, and to conserve and western Africa, 6 had no budget for (Act No. 53 of 1972; Chapter IV, sections regulate the take, hunting, and trade of management activities or the budget was 27, 28, 33, 35), entry into protected wildlife, including parts and products, too low to conserve lion populations; areas is regulated and certain activities but could find no legislation specific to nine reported having either no law are controlled and managed, including lions, or to the main threats affecting enforcement activity or major security of wild animals and grazing of lions: habitat loss, human–lion conflict, deficiencies in staff and resources to livestock. In 2012, India’s Ministry of and loss of prey base (Ecolex 1 conduct patrols. In Comoe´ National Environment and Forests (2012, p. 22) information last accessed November 6, Park, the staff was found to be too small declared the area 5 km from the 2015). for the size of the park (Henschel et al. boundary of Girnar Wildlife Sanctuary National and international 2014, p. 7). Protected areas in Guinea an Eco-sensitive Zone for the long-term conservation strategies rely on protected are essentially parks on paper only. protection and conservation of the lion. areas to protect natural resources from They have no staff, management plan, or This designation prohibits certain negative impacts of human populations operating budget (Brugie´re 2012 in activities within the designated zone, (Craigie et al. 2010, p. 2221). The lion Henschel et al. 2014, p. 7). Although the such as mining, unregulated tourism, is largely limited to protected areas; WAP complex has received high scores polluting industries, and unregulated therefore, effective management is for management effectiveness, the felling of trees. crucial to the survival of the species. presence of 50,000 head of cattle inside Because of the protections afforded by However, weak management of W National Park indicates weak the Government of Gujarat, threats that protected areas has been documented management. Livestock are rare in Arly- contributed to the decline of this across its range, especially in western Pendjari, and lion density is higher; a population have been ameliorated and Africa where most protected areas are higher management budget allocation is most threats faced by lions are not an experiencing severe management suspected to be the cause of the immediate threat. Protections ensure deficiencies (Henschel et al. 2015, observed differences (Henschel et al. food security, water availability, habitat unpaginated; Henschel et al. 2014, pp. 2014, pp. 5–6). Across the lion’s range, suitability, and safety for these lions 5, 7; Brugie´re 2012 in Henschel et al. Africa’s protected areas have generally (Meena 2014, p. 26). However, because 2014, p. 7; Craigie et al. 2010, entire). failed to mitigate threats to large this population is small and isolated, it The WAP complex in western Africa mammal populations, including the lion is vulnerable to extinction from had received high scores for and its prey (Craigie et al. 2010, entire). stochastic events. Although a second management effectiveness (Henschel et Poor management leads to many of location has been proposed to establish al. 2015, p. 7). the threats that lions face, including another free-ranging population Effective management requires encroachment by pastoralists, increased geographically separate from Gir to adequate funding, resources, and staff. poaching pressure, collapse of prey reduce the risk of extinction of this Packer et al. (2013a, pp. 638–639) found populations, and persecution by population, translocation of lions from that lion densities were highest in pastoralists (Brugie´re et al. 2015, pp. Gujarat are still pending (see Deleterious protected areas with the highest 519–520; Henschel et al. 2015, Effects Due to Small Population Sizes). unpaginated; Henschel et al. 2014, pp. 1 ECOLEX is a comprehensive database on 5, 7; Henschel et al. 2010, p. 38). Climate Change environmental law, maintained by the International Consideration of ongoing and Union for Conservation of Nature (IUCN), the Therefore, it can be said that United Nations Environment Programme (UNEP), management of protected areas that still projected climate change is a and the Food and Agriculture Organization of the harbor lions is inadequate to address the component of our analysis under the United Nations (FAO). Our search terms used with threats impacting lions, especially those Act. The term ‘‘climate change’’ refers to respect to wildlife laws were ‘‘African lion,’’ ‘‘Asiatic lion,’’ ‘‘Panthera leo leo,’’ ‘‘Panthera leo in western Africa (Henschel 2015, a change in the mean, variability, or persica,’’ and ‘‘country,’’ e.g., ‘‘Angola,’’ ‘‘Benin,’’ unpaginated). Overall, investment in seasonality of climate variables over etc. Information accessed at http://ecolex.org. conservation activities is extremely low time periods of decades or hundreds of

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years (Intergovernmental Panel on in Africa and Asia and rainfall will events related to monsoons are very Climate Change (IPCC) 2013, p. 1255). continue to decrease in Africa but likely (Hijioka et al. 2014a, p. 1334; Climate change models, like all other increase in India, although regional Gosling et al. 2011, pp. 123–124). scientific models, produce projections variations exist (Hijioka et al. 2014a, p. Increases in precipitation are expected that have some uncertainty because of 1334; Peterson et al. 2014, p. 562; by the 2030s and all regions of India are the assumptions used, the data Gosling et al. 2011, pp. 64–65). expected to experience between 10 and available, and the specific model Warming in Africa is expected to be 30 percent increases in magnitude of features. The science supporting climate greater than the global annual mean pluvial flooding (flooding derived model projections as well as models warming throughout the continent and directly from heavy rainfall and results assessing their impacts on species and all seasons (Chidumayo et al. 2011, p. in overland flow) and an average across habitats will continue to be refined as 22). Future projections expect the India of approximately 50 percent more information becomes available. average temperature in Africa to be greater risk of fluvial flooding (floods as higher by 1.5–3 °C by 2050 (Niang et al. a result of river flows exceeding river Temperature and Precipitation Trends 2014, p. 1206; Carr et al. 2013, p. 16; channel capacity, breaking through Within the past 50–100 years, the UENP 2007, p. 2), while temperatures in riverbanks, and inundating the surface temperature in Africa and Asia Gujarat are expected to increase floodplain) (Gosling et al. 2011, pp. 122, has increased (Hijioka et al. 2014a, p. between 3.0 and 3.5 °C by 2100 (Gosling 123, 126, 130). Gosling et al. (2011, pp. 1333; Niang et al. 2014, p. 1206). Across et al. 2011, pp. 64–65). 65–66) predict increases in average Africa, surface temperature has Annual precipitation shows greater annual rainfall of up to 20 percent in increased by 0.5 °C over the past regional variations, although predictions Gujarat by 2100. century (Niang et al. 2014, p. 1206), of precipitation contain high levels of although there are regional differences. uncertainty. Generally speaking, both Impacts of Climate Change For example, decadal warming rates in Africa and Asia are expected to Climate change is likely to become a South Africa have ranged from 0.1 °C to experience harsher drought and stronger main driver of change in large mammal 0.3 °C (Chidumayo et al. 2011, p. 18) floods during the wet season (Hijioka et populations in the future (Scholte 2011, and 0.23 °C in Tanzania (Carr et al. al. 2014a, p. 1334; Carr et al. 2013, p. p. 7). In the mid-Holocene, 2013, p. 16). The mean annual 12). Precipitation has been projected to responded rapidly to climate change temperature in Burundi has increased decline in western, central, and with a series of local extinctions and by 0.7–0.9 °C since the 1930s, while the southern Africa. The areas of southern near-extinctions, driving a decrease in mean annual temperature in Uganda has Africa expected to experience a decline species richness, and a dramatic increased by 1.3 °C since 1960 (Carr et in precipitation is projected to expand increase in xerophytic taxa (Grayson al. 2013, p. 16). In India, annual mean during the second half of the 21st 2000 and Graham 1992 in Thuiller et al. temperatures increased by 0.56 °C century (Niang et al. 2014, p. 1210; 2006, p. 425). It is likely that many during the 20th century (Hijioka et al. Hijioka et al. 2014a, p. 1333; Carr et al. species and ecosystems will endure 2014a, p. 133; Hijioka et al. 2014b, p. 2013, pp. 12, 14; The Heinz Center similar impacts in response to predicted SM24–2). 2012, p. 13). climate change in the 21st century, Across Africa, trends in annual In contrast, eastern Africa and which will act synergistically with the precipitation indicate a small but northern India are expected to predicted increase in anthropogenic statistically significant decline in experience an increase in mean annual pressures (Fischlin et al. 2007, in Carr rainfall (Niang et al. 2014, p. 1209; precipitation (Niang et al. 2010, p. 1210; et al. 2013, p. 10; Thuiller et al. 2006, Chidumayo et al. 2011, p. 20). Eastern Hijioka et al. 2014a, p. 1334; Carr et al. p. 425). For lion, impacts described Africa has experienced an increase in 2013, pp. 12, 14; Gosling et al. 2011, p. above from existing and predicted extreme precipitation changes, with 65). Some General Circulation Models anthropogenic pressures on the species increasingly frequent droughts followed predict that, by the end of the 21st and its habitat are likely to be by increasingly intense heavy rainfall, century, eastern Africa will have a exacerbated by climate change. The for the last 30 to 60 years; however, wetter climate with more, intense wet general warming and drying trend overall levels of precipitation have been seasons and less severe droughts from projected for Africa could further reduce declining. The intense rainfall events October to December and March lion range, numbers, and prey base. have caused more frequent flooding and through May, a reverse in observed Lions may also have to travel greater soil erosion and degradation (Niang et trends described above. Other models distances to find food or shift their diet al. 2014, pp. 1209, 1211; Carr et al. suggest drying in most parts of Uganda, to livestock, increasing conflict with 2013, p.16). Attri and Tyagi (2010 in Kenya, and South Sudan in August and humans and the risk of retaliatory Hijioka et al. 2014b, p. SM24–3) report September by the end of the 21st killings (Peterson et al. 2014, pp. 562– no significant national trends in century (Niang et al. 2014, p. 1210). Carr 563; Tuqa et al. 2014, p. 8; Tumenta et precipitation for India, although there et al. (2013, p. 15) state that levels of al. 2013, p. 240). Additionally, changes has been a decrease in the number of increased precipitation predicted for the in climate may increase the number and monsoon depressions and an increase in Albertine Rift, located mainly within intensity of disease outbreaks in lions the number of monsoon break days, the eastern African region, are not and its prey (Peterson et al. 2014, pp. which is consistent with an overall predicted to be sufficient to counter the 562–563; The Heinz Center 2012, p. 12; decrease in seasonal mean rainfall effects of warming temperatures; Baylis 2006, p. 4). (Hijioka et al. 2014a, p. 1333). therefore, an overall drying effect is Peterson et al. (2014, pp. 555, 561– Throughout the 20th century, droughts likely to occur, which will be more 562) evaluated the magnitude of were frequent in the Gir area. However, pronounced between February and May. potential changes in lion distribution in in the last two decades average rainfall They also state that November and Africa under different climate change has increased due to increased western December will experience the largest scenarios between the years 2040 and monsoons (Singh and Gibson 2011, p. increases in precipitation. 2070. They found little optimism for the 1756). In South Asia, including India, future future of lions. No broad new areas will Overall, projections indicate declines in the number of rainy days become suitable for lion. Southern temperatures will continue to increase and increases in extreme precipitation Africa, where the broadest areas of

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suitable conditions occur, is projected to will increase this pressure as land 68). Additionally, Van Vuuren et al. become less suitable because of climate becomes more arid and food security (2005 in Celesia et al. 2009, p. 68) found change. Specifically, park areas, concerns are exacerbated (Carr et al. in a study of Kgalagadi Transfrontier including the ‘‘Etosha Pan, Lake 2013, p. 20). Impacts to the socio- Park that adult and cub mortality Opnono, Cuvelai Drainage, Kalahari economic and physical well-being of reached 70 to 90 percent in poor years , and Kgalagadi Transfrontier humans will cause adaptive responses, (defined as years in which average Park areas’’ are projected to decline eliciting changes in the way much of the annual rainfall in the previous 2 years substantially in suitability for lions. A land is used, including further was less than 165 mm). Mortality broad swath of potential distributional encroachment of urban environments decreased to 10 to 40 percent in good area in western Africa is projected to and agricultural land into existing years (years in which average annual become ‘‘distinctly less suitable or even natural habitats (Carr et al. 2013, pp. 10, rainfall in the previous 2 years was uninhabitable.’’ A decrease in the lion’s 19), including protected areas where greater than or equal to 237 mm). These range could mean that stochastic events lions occur. Additionally, land impacts on demography result in impact a larger portion of the whole conversion restructures the landscape reduced numbers of lions and pride species, especially when the species and and may disrupt prey migrations that sizes (Celesia et al. 2009, p. 68). Given its habitat are fragmented (Thuiller et al. are induced by climate change (Thuiller the predicted warming and drying trend 2006, p. 434). Additionally, reductions et al. 2006, p. 425), decreasing or for the 21st century, additional lions in populations and geographic range altering prey available to the lion. could be lost and pride sizes reduced. may limit the lion’s ability to respond Although lions occur in a variety of Furthermore, loss of these lions reduces to climate change (The Heinz Center temperature and precipitation regimes, reproductive potential and recruitment, 2012, p. 12). However, climate change suggesting the species may be tolerant of further contributing to the decline of effects on potential lion distribution are some climatic changes (The Heinz existing populations. The loss of lions projected to be more neutral in eastern Center 2012, p. 13), lions appear to could also mean the loss of genetic Africa than across the entire range. thrive under specific climate parameters variation. Combined with declining Reserves in this region are more likely (Leighton-Jones 2004 in Celesia et al. populations, the risk of inbreeding and to sustain lion populations under 2009, p. 63) and abundance is associated complications could climate change scenarios (Peterson et al. significantly determined by temperature increase. 2014, pp. 555, 561–562). and rainfall (Celesia et al. 2009, pp. 67, Drought conditions can also In India, an increase in average 68). Large felids, including lions, occur contribute to reduced prey availability rainfall in the past two decades has in biomes with an average annual by altering the timing of migration resulted in the conversion of dry temperature of 13 °C or higher; lion (Peterson et al. 2014, p. 562). For savanna to forestland (Hijioka et al. demography is best when mean annual migratory species such as the wildebeest 2014a, p. 1333; Singh and Gibson 2011, temperatures are 16–18 °C (Celesia et al. or zebra, an earlier and more frequent p. 1756). However, the lion population 2009, p. 68). Lion density is influenced onset of the dry season may lead to the in India has shown to be able to use by multiple natural ecological factors species undertaking more migrations, both forestlands and savannas (Singh including herbivore biomass, annual which can lead to increases in mortality and Gibson 2010, p. 1753). Therefore, mean rainfall, soil nutrients, annual and disruption of seasonal hunting this type of habitat conversion due to mean temperature, and interactive patterns of lion (The Heinz Center 2012, changes in climate may not be as effects between rainfall and soil p. 42). Climate change may already be detrimental to lions in India population. nutrients (Celesia et al. 2009, pp. 67, having an impact on the wildebeest as However, increased risks of flooding 69). These factors explain regional Dobson (2009, as cited in Chidumayo et could pose problems for lions. variations in lion densities, where low al. 2011, p. 144) found that, due to the Following a recent flood in Gujarat, nine densities are found in desert or semi- wet season slowly getting drier and the lions drowned in a stream that flows desert ecosystems and higher densities dry season getting wetter, the species is alongside Gir Wildlife Santuary. in moist savannas (Celesia et al. 2009, migrating 2 months earlier than usual, Additionally, lions could face serious p. 67). Lion densities decrease with throwing off timing of migrations and threats following flood events, such as increasing mean temperature and conception times that are set by lunar an outbreak of a disease epidemic (The decreasing rainfall. Therefore, lion cycles. If the wet season rains are Economic Times 2015, unpaginated). density, or carrying capacity of diminishing there will be a reduction in This population of lions is small, protected areas, in sub-Saharan Africa is high-quality forage needed to support isolated, and less genetically diverse; likely to decline with climate warming lactation. This reduction has a therefore, it is more vulnerable to and drying (Chidumayo et al. 2011, p. detrimental effect not only on the stochastic events such as disease 144). survival of the calf but also for the outbreaks and flooding and more prone Lion demography is also influenced population as a whole (Dobson 2009, as to local extinction events (Banerjee and by environmental factors. Many cited in Chidumayo et al. 2011, pp. 144– Jhala 2012, p. 1428; Meena 2010, p. 209; variables are associated with aspects of 145). Johnsingh et al. 2007, p. 93). demography, but the strongest Climate conditions also influence Current lion habitat and suitable associations are with rainfall, prey abundance. In Kruger Park, South habitat predicted to remain under temperature, and landscape features Africa, almost all species are climate change scenarios will be under (e.g., elevation, slope, direction of slope, extremely sensitive to lack of rainfall increasing pressure due to land and compound topographic index) during the dry season, which is conversions to meet the needs of the (Celesia et al. 2009, pp. 63, 68). Impacts predicted to increase in the future. This growing human population. As stated to lion demography have been noted factor may be important to retain green earlier, and supported by Carr et al. with the longer dry spells occurring. For forage during a period when the risk of (2013, p. 20), demand for agricultural example, when prey become scarce at malnutrition is higher (Thuiller et al. land is likely to increase to meet the the end of the dry season, subadult 2006, p. 432). Similarly, reproduction in needs of the growing human population, females may be forced out of prides. Cape buffalo is strongly related to putting pressure on natural landscapes. Furthermore, older lions and cubs may season. Changes in the timing, Projected changes in Africa’s climate die of starvation (Celesia et al. 2009, p. frequency, or intensity of seasonal rains

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could negatively affect reproduction. southern Africa (Niang et al. 2014, p. hosts that may further influence the This species is also sensitive to rainfall 1209; Hijioka et al. 2014a, p. 1333; Carr spread of diseases (Baylis 2006, pp. 9, due to its high water consumption rate et al. 2013, p. 14; Chidumayo et al. 11) and increase risks of extinctions (up to 30–40 liters per animal per day) 2011, p. 21). Drought in the western and (Thuiller et al. 2006, p. 435). (Du Troit 2005, as cited in The Heinz central African regions is expected to Additionally, rainfall conditions also Center 2012, p. 15; Whyte et al. 1995, increase by a rate of 5–8 percent by 2080 affect the susceptibility of animals to pp. 84–85). Variation in the buffalo (UNEP 2007, p. 2). Although drier disease outbreaks (Thuiller et al. 2006, population then is tied to rainfall conditions might initially lead to the p. 435). Munson et al. (2008) concluded conditions year-to-year. Funston and lion home range shrinking as prey that severe climate change could Mills (2006, p. 20) observed that the congregate around remaining water synchronize temporal and spatial buffalo population increases only sources (Sogbohoussou 2011, p. 133), convergence of multiple infectious during periods of average to above- Tuqa et al. (2014, p. 8) found that lion agents, triggering epidemics with greater average rainfall, which means that home ranges expand in the time after a mortality than infections from a single climate projections for a drier Africa drought. The reason for this expansion pathogen. will have detrimental impacts on the may be that, as prey populations around Conservation Measures in Place To buffalo population. Lions are water sources are depleted, the lion has Protect Lions opportunistic predators that feed on a to travel greater distances to find prey. variety of prey. This flexibility in prey In addition, researchers found that lions There has been awareness for several may aid lions in exhibiting some move beyond reserve boundaries and years that conservation strategies need resiliency to changes in prey into communal ranches where there will to be implemented for the lion due to populations (The Heinz Center 2012, p. be greater conflict with humans (Tuqa et the apparent decrease in its population 12). However, as discussed under Loss al. 2014, p. 9). It is likely that lions prey numbers (Hamunyela et al. 2013, p. 1; of Prey Base and Human–Lion Conflict, on livestock, which will intensify Henschel et al. 2010, p. 34; Gebresenbet the loss of prey species can result in human–lion conflict. To compound the et al. 2009, p. 5; IUCN 2006a, b, entire). lions shifting their diet towards issue, pastoralists in sub-Saharan Africa Prior to 2006, institutional livestock which may increase retaliatory will often lead their herds into protected inconsistencies throughout the lion’s killings by humans (Bauer and Kari areas where lions occur during a African range resulted in poor lion 2001, as cited in Tumenta et al. 2013, drought in search of water, which conservation policies and little to no p. 241; Whyte et al. 1995, p. 85). increases the risk of lion predation enforcement of existing laws (IUCN Variation in lion home ranges may (Tumenta et al. 2013, p. 240). 2006b, p. 18). As mentioned, in 2005 have an impact on the frequency of When lion prey on livestock, they and 2006, nongovernmental human–lion conflict especially in primarily focus on cattle (Patterson et al. organizations (NGOs) and several situations where lion home ranges 2004, p. 510). Out of all livestock that governments at various levels organized expand into areas inhabited by humans are domesticated in Africa, cattle have two regional lion conservation (Peterson et al. 2014, p. 562). The the highest monetary value, which workshops. Species specialists, wildlife interplay between the types of climate, means the loss of cattle to lion predation managers, and government officials the density of prey, and seasonal will have the most adverse effect on attended these regional workshops in variation in temperature and pastoralists (Tumenta et al. 2013, p. order to provide range country precipitation all affect lion home range. 240). Additionally, droughts affect the governments with frameworks for Areas with a more arid climate and survival of livestock (Peterson et al. developing their own national action small prey density are associated with 2014, p. 562). A study of the drought plans for the conservation of lions. Over larger home ranges, while temperate or that occurred in Kenya in 2008–2009 50 lion specialists, representing all lion tropical regions with higher prey found that mortality rates among the range countries, participated in these density are associated with smaller cattle population varied between 57 and workshops (Henschel et al. 2010, p. 34). home ranges. In addition, prey living in 64 percent in six districts (Dolrenry During the workshops, lion experts an arid climate tend to disperse, while 2013, p. 47; Zwaagstra et al. 2010, p. collectively assessed what they believed prey in a wetter climate are more 21). Such high mortality may make to be the then-current status of African concentrated, leading to a larger and pastoralists less tolerant of lion lions based on a variety of information, smaller home range, respectively (Tuqa predation and may increase the and subsequently identified 86 African et al. 2014, p. 2; Celesia et al. 2010, pp. frequency of retaliatory killings LCUs. This information was then used 63, 67; Sogbohossou 2011, p. 17; (Peterson et al. 2014, p. 562). as a framework to identify lion areas, Loveridge et al. 2009, p. 953). In Climate change may increase the strongholds, and potential strongholds southern Africa, where most of the lion number and intensity of disease by Riggio et al. (2013, p. 32). populations are enclosed (fenced), outbreaks in lion prey species, as well Many African countries with very variation in the species’ home range as lions (The Heinz Center 2012, p. 12; small lion populations have developed may be more limited. Lion home ranges Baylis 2006, p. 4). Diseases can be or updated their conservation plans for are also influenced by the season with directly and indirectly affected by the lion. Some of these include Benin, ranges being smaller during the dry climate change by impacting Cameroon, Uganda, and Malawi. Some season and larger during the wet season. distribution, the timing of outbreaks, range countries participate in During the dry season, prey congregate and the intensity of outbreaks (Baylis transboundary conservation projects around the few remaining water 2006, p. 4). Higher temperatures may and are collaborating on transboundary sources, concentrating prey species in a increase the rates of development of lion conservation initiatives for shared smaller area, shrinking the home range pathogens and parasites, shorten lion populations. Most range countries needed by the lion to find food. generation times, and increase the have a national lion action plan or Conversely, home ranges expand during number of generations per year, strategies in place, particularly if there the wet season due to prey dispersal increasing the population (Baylis 2006, are economic incentives for them to (Tuqa et al. 2014, p. 8). p. 8; Thuiller et al. 2006, p. 435). have viable lion populations (Groom Climate projections point toward a Temperatures can have impacts on 2013, p. 4; Namibia 2013, pp. 11–12; drier climate for western, central, and vectors (e.g., ticks and mosquitoes) and Zambia Wildlife Authority 2012, p.3;

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LionAid 2011, pp. 1–2; Mesochina et al. habitat across national boundaries. Trust 2011, p. 1). The corridor also 2010a, pp. 40–49; Mesochina et al. Corridors are being restored, fences are allows other wildlife such as lions to 2010b, pp. 33–38; Government of being removed, and protected areas are disperse through habitat that otherwise Tanzania 2010, pp. 3–17; Begg and Begg being connected. Restoration of these would have been unfavorable for 2010, entire). Range states have also corridors allows wildlife to travel wildlife to travel through (Mount Kenya implemented a number of conservation between areas of suitable habitat (Jones Trust 2011, p. 1). It was an expensive strategies designed to conserve habitat, et al. 2012, pp. 469–470). In some areas, project, but the effort appears to have reduce human–lion conflict, and fences have been constructed to protect served its purpose: Elephants are using preserve the lion’s prey-base. grazing resources for domestic livestock the corridor on a regular basis as well as to provide barriers to disease (particularly an underpass under a Conservation Measures To Stem Habitat (Gadd 2012, pp. 153, 176). One aspect highway), and humans are reporting less Loss of these fences is that they separate human–wildlife conflict (Mount Kenya Habitat loss represents one of the lions from their prey. In southern Trust 2011, p. 1). main threats facing lions in Africa Africa, fences are being taken down to However, connectivity alone does not (Bauer et al. 2008, unpaginated). increase the size of connected habitat ensure the dispersal of animals (Roever Attempts by range countries to address and link it to reserves and national et al. 2013, pp. 19–21). The Tanzania this decline in habitat are manifested in parks (IUCN 2009, p. 101; IUCN 2008, Wildlife Research Institute (TAWIRI) is a number of ways, such as the creation various). The Limpopo Transfrontier an organization under Tanzania’s of protected areas and the establishment Park is another example of where this Ministry of Natural Resources and of wildlife corridors to connect practice is being implemented Tourism, and is responsible for fragmented habitats. (Newmark 2008, p. 327). Boundary conducting and coordinating wildlife Two conservation tools used by fences along national borders that research activities in Tanzania. In this African range countries for lions include separate many reserves are being role, TAWIRI has been actively involved the establishment of protected areas and removed to form a 35,000-km2 park. in promoting the development of and the enforcement of protections in these Limpopo National Park (formerly monitoring the use of wildlife corridors areas (Mesochina et al. 2010a and b; known as Coutada 16) in Mozambique, in Tanzania. Surveys conducted in 2009 Treves et al. 2009, pp. 60, 64). However, Kruger National Park in South Africa, and 2010 suggest that the Nyanganje several problems have emerged. For and Gonarezhou National Park, Manjinji Corridor in Tanzania is no longer being example, certain land-tenure systems do Pan Sanctuary, and Malipati Safari Area used by elephants and other wildlife. not recognize community ownership of in Zimbabwe will all be connected, as This corridor is at a narrow passage in land and wildlife and undermine the will be the area between Kruger and the Kilombero Valley and is the shortest extent to which benefits are converted Gonarezhou, and the Sengwe communal distance for animals to cross between into incentives for conservation. land in Zimbabwe and the Makuleke the Udzungwa and Selous ecosystems. Protected-area ‘‘boundaries’’ are not region in South Africa (Newmark 2008, Despite efforts in place, much of the always visible. Additionally, law p. 327). However, in some locations, corridor is being encroached upon by enforcement in protected areas can be areas that have previously been conversion of land to rice farming and sporadic, and parks are often designated as corridors have been cattle grazing (Jones et al. 2012, p. 469). understaffed (Pfeifer et al. 2012, pp. 1, encroached upon by human settlements Because these activities often deter 7). More recent evidence suggests that and agriculture (Estes et al. 2012, pp. wildlife from passing through, the some protected areas are being more 258–261; Jones et al. 2012, p. 469). corridor is ineffective (Jones et al. 2012, commonly encroached upon as human Tanzania is an example of a country p. 469). populations expand and search for attempting to reconnect habitat. As of In the latter half of the 20th century, resources. 2002, the Tanzanian Government, with lions in India were on the verge of Despite encroachment, protected donor and NGO support, was extinction. However, conservation areas are somewhat effective at reconnecting the nine largest blocks of measures were put in place to protect protecting wildlife and habitat as rates forest in the East Usambara Mountains lion habitat. In 1965, Gir Wildlife of habitat loss tend to be lower in using wildlife corridors (Newmark 2002, Sanctuary was created and became the protected areas than outside them various). Additionally, the 2009 first protected area in Gujarat. In 1972, (Green et al. 2013, p. 70; Pfeifer et al. Wildlife Act of Tanzania allows the the Gir Lion Sanctuary Project began. 2012, p. 2). African countries are Minister, in consultation with relevant Two-thirds of the pastoral families realizing the benefits of managing their local authorities, to designate wildlife living in the Sanctuary, and their wildlife populations and parks for corridors, dispersal areas, buffer zones, livestock, were relocated outside Gir tourism; however, conservation of vast and migratory routes. The 2010–2015 forests (Singh and Gibson 2011, p. areas of land for megafauna such as the National Elephant Management Plan of 1754). The area of Gir Wildlife lion is not only complex, but also Tanzania indicates that corridors are the Sanctuary was expanded and the core expensive. As an example, the 28-km primary objective of the plan, and area designated as Gir National Park in (17-mi) elephant corridor, completed in although primarily designed for 1975. 2011 in Kenya, cost $1 million USD elephants, these corridors allow for Following these actions, habitat began (The Nature Conservancy 2013, continuity of populations of other large to recover, the wild ungulate population unpaginated). Additionally, the overall mammal species such as lions (Jones et increased, and, subsequently, lion costs of anti-poaching and al. 2012, p. 470). numbers increased (Singh and Gibson compensation is expected to increase in In 2011, Kenya (which neighbors 2011, pp. 1754, 1755). Habitat adjacent range states concurrently with growing Tanzania to the North), completed a 28- to Gir was also declared a Sanctuary human populations, declining km corridor through an area that had (Pania Sanctuary) in 1989. This area and purchasing power of external funds, and been heavily impacted by human– surrounding community lands were corruption (Garnett et al. 2011, pp. 1–2; wildlife conflict. The purpose of the declared protected forests to serve as a Wittemyer et al. 2008, pp. 123, 125). corridor was primarily to reduce buffer area to the Gir Forests (Singh and Another mechanism for protecting human–elephant conflict and appears to Gibson 2011, p. 1754). As the lion habitat is to reconnect fragmented have been successful (Mount Kenya population began to increase, lion

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dispersed into satellite forest patches. along national borders that separate independence in 1990, Namibia began These reclaimed patches of habitat were many reserves in addition to creating or to turn over ownership of wildlife areas protected and the Mitiyala Sanctuary improving corridors to link good-quality to local communities (van Vliet 2011, p. was created in 2002, and the Girnar habitat for wildlife (Gadd 2012, p. 179; 29; Bandyopadhyay et al. 2010, p. 6). By Sanctuary, in 2007 (Singh and Gibson Newmark 2008, pp. 323–324). 2011, Namibia had 64 communities that 2011, p. 1754). To address the increasing covered 17 percent of the country total After 40 years, the protected areas of consumption of bushmeat, host area (van Vliet 2011, p. 29; Connif 2011, India have experienced habitat recovery, countries have employed a variety of unpaginated; NASCO 2011, p. 4). The a 10-fold increase in ungulates, and an different strategies, including the majority of the incomes from these increase in lion numbers (Singh and development of alternative industries conservancies come from ecotourism, Gibson 2011, pp. 1754, 1756). Since for communities. Helping local followed by trophy hunting (NASCO 1968, India’s Forest Department has communities develop alternate 2011, p. 22). These incomes are then conducted wildlife censuses every 5 industries represents one of the ways used to support infrastructure years (Singh and Gibson 2011, p. 1754), range countries can reduce their improvement in the community. In documenting a steady increase in the dependence on bushmeat. Throughout addition, legal bushmeat acquired lion population. Community pride and Africa, several ideas have been within conservancy lands is distributed love of lions, the media, and political attempted with varying levels of to local families (NASCO 2011, p. 25). pressure has ensured efforts are made to success. For example, the Anne Kent The success of the program in Namibia protect these lions. When problems Taylor Fund (AKTF) helps local Maasai has been attributed to Namibia’s unique arise, they are quickly assessed and a women to buy beads and other supplies characteristics, including low solution found. For example, when 6 to produce traditional items for the local population density and favorable lions were hit and killed by trains, tourist industry (AKTF 2012, p. 7; seasonal rain, which helps prey species immediate action was taken to rectify Lindsey et al. 2012b, p. 45; van Vliet recover (van Vliet 2011, p. 30). Despite the problem (Meena 2014, p. 26). 2011, p. 17). In addition, AKTF helps the successes in Namibia, the country’s Because of these actions, lions in India organize local men into anti-poaching unique characteristics mean that now number 523 (BBC 2015, and de-snaring teams (AKTF 2012, p. 5; adapting Namibia’s success to other, unpaginated). van Vliet 2011, p. 17). By creating more densely populated countries will programs targeting both men and Conservation Measures in Place To be difficult. women, AKTF creates an environment Stem the Loss of Prey Base that provides communities with Conservation Measures to Stem Human- Lions, like most large carnivores, prey financial stability as well as direct Lion Conflict upon a variety of species including community interest in protecting local As the human population expands, buffalo, plains zebra, wildebeest, , wildlife. With 13 years assisting local the potential for conflict with wildlife gemsbok, kob, and warthog (Kenya communities, the AKTF represents one increases. In Africa, conflict between Wildlife Service 2013, p. 13; Beg and of the more successful attempts to villagers and lions, who prey upon Beg 2011, p. 4; Nowell and Jackson encourage locals to shift away from livestock, represent a threat to the 1996, p. 18). Depletion of these prey relying on bushmeat. species (Chardonnet et al. 2010, p. 12; species due to competition with humans Studies compiled by Hazzah (2013 Moghari 2009, p. 14; IUCN 2006a, p. represents a threat to the lion pp. 1, 8) have shown that local 23). In addition, habitat loss due to (Chardonnet et al. 2005, pp. 8–9). As communities who live near protected conversion of land increases the chance noted, the increase in the human areas with more lenient policies have a of villagers coming into direct contact population in Africa is a major more positive attitude and relationship with lions (Chardonnet et al. 2010, p. contributor to the increase in demand with both the manager and the protected 24). In an attempt to address these for bushmeat, which in turn increases area as a whole. This open approach to problems, range countries have human encroachment into wildlife protected area management reflects a employed a variety of different territory (Lindsey et al. 2012b, p. 36). In trend in recent years to bring in local strategies to help the lion. Such addition to the increase in the human communities to assist in the strategies involve education, an effective population, lack of an alternative management of protected areas (Lindsey conservation plan, and interacting with livelihood, lack of alternate food et al. 2012b, p. 53). Wildlife the local community. sources, and lack of clear rights over management programs run by local Historically, range countries seek to land or wildlife are contributing factors communities are defined by two goals: mitigate human-lion conflict through toward the increase in demand for conserving wildlife and providing controlling rather than conserving the bushmeat (Lindsey et al. 2012b, pp. 36– economic aids to the community predator population. In countries such 41). The advent of automatic weapons (Bandyopadhyay et al. 2010, p. 5). With as Malawi, for example, the Department in the bushmeat trade impacts the lion’s regard to discouraging the consumption of Game, Fish and Tsetse Control would prey base, which is being hunted at of bushmeat, this new approach is seen shoot large carnivores that preyed upon unsustainable levels. in the creation of community-based livestock. Because of this policy, more Reconnecting fragmented habitat has wildlife management programs (van than 560 predators (which include the additive effects of not only Vliet 2011, p. 26). The purpose of these lions) were killed in the country conserving the biodiversity of the lion’s programs is to give the local community between 1948 and 1961, (Mesochina et habitat, but also that of its prey base a direct stake in the management of al. 2010b, p. 35). While this department (Lindsey et al. 2012b, p. 43). These wildlife areas. One use for these areas is was disbanded in 1963 and jurisdiction types of restoration practices enhance to turn them into game ranches. These shifted to the new Department of the health of species by allowing genetic areas are used both for legal bushmeat Forestry, crop and livestock protection interchange to occur and, thus, conserve production as well as trophy hunting still remains an important part of its the genetic diversity of all wildlife. and ecotourism. function. Despite the department Wildlife management entities are Namibia has had great success in focusing on protecting crops and linking many of the major protected setting up community-run livestock, the number of lions killed in areas by removing boundary fences conservancies. After gaining the country has declined. Between 1977

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and 1982, eight lions were killed, lion hunts were stopped or prevented countries continue to recognize and whereas six lions were killed between between 2010 and 2014 (Lion Guardians support the role that local communities 1998 and 2007 (Mesochina et al. 2010b, 2014, p. 6; Lion Guardians 2013, p. 5). play in lion conservation. Greater p. 35). While fewer lions are being Lastly, in the years of Lion Guardians support by countries to address the killed than in the previous decades, operations, lion kills have decreased by needs of local communities, and thereby problems remain, including lack of 95 percent and the number of lions has address the needs of lions, may be the resources, lack of manpower, and steadily increased; a total of 286 lions single-most important role these corruption within the range countries. have been documented in the Amboseli- countries can play in changing the Current governmental management of Tsavo ecosystem (Lion Guardians 2014, trajectory of lion declines. lions in countries such as Malawi, p. 6; Lion Guardians 2013, p. 5). Tanzania, and Zambia are managed by In addition, Lion Guardians work Finding the Problem Animal Control units with tribal elders to dissuade young Section 4 of the Act (16 U.S.C. 1533) (Mesochina et al. 2010a, p. 41; men from killing lions for ceremonial and implementing regulations (50 CFR Mesochina et al. 2010b, p. 36). When purposes. Historically, the killing of part 424) set forth procedures for adding lion attack incidents occur, Problem lions through ritualized lion hunts species to, removing species from, or Animal Control dispatches officials to called ilmurran is rewarded with gifting reclassifying species on the Federal investigate the problems. If the problem of cows and other rewards (Lion Lists of Endangered and Threatened lion is located, it is either removed or Guardians 2012, p. 5; Goldman et al. Wildlife and Plants. Under section eliminated. When properly funded, this 2010, p. 334). After introducing village 4(a)(1) of the Act, a species may be program has helped in reducing not elders to the Lion Guardians program determined to be an endangered species only conflicts between lions and first hand, many return home to their or a threatened species based on any of humans but also has driven down the village and give their blessings to the the following five factors: numbers of lions killed. Between 2005 project. This education led to significant (A) The present or threatened destruction, and 2009, there were 116 reported cases results; on August 11, 2013, two Lion modification, or curtailment of its habitat or of lions killed, with the number of lions Guardians stopped a group of hunters range; killed being less than 50 per year in who were planning to hunt a lion in (B) Overutilization for commercial, Tanzania (Mesochina et al. 2010a, p. retaliation for the lion preying on their recreational, scientific, or educational 41). However, limitations of resources livestock. The local village elders fined purposes; (including both manpower and funds) the potential hunters two cattle each for (C) Disease or predation; have hampered the effectiveness of going on a lion hunt, marking a gradual (D) The inadequacy of existing regulatory these officials in responding to these but significant shift in the cultural mechanisms; or (E) Other natural or manmade factors incidents. In addition, many Problem attitudes regarding the lion (Hazzah et affecting its continued existence. Animal Control interventions resulted al. 2014, p. 858; Lion Guardians 2013, in the death of the lion (Mesochina et p. 20). Between 2007 and 2014, only A species is ‘‘endangered’’ for al. 2010a, p. 41; Chardonnet et al. 2009, five lions had been killed in territories purposes of the Act if it is in danger of p. 36). Even in cases of translocation, where Lion Guardians operates, in extinction throughout all or a significant the lions that were being transported contrast to more than 100 lions killed in portion of its range and is ‘‘threatened’’ often end up injured or continue to pose adjacent areas (Lion Guardians 2013, p. if it is likely to become endangered problems to the community (Bauer et al. 5). Furthermore, reduced lion mortality within the foreseeable future throughout 2007, p. 91). was sustained across multiple years, all or a significant portion of its range. NGOs are also assisting in protecting resulting in the reserve having one of The ‘‘foreseeable future’’ is the period of lions. Intervention by NGOs often takes the highest lion densities in Africa time over which events or effects the form of interacting with the local (Hazzah et al. 2014, p. 857; Schuette et reasonably can or should be anticipated, community (Winterbach et al. 2010, p. al. 2013, p. 149). Despite the success of or trends extrapolated. 98). Lion Guardians, which operates in this program, retaliatory as well as As required by the Act, we conducted Kenya and Tanzania, recruits and ceremonial killings of lions outside the a review of the status of the species and educates local young men to monitor program areas remain a threat to the considered the five factors in assessing and track lion movement and warn species. whether the lion is in danger of herders of lion presence in the area, We found that many of the lion range extinction throughout all or a significant recover lost livestock, reinforce states are trying to address lion portion of its range or likely to become protective fencing, and intervene to stop conservation through the establishment endangered within the foreseeable lion hunting parties, thereby mitigating of protected areas, wildlife management future throughout all or a significant or preventing possible human-lion areas, wildlife corridors, and portion of its range. We examined the conflict (Hazzah et al. 2014, p. 853; Lion reconnecting habitat. In some areas, best scientific and commercial Guardians 2013, p. 7; Lion Guardians creating incentives for lion conservation information available regarding the past, 2012, p. 3). From 2010 to 2013, Lion is occurring through community present, and future threats faced by the Guardians maintained a recovery rate of conservation programs in range lion. We reviewed the petition, lost livestock of more than 85, totaling countries. In other cases, participatory information available in our files, other over $1.5 million USD; in 2014 alone, strategies have been implemented to available published and unpublished more than 20,000 livestock (93 percent) enhance local tolerance for large information, and comments received were recovered (Lion Guardians 2014, p. carnivores in Africa. An increasing from peer reviewers and the general 7; Lion Guardians 2013, p. 6). Since number of programs encourage local public. 2010, 1,700 bomas have been reinforced communities to solve problems that When considering what factors might to reduce depredation of livestock. End- arise from human–lion conflict without constitute threats to a species, we must of-year sampling shows that more than killing lions. However, the effectiveness look beyond the mere exposure of the 90 percent of reinforced bomas sampled of these measures still ranges from species to a factor to evaluate whether did not experience further depredation successful to unsuccessful, due in part the species may respond to the factor in (Lion Guardians 2014, p. 7; Lion to lack of resources, political will, and a way that causes actual impacts to the Guardians 2013, p. 6). Additionally, 103 infighting. It is imperative that range species. If there is exposure to a factor

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and the species responds negatively, the potential strongholds, and are sustenance, leading to increased factor may be a threat and we attempt particularly a threat in western, central, presence of hunters in protected areas to determine how significant a threat it and eastern Africa and some parts of and higher than average declines in is. The threat is significant if it drives, southern Africa. Lions are generally wildlife. or contributes to, the risk of extinction incompatible with humans and human- Due to growing demand and of the species such that the species may caused habitat alteration and loss; they availability of modern weapons, many warrant listing as endangered or are the least successful large African wildlife species, including the lion’s threatened as those terms are defined in carnivore outside conservation areas. In prey base, have become depleted in the Act. order to survive, they require larger many areas. Hunters are increasingly Overall, the lion population has contiguous habitats with fewer negative focusing on protected areas since declined and is expected to continue to human impacts than other more wildlife has been depleted in non- decline. Across its range, the lion is resilient species. Expansion of human protected areas. Bushmeat hunting is facing threats stemming from human settlements and activities into lion illegal, yet weak management and population growth. We find a number of habitat renders it unsuitable for lions, inadequate law enforcement have factors are currently impacting the primarily because human expansion facilitated poaching of bushmeat in species and will impact the species in results in reduced availability of wild protected areas. Significant decreases in the future. In general, these factors prey and lion mortality due to increases large mammal populations, including include: Habitat fragmentation, in human–lion conflict. Both of these lion prey species, have occurred in degradation, and loss (Factor A); factors influence the distribution and protected areas throughout Africa. excessive mortality due to trophy population viability of lions. Overall, the large mammal population hunting and trade in lion bone (Factor Furthermore, fragmentation and has declined 59 percent. Regional B); disease (Factor C); loss of prey base, isolation of lion habitat and populations differences in herbivore population retaliatory killing due to human–lion can also impact dispersal and genetic abundance were also detected. Because conflict, deleterious effects due to small viability. prey availability is an important factor populations, and climate change (Factor Prey availability is essential to lion for lions, decreases in prey densities E); and inadequate regulatory survival as it affects reproduction, result in decreases in lion density. mechanisms and weak management of recruitment, and foraging behavior and, Expansion of human settlements and protected areas (Factor D). therefore, also impacts lion movement, agricultural and pastoral activities into Overall, the lion population has abundance, and population viability. protected areas not only decreases prey decreased by 43 percent over the last 21 Prey abundance does not appear to be availability, it increases exposure of years. Regional variations indicate an 8 a concern for lion populations in India. livestock and humans to lions, thus percent increase in southern Africa and Conservation initiatives have ensured resulting in human-lion conflict. Most a 55 percent increase in India; however, that ample prey is available, and the conflict occurs at protected area the eastern region and western and pastoral communities that cohabitate boundaries where villages are central region (combined) decreased by with lions are primarily vegetarian; established and human encroachment 59 and 66 percent, respectively, in the therefore, there is no competition for occurs, which increases the chance of past 21 years. Furthermore, almost all food and no demand for bushmeat. In human-lion encounters. Furthermore, lion populations in Africa that Africa, lions are under serious threat cattle herders enter protected areas, and historically exceeded 500 individuals, due to decreased prey abundance. lions move beyond the borders of the minimum number estimated to Widespread decreases in prey species protected areas in search of food, constitute a viable population, are have been driven by human population increasing interactions between humans declining. growth and unsustainable, increasingly and lions and the risk of human-lion Human population growth has led to commercialized bushmeat hunting in conflict. a substantial decrease in lion habitat and around protected areas. The most significant cause of human- over the past 50 years. Current savanna Bushmeat is an important source of lion conflict is livestock depredation habitat that is suitable for lions is protein and livelihood in Africa. The and, to a lesser extent, attacks on fragmented and totals only 25 percent of growing human population increases humans. As a result of prey species African savanna habitat. This loss of the demand for bushmeat, fueling trade, becoming depleted in many areas, lions habitat has resulted in local and urban markets, and international will seek out livestock. Additionally, regional lion population extirpations, markets. Bushmeat sold at elevated when pastoralists graze increasing reduced lion densities, and a prices increases commercialization and numbers of livestock in and adjacent to dramatically reduced range; this the number of hunters. These hunters, protected areas and cultivate land up to decrease in habitat also partially who are often poor, are enticed by the and within the boundaries of protected explains why lions are now largely quick income to find more efficient areas, humans and livestock are limited to protected areas. Due to good hunting methods, putting subjected to lions, and the risk of protection and management, lions in unprecedented pressure on wildlife. predation and the number of livestock India have dispersed to additional Bushmeat contributes significantly to lost to predation increases. Conversion forested habitat outside the protected food security, and is often the most of rangeland to agricultural land has area, extending their range. Lion habitat important source of protein in rural blocked migratory prey routes, forcing in Africa, however, continues to be areas. It comprises between 6 percent lions to rely more on livestock. threatened by expansion of human (southern Africa) and 55 percent (CAR) Additionally, because most protected settlements, despite occurring within of a human’s diet within the lion’s areas are too small to support a lion’s protected areas. African range. In western Africa, large home range, adjacent dispersal Expansion of human settlements, bushmeat is a secondary source of areas are often used by lions in search agriculture, and/or livestock grazing are protein, with fish being the primary of prey, putting them into greater reported as occurring in or on the source. However, when widespread loss contact with livestock and humans. periphery of several areas identified by of jobs and income occurs due to poor Conditions worsen as livestock numbers Riggio et al. (2013, suppl. 1) as lion fish harvests, bushmeat becomes an and areas under cultivation increase, strongholds (viable populations) and important source of income and leading to overgrazing, further habitat

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destruction, and greater depredation Six management weaknesses have of the surviving male coalition. rates. Attacks on humans appear to be been identified in the current Additionally, a study found a 100 more frequent in southern and eastern management of lion hunting. These percent fatality rate for males that are Africa and rare in western and central weaknesses include: (1) A lack of prematurely forced to disperse due to a Africa. scientifically based quotas, which new male takeover. A lack of mature Livestock provide an economic value results in excessive harvests; (2) a lack males dispersing, whether it’s due to to humans, particularly those in extreme of enforcement in age restrictions, trophy hunting or retaliatory killing, poverty. When lions have no economic which leads to unsustainable harvests, reduces the genetic viability of value to local communities and they kill increased rates of infanticide, and populations and may contribute to local or are perceived to kill livestock, the population declines; (3) hunting of population extinctions. economic impact to local communities female lion in Namibia, which decreases Lion experts recommend age-based can be significant. Impacts on victims of reproduction success, thereby strategies be incorporated into lion lion attacks create resentment towards decreasing males available for trophy management action plans. Although the lions and lion conservation, and a hunting; (4) the use of fixed quotas that, 6-year method has the potential to greater likelihood of retaliation. The which encourages hunters to be reduce the rate of infanticide in lion most common solution to lion attacks is unselective in their take of a trophy (i.e., populations subject to trophy hunting, retaliatory killing. Spearing, shooting, they will kill younger, less desirable the issue of incorporating this strategy trapping, and poisoning of lions occur males); (5) a lack of minimum hunt into each country’s conservation regularly. Retaliatory killings have been lengths or minimum lengths that are too strategy and/or action plan, and reported as a significant threat to lion short to allow hunter the time needed to following up with implementation, populations in protected areas of be more selective in their take of enforcement, and transparency, has yet western and central Africa, Botswana, trophies; and (6) general problems to be observed in many of the lion’s South Africa, Cameroon, Kenya, associated with management of trophy range countries. Lack of implementation Tanzania, and Zimbabwe. Despite close hunting, including corruption, of age-based strategies may undermine occupation of India’s lion population allocation of concessions, and lack of the successful use of trophy hunting as with human settlements, increased benefits to communities and recognition a sustainable conservation strategy. predation on livestock, and some of the important role they play in Trade in lion parts and products are retaliatory killing of lions, human-lion conservation. common in western and central Africa. conflict and associated retaliatory Documented declines in lion Lion populations in these regions are killing is not a major source of lion populations of Africa are a result, in small and declining and, therefore, the mortality for that population. part, of mismanaged trophy hunting. common use of lions in these regions for Every year, human-lion conflicts Multiple researchers have documented their parts and products is likely intensify due to habitat loss, poor declines in lion populations across the unsustainable. Further, there seems to livestock management, and decreased range of the species as a result of be a burgeoning trade in lion bone to availability of wild prey. Because most mismanaged trophy hunting. supplement or replace tiger bone. There human-lion conflict occurs at the Specifically, negative impacts to lions is potential that the current legal trade borders of protected areas, only those from excessive offtakes have been in lion bone will eventually not be prides that occur near the borders are documented in Benin, Cameroon, enough to supply demand, resulting in subjected to human-lion conflict. Tanzania, Zambia, and Zimbabwe. poaching of lions in the future for the However, when these lions are removed Additionally, the effects of over- Asian medicinal trade. via retaliatory killing, territorial gaps are harvesting can extend into adjacent As a result of human population then filled with lions that may have national parks where hunting is expansion into lion habitat, lions are occurred closer to the core of protected prohibited. increasingly exposed to diseases from areas, causing these border areas to Except in Mozambique, trophy domestic animals. Because lions are a serve as population sinks and exposing hunting quotas are higher than the top predator, they are at a particularly more lions to human-lion conflict and recommended maximum harvest of 1 high risk of exposure to pathogens. retaliation. Retaliatory killing of lions lion per 2,000 km2. Additionally, the Available studies do not indicate that continues in many areas, and this mean actual harvests in Burkina Faso, infection with a single disease is practice impacts the viability of lion Zambia, Namibia, and Zimbabwe are causing detrimental impacts to lions at populations across their range. The higher than the recommended 1 lion per the species level, although general body killing of lions due to human-lion 2,000 km2 offtake. condition, health, and lifespan may be conflict is enough to result in the local In the absence of reliable population compromised and result in negative extirpation of lion populations. estimates, age restriction on trophy impacts at the individual or population Lions are a key species in sport harvests can ensure sustainability. If level. Co-infections, however, could hunting, or trophy hunting, which is offtake is restricted to males older than have synergistic effects that lead to carried out in a number of range 6 years of age, trophy hunting will likely greater impacts on lions than a single countries. If managed correctly, trophy have minimal impact on the pride’s infection. hunting can be an important social structure and young. By removing Disease appears to be a secondary management tool for conserving land only males 6 years of age or older, factor influencing the decline of lions and providing financial resources for younger males remain in residence long when co-infections occur or when lion conservation. However, enough to rear a cohort of cubs disease is combined with other factors, management programs are not always (allowing their genes to enter the gene including environmental changes, sufficient to deter unsustainable pool; increasing the overall genetic reduced prey density, and inbreeding offtakes, which has resulted in declines diversity); recruitment of these cubs depression. Diseases weaken in lion populations in many areas. The ensures lion population growth and, individuals and allow them to succumb main problem with mismanaged trophy therefore, sustainability. However, to other diseases or factors. Although hunting stems from excessive harvests harvesting males that are too young disease does not appear to be a major because of impacts associated with causes male replacements, which results driver in the status of the lion, removal of males. in increased infanticide rates and death populations can suffer significant losses;

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some may recover to pre-outbreak events. The establishment of another Increases in average rainfall in the levels, others may not. Given the small geographically separated, free-ranging past 20 years have resulted in the and declining lion populations that population would reduce the risk of conversion of dry savanna to forestland remain, any loss of individuals from the extinction. Establishment of a new in India; however, these lions have used populations could be highly population at Kuno Wildlife Sanctuary both habitats. Therefore, habitat detrimental. in Madhya Pradesh State has been conversion due to climate change may The viability of a lion population proposed. However, the Government of not be as detrimental to lions in India. partly depends on the number of prides Gujarat has refused to allow any lions However, increased risks of flooding and ability of males to disperse and from Gir to be transferred. could pose a problem for lions. interact with other prides, which affects As human populations continue to Additionally, lions could face threats exchange of genetic material. Without rise in sub-Saharan Africa, the amount following flood events, such as an genetic exchange, or variation, of land required to meet the expanding outbreak of disease. Because this individual fitness is reduced and human population’s needs is constantly population is small, isolated, and less species are less able to adapt to increasing. Lions are increasingly genetically diverse, it is more vulnerable environmental changes and stress, limited to protected areas, and human to stochastic events and more prone to increasing the risk of extinction. population growth rates around local extinction events. Male dispersal plays an important protected areas in Africa tend to be Current lion habitat and suitable role in determining the level of higher than the average rural growth habitat predicted to remain under inbreeding in lion populations. The rate. Considering the majority of the climate change scenarios will be under fewer number of males present to human population in sub-Saharan increasing pressure due to land contribute genes to the next generation, Africa is rural, and land supports the conversions to meet the needs of the the more inbred the population will be. livelihood of most of the population, growing human population. Projected Therefore, not only does dispersal loss and degradation of lion habitat, loss changes in Africa’s climate will increase impact inbreeding, so does the loss of of prey base, and increased human-lion this pressure as land becomes more arid male lions due to excessive trophy conflict can reasonably be expected to and food security concerns are hunting and infanticide. Because the accompany the rapid growth in sub- exacerbated. Adaptive responses may number of prides and male dispersal are Saharan Africa’s human population into result in further encroachment into the most important factors for the foreseeable future. natural habitats. Land conversion will maintaining viability, sufficient areas Impacts described above from existing restructure the landscape, disrupt prey are needed to support at least 50 prides, and predicted anthropogenic pressures migration, and decrease prey available but preferably 100 prides, and allow on the species and its habitat are likely to lion. Lion densities decrease with unrestricted male dispersal. to be exacerbated by climate change. increasing mean temperature and Unfortunately, few lion populations The general warming and drying trend decreasing rainfall. Therefore, lion meet these criteria as almost all lion projected for Africa could further reduce density, or carrying capacity of populations in Africa that historically lion range, numbers, and prey base. protected areas, in sub-Saharan Africa is exceeded 500 individuals are declining, Lions may also have to travel greater likely to decline with climate warming and few protected areas are large distances to find food or shift their diet and drying. enough to support viable populations. to livestock, increasing conflict with The loss of lions could also mean the Furthermore, research indicates that humans and the risk of retaliatory loss of genetic variation. Combined with there is a general lack of gene flow in killings. Additionally, changes in declining populations, the risk of most lion conservation units. climate may increase the number and inbreeding and associated Lack of dispersal and genetic intensity of disease outbreaks in lions complications could increase. Drought variation can negatively impact the and their prey. conditions can also contribute to reproductive fitness of lions in these Under different climate change reduced prey availability by altering the populations and local extirpation is scenarios between the years 2040 and timing of migration. Climate conditions likely. Loss of fecundity leads to a 2070, no broad new areas will become also influence prey abundance, and the decrease in population size, fewer suitable for lion. Southern Africa, where loss of prey species can result in lions prides in a population, and increased the broadest areas of suitable conditions shifting their diet towards livestock, inbreeding which contributes to a occur, is projected to become less which may increase retaliatory killings decline in the population and increases suitable because of climate change. A by humans. the risk of extinction. Additionally, lack broad swath of potential distributional Diseases can be directly and of genetic variation can impact the area in western Africa is projected to indirectly affected by climate change by ability of lions to withstand stochastic become ‘‘distinctly less suitable or even impacting distribution, the timing of events or limit the lion’s ability to uninhabitable.’’ A decrease in the lion’s outbreaks, and the intensity of evolve responses to climate change. range could mean that stochastic events outbreaks. Severe climate change could India’s lion population is isolated and impact a larger portion of the whole synchronize temporal and spatial genetically less diverse. Currently, there species, especially if it occurs where the convergence of multiple infectious is no evidence of depressed species and its habitat occur. agents, triggering epidemics with greater demographic parameters. However, Additionally, reductions in populations mortality than infections from a single intense management may interfere with and geographic range may limit the pathogen. natural selection by ensuring survival of lion’s ability to respond to climate National and international unfit lions, which facilitates the change. Conversely, climate change conservation strategies rely on protected propagation of deleterious genes in the effects on potential lion distribution are areas to protect natural resources from population. Being a small, isolated projected to be more neutral in eastern negative impacts of human populations. population and less genetically diverse, Africa than across the entire range. The lion is largely limited to protected therefore, it is more vulnerable to the Reserves in this region are more likely areas; therefore, effective management is loss of any individuals due to to sustain lion populations under crucial to the survival of the species. environmental and stochastic events, climate change scenarios in the However, weak management of and more prone to local extinction medium-term. protected areas has been documented

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across its range, especially in western availability of wild prey and lion including Nigeria, Burkina Faso, and Africa where most protected areas are mortality due to increases in human– Cameroon, maintain local markets in experiencing severe management lion conflict. Both of these factors lion products. Trade in lion skins and deficiencies. influence the distribution and partial skins is described as ‘‘frequent’’ Based on the best scientific and population viability of lions. in street markets in Abidjan, Coˆte commercial information, we find that Significant decreases in prey d’Ivoire, and the scale of domestic trade several factors are negatively impacting abundance have occurred in protected in illegal lion products is described as the lion and contributing to the risk of areas throughout Africa. In western ‘‘massive’’ in Nigeria. In the central extinction. However, we find there is a Africa, specifically, herbivore African country of Cameroon, the substantial difference in the magnitude populations have decreased by 85 estimated value of a single lion carcass of these threats to the risk of extinction percent. As a result of prey species exceeds the trophy fee, and at a lion between the subspecies P. l. leo and P. becoming depleted in many areas, lions conservation conference, the l. melanochaita. Based on current seek out livestock for food; attacks on Government of Cameroon identified population estimates, projected livestock occur at the highest frequency trade in lion skins as a major cause of population trends, and the threats in areas where natural prey abundance the decline in lion populations in described herein, we find that the is lowest. Traditional livestock western and central Africa. Trade in subspecies P. l. leo and P. l. husbandry practices can reduce lion skins is most likely one of the melanochaita qualify for different depredation rates, but these traditional biggest threats to lion survival in statuses under the Act. practices are being replaced with less western Africa due to the rarity of lions diligent practices. For example, in the Finding for Panthera leo leo in the region, the extent of the trade, Pendjari area of Benin, traditional and the high price of lion skins. Lion The range of P. l. leo includes the enclosures are low with few branches. populations in western and central western and central African regions and These structures and the lack of Africa are small and declining and, India. This subspecies has experienced enclosures encourage livestock therefore, the common use of lions in a reduction in range, a reduction in total predation. People do not invest much these regions for their parts and number of populations, and a reduction into improving enclosures even though products is likely unsustainable. in number of lions. There are they appear to be economically efficient, The viability of a lion population approximately 1,500 lions distributed ecologically effective, and culturally partly depends on the number of prides among 15 populations; 14 in Africa and acceptable. Even enclosures that were and the ability of males to disperse and 1 in India. The population in western built as part of a conservation project interact with other prides, which affects and central Africa has declined by 66 were not used full time due to lack of exchange of genetic material. Without percent since 1993. The current labor and, in some cases, the herd being genetic exchange, or variation, the more population estimate for this portion of too large for the enclosures. When lions inbred the population will be, its range is approximately 915 lions. in Africa cause or are perceived to cause individual fitness is reduced, None of the lion populations in these damage to livestock, property, or reproductive fitness is reduced, and regions meet the MVP, although we do people, the response is generally to kill species are less able to adapt to note that the WAP complex qualifies as them. Retaliatory killings are reported to environmental changes and stress or a potential stronghold where a viable be a significant threat to lion stochastic events. Loss of fecundity population could occur if immediate populations in western and central leads to a decrease in population size, interventions are implemented. Between Africa. fewer prides in a population, and 1993 and 2014, the Indian population Some countries in the African range increased inbreeding which contributes increased by 55 percent. A census of this subspecies allow hunting of P. l. to a decline in the population and may conducted in 2015 indicates the leo. Management programs do not result in local extirpation. The entire P. population has increased by 27 percent appear to be sufficient to deter l. leo subspecies comprises small, since 2010, with lions now numbering unsustainable offtakes, which has isolated populations. Research indicates 523. Although this population is found resulted in declines in lion populations that there is a general lack of gene flow within a protected area, its single, small in many areas. Specifically, negative in most lion conservation units. population of 523 animals continues to impacts to lions from excessive offtakes Furthermore, the suggested minimum be highly vulnerable to disease and have been documented in Benin and number of lions estimated to constitute other stochastic events. Due to weak Cameroon. Additionally, hunting quotas a viable population is at least 250 lions, management in Africa and small in Benin and Burkina Faso are too high but preferably 500 lions, or 50–100 populations throughout its range, this for sustainability, although Burkina prides. This threshold may be smaller subspecies continues to face threats. Faso has proposed to reduce their quota for P. l. leo as pride sizes are generally Remaining African populations are in the 2015–2016 season. Actual smaller than those for P. l. particularly threatened by expansion of harvests in Burkina Faso were also melanochaita. However, given the size human settlements, agriculture, and/or found to be higher than recommended of the remaining populations, few could livestock grazing. Expansion of levels. Although experts recommend be considered potentially viable. agriculture and livestock grazing are age-based strategies be incorporated into Additionally, few protected areas are reported in or around two of the larger lion management plans to reduce large enough to support viable African populations of P. l. leo, WAP excessive harvests and reduce the rate of populations. Complex and a Chad–CAR population; infanticide, Benin and Burkina Faso Although there are laws meant to management in portions of both have yet to implement an age-based protect wildlife, including lions and protected areas is reported as weak, strategy. As a result, species experts their prey species, the drastic and raising concern for the persistence of agree that there is no level of offtake that continuing decline of the species and its lions and their habitat. Expansion of would be sustainable for P. l. leo prey indicate these regulatory human settlements and activities into populations in their current condition. mechanisms are not adequate to lion habitat renders it unsuitable for Trade in lion parts and products is ameliorate threats to P. l. leo. lions, primarily because human very common in western and central Furthermore, national and international expansion results in reduced Africa. Many African countries, conservation strategies rely on protected

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areas to protect natural resources from conducive to disease outbreaks and are Africa, Tanzania, Zambia, and negative impacts of human populations. a serious concern to the persistence of Zimbabwe. Between 1993 and 2014, the However, weak management of the lion population as this population is lion population in eastern Africa protected areas has been documented more vulnerable to stochastic events declined by 59 percent. In southern across the lion’s range, especially in and local extinction. Africa the lion population increased by western Africa where most protected Threats acting on P. l. leo have 8 percent during the same time period. areas are experiencing severe contributed to large reductions in the Most of the increasing populations management deficiencies, including the subspecies’ range and suitable habitat, contributing to this trend are small, lack of a budget or a budget insufficient abundance, and number and fenced reserves. However, one of the to carry out management activities. connectivity of populations. The largest populations in southern Africa, The lion population in India is one subspecies has reached critically low Okavango, and populations in 6 population of P. l. leo that is increasing numbers of individuals and potentially unfenced reserves in Botswana, and could potentially be considered a viable populations. Furthermore, while Namibia, and Zimbabwe declined. viable population based on the number one small population may be increasing, Although there are larger populations of of lions. However, intense management, we are not aware of any information P. l. melanochaita that may meet the including healthcare interventions, may indicating that the overall trend of large suggested MVP, almost all lion interfere with natural selection declines in the subspecies range, populations in Africa that historically processes by ensuring the survival of abundance, and connectivity, will exceeded 500 individuals, are declining. unfit lions, which facilitates the reverse course. Expansion of human settlements, propagation of deleterious genes in the Threats continue to act on this agriculture, and/or livestock grazing is population. This population is also subspecies. Due to small population size occurring in or on the major populations running out of area to expand. Being a and lack of connectivity between and is particularly a threat in eastern small, isolated population and less populations, most populations are not Africa and some parts of southern genetically diverse, it is more vulnerable able to recover from the loss of suitable Africa. In particular, expansion of to the loss of any individuals due to habitat or individuals. Furthermore, agriculture and livestock grazing is environmental and stochastic events, because all populations are small and occurring in or around major and more prone to local extinction isolated, the subspecies lacks resiliency populations in Kenya, Tanzania, and events. to recover from stochastic or Zambia and both are major threats to As previously stated, threats to the catastrophic events and is thus highly lion survival in these countries. lion are expected to continue or increase vulnerable to extirpation. Threats are Expansion of human settlements and in conjunction with predicted human currently affecting the subspecies and activities into lion habitat renders it population growth. The human the impacts on the subspecies are unsuitable for lions, primarily because population, and thus negative impacts expected to continue or even intensify human expansion results in reduced to lions, as well as decreases in lion over time as the human population availability of wild prey and lion populations, associated with human increases and as climate change mortality due to increases in human– population growth, is expected to progresses, negatively impacting lion conflict. Both of these factors increase substantially by 2050. If availability of suitable habitat, lion influence the distribution and regional trends continue at their current distribution, and lion numbers. Based population viability of lions. However, rate, western and central Africa will on the current distribution and size of in some parts of southern Africa, lions likely lose a third of its population in 5 P. l. leo populations, the current threats are repopulating areas where lions were years and half the population in 10 acting on this subspecies, the impacts of recently extirpated due to adequate years. Lion bone may be increasingly those threats, and the impacts of future protection of habitat and prey. used as a replacement for tiger bone in threats and climate change on lion Significant decreases in prey traditional Asian medicine and in Asian distribution, lion numbers, habitat, prey abundance have occurred in protected luxury products. Therefore, trade in lion availability, susceptibility to disease, areas throughout Africa, including bone could become lucrative, spur loss of lions via human–lion conflict Botswana, Kenya, Mozambique, Sudan, considerable demand from suppliers of and trophy hunting, and resiliency to Zambia, and Zimbabwe. Herbivore the black market, result in extensive stochastic and catastrophic events, we populations have decreased by 52 poaching of wild lions, and have find that the viability of this subspecies percent in eastern Africa, although they significant impacts to lion populations. is compromised and will not be have increased by 24 percent in Additionally, future development in resistant or resilient to ongoing and southern Africa. Protected areas in India could alter habitat vital for future threats. Therefore, we find that P. Ethiopia, Mozambique, Tanzania, and dispersal. Tolerance to loss of livestock l. leo is in danger of extinction Zambia are increasingly settled; may also wane as traditional beliefs and throughout its range and list the decreases in prey abundance in African traditional value systems are rapidly subspecies as endangered. protected areas are driven by human changing under the influence of population growth, especially along the globalization. Furthermore, effects of Finding for Panthera leo melanochaita boundaries of protected areas where climate change on lion habitat are The range of P. l. melanochaita human population growth rates are projected to manifest as early as 2040. includes the southern and eastern high, encroachment and habitat loss Under climate change scenarios, a broad African regions. Although this occurs, and people are dependent on swath of potential distributional area in subspecies has experienced range bushmeat. Additionally, many western Africa is projected to become reduction, a decline in the number of communities lack the rights over land distinctly less suitable or even populations, and a decline in the and in most cases in Botswana, uninhabitable. Increases in rainfall number of lions, it remains relatively Tanzania, Zambia, and Zimbabwe, the predicted for India may not have widespread. Currently, there are government retains a significant portion detrimental impacts on lion habitat; approximately 17,730 P. l. melanochaita of revenue from wildlife; therefore, however, increased risks of flooding lions distributed among 68 protected those that bear the costs of wildlife do could result in increased mortality, and areas, with larger populations in not receive benefits, and bushmeat post-flooding conditions could be Botswana, Kenya, Namibia, South hunting is the only way to benefit from

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wildlife. Furthermore, conversion of Lack of implementation of age-based for the bone trade may currently exist as rangeland to agricultural use has strategies may undermine the successful prices paid to South African game blocked several migratory routes for use of trophy hunting as a sustainable farmers and landowners for lion bones Tanzania’s wildebeest and zebra conservation strategy. exceeded the per capita GDP (gross populations, which likely forces lions to The captive-breeding industry has domestic product) in many lion range rely more on livestock. publicized captive breeding and states. Thus, the current price paid for As a result of prey species becoming reintroduction of captive-born species lion bone appears to provide incentive depleted in many areas, lions seek out into the wild as a potential solution to in some countries to poach wild lions. livestock for food; attacks on livestock the decrease in wild lion populations. The viability of a lion population occur at the highest frequency in areas However, lions raised in captivity often partly depends on the number of prides where natural prey abundance is lowest. develop a variety of issues that make and ability of males to disperse and Additionally, traditional livestock them unsuitable for reintroduction, and interact with other prides, which affects husbandry practices can reduce reintroduction efforts have not been the exchange of genetic material. depredation rates, but these traditional shown to address the underlying causes Without genetic exchange, or variation, practices are being replaced with less of population declines throughout the the more inbred the population will be, diligent practices. In Kenya and species’ range. Existing research has individual fitness is reduced, Tanzania, social changes are altering generally found that captive-raised lions reproductive fitness is reduced, and traditional Maasai pastoral livelihoods, are not as able to adapt successfully to species are less able to adapt to reducing dependency on livestock, and conditions out of captivity and, environmental changes and stress or reducing traditional livestock care and therefore, the success rate is much stochastic events. Loss of fecundity management, leaving livestock more reduced compared to the use of wild- leads to a decrease in population size, vulnerable to predation. Although lions caught lions. fewer prides in a population, and While it is argued that South Africa’s generally avoid people, they will increased inbreeding, which contributes captive-bred lion industry may reduce occasionally prey on humans, causing to a decline in the population and local pressures of trophy hunting on wild serious injury or death. Attacks on extirpation. Research indicates that South African populations, there is no humans appear to be more frequent in there is a general lack of gene flow in substantial or peer-reviewed science to the range of P. l. melanochaita than P. most lion conservation units. support such a claim. Likewise, there is l. leo. When lions cause or are perceived Furthermore, the suggested minimum no record or evidence to support claims to cause damage to livestock, property, number of lions estimated to constitute or people, the response is generally to that the captive-bred lion industry is a viable population is at least 250 lions, kill them. Retaliatory killings are supporting reintroduction into the wild but preferably 500 lions, or 50–100 reported to be a significant threat to lion in any significant way. However, future prides. Almost all lion populations in populations in Botswana, South Africa, efforts to control hunting of captive-bred Africa that historically exceeded 500 Kenya, Tanzania, and Zimbabwe. lions could potentially increase the Some P. l. melanochaita range demand for wild lion trophies and individuals are declining, and few countries allow hunting of lions. result in excessive harvests. protected areas are large enough to Although some management programs Additionally, trade in bones of captive support viable populations. appear to follow recommended lions could stimulate harvest of wild While the lion bone trade appears to practices for sustainability, most do not lions to supply a growing bone trade. currently be based primarily in South appear to be sufficient to deter Hunting of captive lions could also Africa’s captive-bred lion hunting unsustainable offtakes, which has potentially undermine the price of wild industry, the trade appears to be having resulted in declines in lion populations hunts and reduce incentives for little or no impact on wild lion in many areas. Specifically, negative conservation of wild lions in other populations in South Africa at this impacts to lions from excessive offtakes African countries. time—lion populations in South Africa have been documented in Tanzania, Lion parts and products are used in are stable or increasing and there is little Zambia, and Zimbabwe. Additionally, many African countries as medicine, poaching of wild lions in the country hunting quotas in most countries are nutrition, talismans, and decorations, (Funston and Levendal 2014, pp. 1, 26; higher than the recommended offtake and in traditional ceremonies and Williams et al. 2015, pp. 79–80). for sustainability. Actual harvests in rituals. Kenya and Somalia maintain However, the impact of the lion bone Namibia, Zambia, and Zimbabwe were local markets in lion products. Lion trade on lion populations outside South also found to be higher than skins and canines are also described as Africa is unknown and most wild lions recommended levels. Experts ‘‘easily found’’ in the markets of Dakar, occur outside South Africa (see recommend age-based strategies be Senegal. In southern and eastern Africa, Distribution and Abundance). While incorporated into lion management trade in lion parts, particularly lion wild tiger populations are declining, the plans to reduce excessive harvests and bone, to Asia is generally considered a demand for tiger parts in Asia is reduce the rate of infanticide and severe potential threat to the species. increasing. With tigers difficult to several countries, including According to CITES, there is ‘‘clear obtain, lion bone may be increasingly Mozambique (only Niassa National scope for the international trade in lion used as a replacement for tiger bone. Reserve), Tanzania, and Zimbabwe have body parts for [traditional Chinese Considering the sharp and continuing committed to implementing an age- medicine and traditional African increases in demand from Asia for lion based strategy. Of these, only Niassa medicine] to grow uncontrollably, as it bone and the effect of the tiger bone National Reserve and Zimbabwe have has done for other big cats.’’ According trade on tiger populations, there is fully implemented age restrictions and to Kenya, the declared exports of bones, potential for demand to surpass the shown reductions in offtake. Tanzania skulls, and skeletons derived from wild availability of legally obtained lion has implemented age restrictions and lions also show an increasing trend bone. Therefore, trade in lion bone shown reductions in offtake; however, through the period 2003–2012, with could become lucrative, spur transparency (in terms of trophy quality total declared specimens in 2012 more considerable demand from suppliers of data) and the scientific objectivity of the than ten times those in 2003. Evidence the black market, result in extensive evaluating body has been questioned. suggests incentive to poach wild lions poaching and unsustainable harvest of

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wild lions to meet demand, and have endangered species within the conventions set forth in the ESA. When significant impacts to lion populations. foreseeable future and we are listing P. a species is listed as endangered, certain Although there are laws in place in l. melanochaita as a threatened species. actions are prohibited under section 9 of lion range countries that are meant to the ESA and are implemented through Significant Portion of Its Range protect wildlife, including lions and our regulations in 50 CFR 17.21. These their prey species, the drastic and Under the Act and our implementing include, among others, prohibitions on continuing decline of the species and its regulations, a species may warrant take within the United States, within prey in some parts of its range indicate listing if it is endangered or threatened the territorial seas of the United States, these regulatory mechanisms are not throughout all or a significant portion of or upon the high seas; import; export; adequate to ameliorate threats to the P. its range. The term ‘‘species’’ includes and shipment in interstate or foreign l. melanochaita throughout its range. ‘‘any subspecies of fish or wildlife or commerce in the course of a commercial Furthermore, national and international plants, and any distinct population activity. Exceptions to the prohibitions conservation strategies rely on protected segment [DPS] of any species of for endangered species may be granted areas to protect natural resources from vertebrate fish or wildlife which in accordance with section 10 of the negative impacts of human populations. interbreeds when mature.’’ We ESA and our regulations at 50 CFR However, weak management of published a final policy interpreting the 17.22. protected areas has been documented phrase ‘‘Significant Portion of its The ESA does not specify particular across the lion’s range. Range’’ (SPR) (79 FR 37578, July 1, prohibitions and exceptions to those As indicated above, P. l. melanochaita 2014). The final policy states that (1) if prohibitions for threatened species. remains relatively widespread and some a species is found to be endangered or Instead, under section 4(d) of the ESA, remaining populations are large enough threatened throughout a significant the Secretary, as well as the Secretary of to be considered viable. Therefore, due portion of its range, the entire species is Commerce depending on the species, to the size of some populations, the listed as endangered or threatened, was given the discretion to issue such number of remaining populations, and respectively, and the Act’s protections regulations as deemed necessary and the stability or increasing status of some apply to all individuals of the species advisable to provide for the populations, we find that P. l. wherever found; (2) a portion of the conservation of such species. The melanochaita is not currently in danger range of a species is ‘‘significant’’ if the Secretary also has the discretion to of extinction. However, the overall species is not currently endangered or prohibit by regulation with respect to population of the subspecies continues threatened throughout all of its range, any threatened species any act to decline and threats to the lion are but the portion’s contribution to the prohibited under section 9(a)(1) of the expected to continue or increase in the viability of the species is so important ESA. Exercising this discretion, the future in conjunction with predicted that, without the members in that Service has developed general human population growth. If regional portion, the species would be in danger prohibitions in the ESA regulations (50 trends in lion populations continue at of extinction, or likely to become so in CFR 17.31) and exceptions to those the current rate, eastern Africa will lose the foreseeable future, throughout all of prohibitions (50 CFR 17.32) that apply a third of its lion population in 20 years its range; (3) the range of a species is to most threatened species. Under 50 and half the population in 30 years. considered to be the general CFR 17.32, permits may be issued to Effects of climate change on lion habitat geographical area within which that allow persons to engage in otherwise are projected to manifest as early as species can be found at the time the prohibited acts for certain purposes. 2040. Although climate change effects Service or the National Marine Fisheries Under section 4(d) of the ESA, the on potential lion distribution are Service makes any particular status Secretary, who has delegated this projected to be more neutral in eastern determination; and (4) if a vertebrate authority to the Service, may also Africa than across the entire range, species is endangered or threatened develop specific prohibitions and southern Africa, where the broadest throughout an SPR, and the population exceptions tailored to the particular areas of suitable conditions occur, is in that significant portion is a valid conservation needs of a threatened projected to become less suitable DPS, we will list the DPS rather than the species. In such cases, the Service issues because of climate change. Specifically, entire taxonomic species or subspecies. a 4(d) rule that may include some or all park areas, including the ‘‘Etosha Pan, We found the lion subspecies P. l. leo of the prohibitions and authorizations Lake Opnono, Cuvelai Drainage, to be in danger of extinction throughout set out in 50 CFR 17.31 and 17.32, but Kalahari Gemsbok, and Kgalagadi its range, and the subspecies P. l. which also may be more or less Transfrontier Park areas’’ are projected melanochaita likely to become restrictive than the general provisions at to decline substantially in suitability for endangered within the foreseeable 50 CFR 17.31 and 17.32. For P. l. lions. In addition, reforms to trophy future throughout its range. Therefore, melanochaita, the Service has hunting have been made to ensure no portions of the species’ range are determined that a 4(d) rule is necessary sustainability of trophy hunting, but ‘‘significant’’ as defined in our SPR and advisable. these reforms have been implemented in policy, and no additional SPR analysis We are adding a 4(d) (special) rule for only a few places. Furthermore, demand is required. P. l. melanochaita at 50 CFR 17.40(r). for lion bone is expected to increase in This 4(d) rule maintains all of the the future and high prices for lion bone 4(d) Rule for Panthera leo prohibitions and exceptions codified in provide incentive to poach wild lions. melanochaita 50 CFR 17.31 and 17.32 with regard to As a result of the likely impacts of these The purposes of the ESA are to this subspecies and supersedes the threats, it is reasonable to conclude that provide a means whereby the import exemption found in 50 CFR 17.8 the population of P. l. melanochaita is ecosystems upon which endangered for threatened wildlife listed in likely to be drastically reduced and species and threatened species depend Appendix II of CITES, such that a fragmented in the foreseeable future, may be conserved, to provide a program threatened species import permit under limiting the ability of the subspecies to for the conservation of such endangered 50 CFR 17.32 is now required for the recover from stochastic and catastrophic species and threatened species, and to importation of all P. l. melanochaita events. Therefore, we find that this take such steps as may be appropriate to specimens. Therefore, through the subspecies is likely to become an achieve the purposes of the treaties and promulgation of this 4(d) rule, the

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presumption of legality provided under species listed pursuant to section 4 of a threatened species import permit. section 9(c)(2) of the Act for the the Act, but that is listed in Appendix While the Service cannot control otherwise lawful importation of wildlife II of CITES, shall be presumed to be in hunting of foreign species such as P. l. listed in Appendix II of CITES that is compliance with provisions of the Act melanochaita, we can regulate their not an endangered species listed and implementing regulations if the importation and thereby require that pursuant to section 4 of the Act does not importation is not made in the course of U.S. imports of sport-hunted P. l. apply to this subspecies. Thus, under a commercial activity. While there has melanochaita trophy specimens are this 4(d) rule, all otherwise prohibited been question as to whether this obtained in a manner that is consistent activities, including all imports of P. l. provision of the Act might automatically with the purposes of the Act and the melanochaita specimens, require prior require allowing the importation of a conservation of the subspecies in the authorization or permits under the Act. species that is both listed as threatened wild, by allowing importation from Under our regulations at 50 CFR 17.32, and in Appendix II, and preclude the range countries that have scientifically permits or authorization to carry out an issuance of more restrictive 4(d) rules sound management programs that otherwise prohibited activity could be covering importation, the Service has address the threats that are facing lions issued for scientific purposes, the concluded that such 4(d) rules may be and are enhancing the survival of the enhancement of propagation or survival issued to provide for the conservation of species in the wild within that country of the species, economic hardship, the involved species. Section 9(c)(2) (see further discussion below on zoological exhibitions, educational does not expressly refer to threatened enhancement of propagation or survival purposes, or special purposes consistent species or prevent the issuance of with regard to authorizing the import of with the purposes of the Act. appropriate 4(d) rules and could not sport-hunted trophies of P. l. Applications for these activities are logically have been intended to allow melanochaita). Further, for the import available from either http://www.fws. the addition of a species to an appendix of parts or products, there is evidence gov/forms/3-200-20.pdf or http://www. of an international convention to that trade in lion parts, particularly fws.gov/forms/3-200-37.pdf. override the needs of U.S. law, where bones, is fast becoming a substitute for The intent of this 4(d) rule is to there is reliable evidence to affect the tiger bones in traditional Asian provide for the conservation of P. l. presumption of validity. Finally, the medicine and Asian luxury products melanochaita consistent with the term ‘‘presumed’’ implies that the (see Traditional Use of Lion Parts and purposes of the Act. Under this 4(d) established presumption is rebuttable Products section). While the primary rule, the prohibitions, in part, make it under certain circumstances, including source of the current bone trade appears illegal for any person subject to the through the promulgation of a protective to be from captive-bred lions from South jurisdiction of the United States to regulation pursuant to section 4(d) of Africa, considering the sharp and ‘‘take’’ (includes harass, harm, pursue, the Act. continuing increases in demand from hunt, shoot, wound, kill, trap, capture, In the case of the P. l. melanochaita, Asia for lion bone, there is potential for or to attempt any of these) within the demand to surpass the availability of United States or upon the high seas; there are substantive grounds on which to challenge the presumption. For the legally obtained lion bone and, import or export; deliver, receive, carry, consequently, result in poaching and transport, or ship in interstate or foreign import of sport-hunted trophies, while there is evidence that some range unsustainable harvest of wild lions to commerce, by any means whatsoever, in meet demand. Based on the effect of the the course of commercial activity; or sell countries are implementing lion management programs, the best tiger bone trade on tiger populations, if or offer for sale in interstate or foreign current conditions continue unchanged, commerce any P. l. melanochaita available information indicates that not there is considerable potential for specimens. It would also be illegal to all lion hunting programs are well extensive poaching of wild lions to possess, sell, deliver, carry, transport, or managed or provide enhancement to occur in order to meet demand. Given ship any such wildlife that has been survival of the subspecies (see Trophy the current threats to the subspecies, taken in violation of the Act. We find Hunting section), Namely, mismanaged unsustainable harvest to supply a trade that these protections, including the trophy hunting is reported to contribute in parts could contribute to the further requirement for a permit for the import, to documented declines in lion decline of the subspecies. export, interstate and foreign commerce populations of Africa (Rosenblatt et al. and take for all P. l. melanochaita 2014, entire; Sogbohossou et al. 2014, Finally, due to our concerns about the specimens, will support and encourage entire; Becker et al. 2013, entire; increasing trade in lion bones and conservation actions for P. l. Lindsey et al. 2013a, entire; Packer et al. evidence that live lions are being melanochaita and require that permitted 2013, p. 636; Croes et al. 2011, entire; exported to Asia, presumably for the activities involving this subspecies are Packer et al. 2011, entire; Loveridge et bone trade, we find that unregulated carried out in a manner that is al. 2007, entire). Depending on how trade and the taking of live lions could consistent with the purposes of the Act trophy hunting is regulated and further contribute to the lion bone trade. and our implementing regulations. managed, trophy hunting can be a tool Further, the noncommercial imports of In connection with this 4(d) rule, the for conservation, but may also have live lions could be a cover for the Service notes that P. l. melanochaita is negative impacts on lions (Bauer et al. establishment of lion bone trade within listed in Appendix II of CITES and, 2015a, unpaginated; Lindsey et al. the United States. As with captive tigers without this 4(d) rule, could be 2013a, p. 1; Whitman et al. 2004, pp. and the use of live animals for the bone imported into the United States 176–177; Loveridge et al. 2007, p. 548). trade, the Service finds that the pursuant to section 9(c)(2) of the Act We want to encourage and support unregulated movement of lions within upon the presentation of a proper CITES efforts by range countries to develop the United States, as well as the import export permit from the country of programs that are based on sound or export of these animals is reasonably export, if such importation is not made scientific information. As noted, the likely to be used as a loophole for the in the course of a commercial activity. 4(d) rule for P. l. melanochaita would bone trade and serve as cover for the Section 9(c)(2) of the Act provides that provide for the importation into the establishment of lion bone trade within the otherwise lawful importation of United States of trophies taken legally the United States. By requiring permits wildlife that is not an endangered in range countries upon the issuance of for all otherwise prohibited activities

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under the Act, such as import, export, with any known program intended to species. It is also imperative that the program interstate and foreign commerce and enhance the survival probabilities of the is part of a legally recognized governance take, including noncommercial imports population from which the wildlife sought to system that supports conservation. be covered by the permit was or would be (c) Socio-Economic-Cultural Benefit: A of live lions, we can ensure that live removed; well-managed hunting program can serve as lions are not used to supplement the (iv) Whether the purpose for which the a conservation tool when it respects the local trade in lion bones. permit is required would be likely to reduce cultural values and practices. It should be Therefore, we find that regulation of the threat of extinction facing the species of accepted by most members of the the importation of all P. l. melanochaita wildlife sought to be covered by the permit; community, involving and benefiting local parts and products, including live (v) The opinions or views of scientists or residents in an equitable manner. The animals and sport-hunted trophies, will other persons or organizations having program should also adopt business practices ensure that imported specimens are expertise concerning the wildlife or other that promote long-term economic sustainability. obtained in a manner that is consistent matters germane to the application; and (vi) Whether the expertise, facilities, or (d) Adaptive Management: Planning, with the purposes of the Act and the other resources available to the applicant Monitoring, and Reporting: Hunting can conservation of the subspecies in the appear adequate to successfully accomplish enhance the species when it is based on wild. the objectives stated in the application. appropriate resource assessments and Our threatened species permitting monitoring (e.g., population counts, trend In addition to these factors, regulations at 50 CFR 17.32 provide data), upon which specific science-based particularly in relation to sport hunting, issuance criteria for threatened species quotas and hunting programs can be we find the IUCN Species Survival permits (50 CFR 17.32(a)(2)), but do not established. Resource assessments should be Commission (SSC) Guiding Principles specify what would constitute the objective, well documented, and use the best on Trophy Hunting as a Tool for science available. Adaptive management of enhancement of propagation or survival Creating Conservation Incentives, Ver. quotas and programs based on the results of with regard to authorizing the import of 1.0 (IUCN SSC 2012), to provide useful resource assessments and monitoring is parts or products of P. l. melanochaita, principles, which, considered in essential. The program should monitor including sport-hunted trophies. hunting activities to ensure that quotas and conjunction with our threatened species Therefore, when making a sex/age restrictions of harvested animals are issuance criteria, will aid the Service determination of whether an otherwise met. The program should also generate when making an enhancement finding prohibited activity enhances the reliable documentation of its biological for importation of sport-hunted trophies propagation or survival of P. l. sustainability and conservation benefits. of P. l. melanochaita. This document (e) Accountable and Effective Governance: melanochaita, the Service will examine sets out guidance from experts in the A biologically sustainable trophy-hunting the overall conservation and field on the use of trophy hunting as a program should be subject to a governance management of the subspecies in the tool for ‘‘creating incentives for the structure that clearly allocates management country where the specimen originated responsibilities. The program should account conservation of species and their and whether that management of the for revenues in a transparent manner and habitats and for the equitable sharing of subspecies addresses the threats to the distribute net revenues to conservation and the benefits of use of natural resources’’ subspecies (i.e., that it is based on sound community beneficiaries according to (IUCN SSC 2012, p. 2) and recognizes scientific principles and that the properly agreed decisions. All necessary that recreational hunting, particularly steps to eliminate corruption should be taken management program is actively trophy hunting, can contribute to and to ensure compliance with all relevant addressing the current and longer term biodiversity conservation and more national and international requirements and threats to the subspecies). In that specifically, the conservation of the regulations by relevant bodies such as review, we will evaluate whether the administrators, regulators and hunters. hunted species. import contributes to the overall The SSC document lays out five The Service’s approach to conservation of the species by guiding principles that, considered in enhancement findings for the considering whether the biological, conjunction with our threatened species importation of sport-hunted trophies of social, and economic aspects of a issuance criteria, will aid the Service P. l. melanochaita is consistent with the program from which the specimen was when making an enhancement finding purpose and intent of the Endangered obtained provide a net benefit to the for importation of sport-hunted trophies Species Act. Before we will authorize subspecies and its ecosystem. of P. l. melanochaita: the importation of a sport-hunted The Service will evaluate any (a) Biological sustainability: The hunting trophy, we must determine that the application received that involves P. l. trophy hunting program is managed to melanochaita in the context of program cannot contribute to the long-term decline of the hunted species. It should not ensure the long-term survival of the enhancement of propagation or survival alter natural selection and ecological species. In many parts of the world, permitting in accordance with our function of the hunted species or any other wildlife exists outside of protected areas threatened species permitting species that share the habitat. The program and must share the same habitat and regulations at 50 CFR 17.32 and should not inadvertently facilitate poaching compete with humans living in these issuance criteria for threatened species or illegal trade in wildlife by acting as a cover areas for space and resources. If permits (50 CFR 17.32(a)(2)). These for such illegal activities. The hunting program should also not manipulate the communities that share these resources include, in addition to the general with wildlife do not perceive any permitting criteria in 50 CFR 13.21(b): ecosystem or its component elements in a way that alters the native biodiversity. benefits from the presence of wildlife, (i) Whether the purpose for which the (b) Net Conservation Benefit: The they may be less willing to tolerate the permit is required is adequate to justify biologically sustainable hunting program wildlife. However, under certain removing from the wild or otherwise should be based on laws, regulations, and circumstances, trophy hunting can changing the status of the wildlife sought to scientifically based quotas, established with address this problem by making wildlife be covered by the permit; local input, that are transparent and more valuable to the local communities (ii) The probable direct and indirect effect periodically reviewed. The program should that issuing the permit would have on the produce income, employment, and other and encourage community support for wild populations of the wildlife sought to be benefits to create incentives for reducing the managing and conserving the hunted covered by the permit; pressure on the target species. The program species, as well as other species. (iii) Whether the permit, if issued, would should create benefits for local residents to When evaluating whether the in any way, directly or indirectly, conflict co-exist with the target species and other importation of a trophy of P. l.

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melanochaita would be authorized conservation. In evaluating these CFR 17.11(h) to add P. l. leo subspecies pursuant to 50 CFR 17.32, in accordance factors, we will work closely with the and the P. l. melanochaita subspecies to with our threatened species issuance range countries and interested parties to the List of Endangered and Threatened criteria, we will examine how a obtain the information. By allowing Wildlife as an endangered species and country’s management program for lions entry into the United States of P. l. a threatened species, respectively. This addresses the three main threats that melanochaita trophies from range rule establishes a 4(d) rule for P. l. have led to the decline of the countries that have science-based melanochaita, which implements all of subspecies: Habitat loss, loss of prey management programs, we anticipate the prohibitions and exceptions under base, and human-lion conflict. When that other range countries would be 50 CFR 17.31 and 17.32 and requires a examining a management program and encouraged to adopt and financially threatened species import permit under whether trophies taken as part of that support the sustainable management of 50 CFR 17.32 for the importation of all program meet the issuance criteria, we lions that benefits both the species and P. l. melanochaita specimens. Under the would study a number of factors. Some local communities. In addition to 4(d) rule, the import exemption found of the factors we would consider addressing the biological needs of the in 50 CFR 17.8 for threatened wildlife include whether the program is based subspecies, a scientifically based listed in Appendix II of CITES does not on sound scientific information and management program would provide apply to this subspecies. Therefore, identifies mechanisms that would arrest economic incentives for local through the promulgation of this 4(d) the loss of habitat or increase available communities to protect and expand P. l. rule, the presumption of legality habitat (i.e., by establishing protected melanochaita habitat. provided under section 9(c)(2) of the areas and ensuring adequate protection As stated, under this 4(d) rule any Act for the otherwise lawful importation from human encroachment). We would person wishing to conduct an otherwise of wildlife listed in Appendix II of consider whether the management prohibited activity, including all CITES that is not an endangered species program actively address the loss of the imports of P. l. melanochaita listed pursuant to section 4 of the Act lion’s prey base by addressing poaching specimens, must first obtain a permit does not apply to this subspecies (See: or unsustainable offtake within the under 50 CFR 17.32. As with all permit 4(d) Rule for Panthera leo country. A component of a management applications submitted under 50 CFR melanochaita). plan from which trophy imports would 17.32, the individual requesting Available Conservation Measures meet the issuance criteria would be authorization to import a sport-hunted whether there are government trophy of P. l. melanochaita bears the Conservation measures provided to incentives in place that encourage burden of providing information in their species listed as endangered or habitat protection by private application showing that the activity threatened under the Act include landowners and communities and meets the requirements for issuance recognition of , incentives to local communities to criteria under 50 CFR 17.32. In some requirements for Federal protection, and reduce the incursion of livestock into cases for imports, such as sport-hunted prohibitions against certain practices. trophies, it is not always possible for the protected areas or to actively manage Recognition through listing encourages applicant to provide all of the necessary livestock to reduce conflicts with lions. and results in public awareness and information needed by the Service to We would examine if the hunting conservation actions by Federal and make a positive determination under the component of the management program State governments in the United States, Act to authorize the activity. For the supports all of these efforts by looking foreign governments, private agencies import of sport-hunted trophies of P. l. at whether hunting concessions/tracts and groups, and individuals. melanochaita, the Service will typically are managed to ensure the long-term consult with the range country to the Section 7(a) of the Act, as amended, survival of the lion, its prey base, and extent practicable and other interested and as implemented by regulations at 50 habitat. As stated previously, hunting parties to obtain necessary information. CFR part 402, requires Federal agencies can generate significant economic The Service has the discretion to make to evaluate their actions that are to be benefits if properly conducted. In the required findings on sport-hunted conducted within the United States or looking at whether we would be able to trophy imports of P. l. melanochaita on upon the high seas, with respect to any authorize the import of a trophy under a country-wide basis, although species that is proposed to be listed or the issuance criteria of 50 CFR 17.32, we individual import permits will be is listed as endangered or threatened. would examine if the trophy hunting evaluated and issued or denied for each Because P. l. leo and P. l. melanochaita provides financial assistance to the applicant. While the Service may make are not native to the United States, no wildlife department to carry out enhancement findings for sport-hunted critical habitat is being proposed for elements of the management program trophy imports of P. l. melanochaita on designation with this rule. Regulations and if there is a compensation scheme a country-wide basis, the Service implementing the interagency or other incentives to benefit local encourages the submission of cooperation provision of the Act are communities that may be impacted by information from individual applicants. codified at 50 CFR part 402. Section lion predation. We would also consider We would rely on the information 7(a)(2) of the Act requires Federal how a U.S. hunter’s participation in the available to the Service and may rely on agencies to ensure that activities they hunting program contributes to the information from sources other than the authorize, fund, or carry out are not overall management of lions within a applicant when making a permitting likely to jeopardize the continued country. decision. existence of a listed species or to Management programs for P. l. destroy or adversely modify its critical melanochaita would be expected to Effects of This Rule habitat. If a proposed Federal action address, but are not limited to, This action revises the taxonomic may adversely affect a listed species, the evaluating population levels and trends; classification of the Asiatic lion responsible Federal agency must enter the biological needs of the species; (currently classified as P. l. persica and into formal consultation with the quotas; management practices; legal listed as an endangered species under Service. Currently, with respect to the protection; local community the Act) to P. l. leo based on a lion, no Federal activities are known involvement; and use of hunting fees for taxonomic change. This rule revises 50 that would require consultation.

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Section 8(a) of the Act authorizes the organizations and invited them to information to be the best available provision of limited financial assistance comment on the proposed listing. We scientific and commercial information; for the development and management of received tens of thousands of comments. therefore, we have accepted this programs that the Secretary of the We reviewed all comments we taxonomic change and incorporated this Interior determines to be necessary or received from the public for substantive decision into this document under the useful for the conservation of issues and new information regarding Taxonomy section of this document. As endangered or threatened species in the proposed listing of this species, and a result, our assessment is of the status foreign countries. Sections 8(b) and 8(c) we address those comments below. of the lion species (both P. l. leo and P. of the Act authorize the Secretary to Overall, most commenters supported l. melanochaita), including the lion encourage conservation programs for the proposed listing, but did not provide population in India. foreign listed species, and to provide additional scientific or commercial data (2) Comment: Several peer reviewers assistance for such programs, in the for consideration. We have not included provided updated information on form of personnel and the training of responses to comments that supported population estimates and trends. Based personnel. the listing decision but did not provide on a time trend analysis of scientific Section 9 of the Act and our specific information for consideration. census data for 46 well-monitored implementing regulations at 50 CFR Most of the commenters that did not populations, an overall 43 percent 17.21 and 50 CFR 17.31 set forth a series support the proposed listing were decline in lion populations across of general prohibitions that apply to all affiliated with the trophy hunting Africa was inferred. Furthermore, endangered and threatened wildlife, industry and opposed the rule due to regional trends emerged, showing that, respectively, except where a 4(d) rule potential impacts on importing trophies. while populations in southern African applies to threatened wildlife, in which These comments are addressed below. increased by 22 percent, populations in case the 4(d) rule contains all the eastern and western and central Africa Peer Review applicable prohibitions and exceptions. combined decreased by 57 percent and Under the 4(d) rule for P. l. In accordance with our policy 66 percent, respectively. The peer melanochaita, all of the prohibitions published on July 1, 1994 (59 FR reviewers also indicated that the actual under 50 CFR 17.31 apply to P. l. 34270), we solicited expert opinions number of lions in Africa is much lower melanochaita specimens. These from ten individuals with scientific than previous estimates. Application of prohibitions, at 50 CFR 17.21 and 17.31, expertise that included familiarity with regional trends to lion estimates made in part, make it illegal for any person the species, the geographic region in in 2002 resulted in an estimate of fewer subject to the jurisdiction of the United which wild members of the species than 20,000 lions, a significant States to ‘‘take’’ (includes harass, harm, occur, and conservation biology difference from the previous estimate of pursue, hunt, shoot, wound, kill, trap, principles. We received responses from 32,000. capture, or to attempt any of these) five of the peer reviewers from whom Our Response: We considered this within the United States or upon the we requested comments. The peer information and note that this high seas; import or export; deliver, reviewers generally supported our rule; information was also included in the receive, carry, transport, or ship in however, they provided updated IUCN Red List Assessment for the lion. interstate or foreign commerce, by any information on taxonomy, current Information on population estimates means whatsoever, in the course of population estimates, and population and trends was incorporated into the commercial activity; or sell or offer for trends. They also found our analysis of Species Information section of this sale in interstate or foreign commerce some of the threats to be inaccurate. document. Assessment of this any lion specimens. It also is illegal to Specifically, they provided comments information led us, in part, to conclude possess, sell, deliver, carry, transport, or and additional information on loss of that the status of the lion is more serious ship any such wildlife that has been prey base, trophy hunting, infanticide, than previously indicated, especially in taken in violation of the Act. Permits corruption, and trade in lion bones. In the western and central regions of Africa may be issued to carry out otherwise some cases, a correction is indicated in (P. l. leo). prohibited activities involving the citations by ‘‘personal (3) Comment: One peer reviewer endangered and threatened wildlife communication’’ (pers. comm.), which commented that the section on prey loss species under certain circumstances. could indicate either an email or does not address the issue of prey loss Regulations governing permits for telephone conversation; in other cases, in protected areas where most lions endangered species, such as P. l. leo, are the research citation is provided. occur. codified at 50 CFR 17.22. Regulations Our Response: The peer reviewer Peer Reviewer Comments governing permits for threatened provided a list of literature on the species, such as P. l. melanochaita, are (1) Comment: Several peer reviewers patterns and trends of prey loss in codified at 50 CFR 17.32. Certain commented on our section of the protected areas that were recently or are exceptions apply to agents of the proposed rule regarding the taxonomic currently occupied by lions. We have Service and State conservation agencies. classification of lion. These peer reviewed these articles and have reviewers confirmed that the IUCN Cat incorporated the findings in this Summary of Comments and Specialist Group recommended a two- document (under Loss of Prey Base). Recommendations subspecies classification: Panthera leo This information did not change our We based this action on a review of leo for lions of India and western and determination, but rather further the best scientific and commercial central Africa, and P. l. melanochaita supported our determination that prey information available, including all for lions in eastern and southern Africa. loss has occurred throughout the information received during the public Our Response: We have reviewed the African range countries and is one of the comment period. In the October 2014 2015 IUCN Red List Assessment for the major threats to lion. proposed rule, we requested that all lion, which proposes the new (4) Comment: One peer reviewer interested parties submit information classification as recommended by the stated that although most lions in Africa that might contribute to development of IUCN Cat Specialist Group, and the persist inside protected areas, the a final rule. We also contacted genetic studies supporting this majority of the protected areas should appropriate scientific experts and classification. We found this be uninhabited by humans; therefore,

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only prides located at the edge of these Asiatic lion (Panthera leo persica) is ensure quotas would be based on the protected areas should come into now classified as Panthera leo leo best available science. We have revised conflict with humans. Because the which is now listed as endangered this section accordingly. proportion of lions subjected to conflict under the Act. Scientific samples of P. (13) Comment: Several peer reviewers with humans is small, it is wrong to l. leo will require permits pursuant to 50 commented that the information state that the greatest threat to lions in CFR 17.22. provided in the proposed rule regarding Africa is human-lion conflict. (9) Comment: Several peer reviewers quotas and offtake trends was incorrect; Our Response: We have considered commented that the information several of the peer reviewers provided the peer reviewer’s comments and have provided in the proposed rule regarding additional information (and citations) altered our discussion of threats to lions quotas and offtake trends was incorrect; on country-specific quota trends, from human-lion conflict by clarifying specifically, several peer reviewers current quotas, and offtake trends. One that it is the lions that persist at the noted several publications pertinent to peer reviewer noted that clarification boundary, or just outside, of protected quotas that should be re-examined and was needed regarding the difference areas that are most subjected to this more thoroughly discussed. between quotas and offtake rates. threat. This information did not change Our Response: We reexamined the Additionally, two peer reviewers our determination; human-lion conflict information available to us during the provided additional information on remains a threat to lion persistence. drafting of the proposed rule and moratoriums in two of the range (5) Comment: Three peer reviewers reviewed the citations provided during countries. indicated that our assessment of the public comment period. We Our Response: We reexamined the corruption within lion range countries consider these publications to be the information available to us during the was not realistic; that corruption in best available science regarding quota drafting of the proposed rule and most of Africa is extensive and setting in the interim while other reviewed information in additional worsening. They pointed out oversights strategies are more fully developed (i.e. citations provided during the public and errors pertaining to this subject in age-based strategies, adaptive comment period. We agree that our proposed rule and provided management systems, etc.). We have clarification was needed, and, based additional citations on the topic. revised this section to include more upon peer review comments and Our Response: We reexamined the discussion accordingly. additional information, we have revised information available to us during the (10) Comment: Several peer reviewers this section accordingly. drafting of the proposed rule and provided additional information on (14) Comment: One peer reviewer reviewed information in additional country-specific management trends; commented that lion trophy hunting citations, and agree that our section on specifically, information was provided could remain as an additive threat if corruption did not accurately reflect on the progress of the commitment to hunting reforms are not implemented corruption in lion range countries. and implementation of the age-based and suggested that ‘‘USFWS and Based on peer reviewer comments and strategy. equivalent bodies in the EU and available information, we have revised Our Response: We appreciate the peer elsewhere could mediate such reforms this section accordingly. reviewers input and have incorporated by imposing reduced quotas, best (6) Comment: Two peer reviewers and this information into the section of the practices and the adherence to age three NGO stakeholders indicated rule accordingly. restrictions on countries wishing to concern that trade in lion parts, (11) Comment: One peer reviewer export trophies.’’ particularly lion bone, from Africa to commented that, although species Our Response: It is not appropriate to Asia may pose a potential threat to the experts do generally support trophy establish specific criteria, such as a set species. hunting as a management tool, quota number, in this final rule because Our Response: We agree and have additional discussion was needed this may not allow for the countries to revised this rule to include information regarding the recommended reforms implement an adaptive management on the lion bone trade. species experts submitted during the strategy based on the current status of (7) Comment: A peer reviewer drafting of the proposed rule. the species within the country. During identified inaccuracies in our review of Our Response: We reexamined the the public comment period we received information on traditional use of lion recommendations as provided by new information regarding infanticide parts and products in west and central species experts and agree that additional and the effects of hunting younger male Africa, and also indicated that trade in discussion was needed. We have lions on pride structure. Therefore, we lion parts and products is very common incorporated the additional discussion agree with the peer reviewer that the in these regions. in the section as appropriate. Service is in a position to proactively Our Response: We appreciate the peer (12) Comment: Four of the peer engage with countries to assure reviewer’s input. We reviewed the reviewers commented that although exported trophies fulfill minimum age available information and revised the species experts support trophy hunting requirements, and we will consider section of this rule pertaining to as a management tool, it needs to be these factors in making our traditional use of lion parts and conducted in a sustainable manner that enhancement findings. products in west and central Africa would require reforms to the current (15) Comment: Two peer reviewers accordingly. practices. Peer reviewers stated that the stated that populations in West and (8) Comment: One peer reviewer quotas set throughout most range states Central Africa are small and isolated, questioned whether ‘‘any lion are above sustainable levels (Packer et and, as a result, sustainable offtake was specimen’’ referred to in the 4(d) rule al. 2011) and that quotas should be not possible. Several peer reviewers also would include Asiatic lion and/or science-based and sustainable. provided additional information and scientific samples. Our Response: We agree that current citations on documented lion Our Response: The 4(d) rule applies quotas are currently set higher than population declines resulting from only to the threatened subspecies, P. l. those recommended by Packer et al. excessive lion quotas and poor melanochaita. Scientific samples of P. l. (2011). Species experts recommend the management of trophy hunting. melanochaita will require permits implementation of an adaptive Our Response: We reexamined the pursuant to 50 CFR 17.32. The former management quota system that would information available to us during the

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drafting of the proposed rule and Our Response: We agree that if into our rule accordingly. It should be reviewed the citations provided during hunting concessions maintained noted, however, that, with this finding, the public comment period. We have sustainable levels of harvest, the Panthera leo leo meets our definition of incorporated the new information situation of harvesting males from an endangered species and, therefore, accordingly. neighboring protected areas would not will be subject to the provisions and (16) Comment: One peer reviewer be expected to occur. Species experts regulations of the Act for endangered commented that our review of have recommended best practices for species. Import of sport-hunted trophies infanticide as a result of trophy hunting sustainable development of quotas and of Panthera leo melanochaita will was incomplete and provided additional offtake (Packer et al. 2011, p. 151) while require issuance of a threatened species literature and citation on the subject for other methods are developed (adaptive import permit under 50 CFR 17.32, our consideration. quota management based upon which will require an enhancement Our Response: We agree that scientific data with an enforceable finding (see 4(d) Rule for Panthera leo additional discussion was appropriate monitoring program, (Lindsey et al. melanochaita, above). regarding the impacts of infanticide, (2013a, pp. 8–9) and Hunter et al. (2013, (22) Comment: One commenter noted including a review of the new studies unpaginated)); these recommended that, although the proposed rule offers provided on evolutionary adaptions and reforms have been incorporated as concrete examples of the role of trophy impacts of subadult early dispersal on appropriate. Additionally, based on hunting in lion conservation, the the species. We agree that infanticide information provided during the public proposal offers only limited support of and associated factors relating to trophy comment period, there currently is no trophy hunting benefits. Additionally, hunting of males may have additive level of offtake that would be one commenter notes that the hunting impacts on the decline of certain sustainable in West and Central Africa community has been a leader in lion populations. Therefore, we have at this time. We have incorporated this conservation in terms of habitat incorporated this information into our information into our rule. For Panthera conservation and states that the success final rule. leo melanochaita, we have developed a of certain populations is largely in part 4(d) rule and clarified factors we will to contributions from the hunting Public Comments consider when making an enhancement community. (17) Comment: One commenter noted finding for importation of sport-hunted Our Response: Based on information received during the formation of the that there are very few reliable or trophies of P. l. melanochaita (see 4(d) proposed rule and based on additional scientifically credible lion population Rule for Panthera leo melanochaita, information received during the public surveys in Africa and as a result, quotas above). comment period, we agree that trophy are not scientifically derived. (20) Comment: Several commenters hunting, if managed in a sustainable and Additionally, the commenter noted that stated that populations in West and scientific manner, can provide benefits quota allocations are largely based upon Central Africa are small and isolated to both local communities as well as to concession operators’ opinions. and as a result, sustainable offtake was not possible. Several commenters also lion conservation. We also agree that Our Response: We consider Packer et provided additional information and trophy hunting has conserved a al. (2011) to be the best available science citations on documented lion considerable portion of lion habitat. regarding quota setting in the interim population declines resulting from However, species experts have while other strategies are more fully excessive lion quotas and poor identified several areas across the range developed (i.e., age-based strategies, management of trophy hunting. of the species where hunting has adaptive management systems, etc.). We Our Response: We reexamined the contributed to the decline of lion have re-examined information provided information available to us during the populations. Species experts have during the development of the proposed drafting of the proposed rule and outlined these flaws and have rule and reviewed new information reviewed the citations provided during developed and introduced several provided during the public comment the public comment period. With the recommended reforms to assure that period on quotas, scientific quota new population estimates, in offtake is sustainable and scientific. We development, and adaptive quota combination with the literature and have incorporated these key issues and management systems. As a result, we citations provided during the public the recommended reforms into this rule have incorporated this information into comment period, we agree that given the as appropriate. Although we our rule accordingly. current state of the populations in West acknowledge the role trophy hunting (18) Comment: One commenter noted and Central Africa (Panthera leo leo), has played in lion conservation, we also that the proposed rule addressed only sustainable offtake is not possible. As a have reviewed additional literature CITES Trade Data exports under the result, we have found that, in their provided that documents the decline of ‘‘trophy’’ category and that many are current condition, sustainable offtake lion populations as a result of exported under the ‘‘skins’’ category. for Panthera leo leo is not possible. mismanaged trophy hunting. At this Our Response: We have reviewed the Therefore, we find that trophy hunting time, based on information received U.S. imports of ‘‘skins’’ for 2013 and does rise to a level of threat for Panthera during the public comment period, have incorporated this information into leo leo. We have incorporated the new based on the current trends of lion our rule. information accordingly. populations in West and Central Africa (19) Comment: One commenter states (21) Comment: Several range (Panthera leo leo), experts suggest that that lion trophies exported are almost countries provided additional there is no level of offtake that is exclusively males and subadult males, information on their progress in considered sustainable in these regions. and as such, are targeted by hunters at implementing the best recommended Regardless, import of sport-hunted unsustainable levels. Additionally, the practices and reforms as outlined by trophies of Panthera leo leo will require commenters note that the situation of species experts. issuance of an endangered species harvesting males from neighboring Our Response: We appreciate the import permit under 50 CFR 17.22, protected areas would not be expected information provided by the range which will require an enhancement to occur if the males were being countries. We have incorporated finding. Import of sport-hunted trophies harvested at sustainable levels. relevant portions of this information of Panthera leo melanochaita will

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require issuance of a threatened species hunting regulations. Other commenters technical issues will arise during the import permit under 50 CFR 17.32, note that these reforms have only been implementation phase. Species experts which will require an enhancement fully implemented in some countries have been working through these issues finding (see 4(d) Rule for Panthera leo and additional reforms are needed by providing research and outreach melanochaita, above). throughout the range. An additional materials detailing the most current (23) Comment: Several commenters commenter noted that the information aging techniques, and by providing noted that excessive lion quotas and presented in the proposed rule on range training to concession operators and offtake was the primary driver for countries implementation of best communities (Begg and Begg 2010, pp. declines in lion abundance. practices is overly optimistic with 8, 14; Packer and Whitman 2006, Our Response: We reviewed the new regard to what has actually been entire). We anticipate additional literature provided and agree that the achieved. research will emerge as this strategy is excessive offtake contributed to the Our Response: Several commenters implemented across the species range. decline of some lion populations provided updates regarding the progress (29) Comment: Several commenters throughout their range. We have of range countries’ reforms to hunting noted that the existing age limit for ‘old incorporated this information into our regulations. Although multiple males’ is not enforced. rule and addressed the recommended countries have begun to implement the Our Response: Enforcement of reforms as provided by Hunter et al. reforms as outlined in this document, wildlife crime continues to be an issue (2013, entire) and Lindsey (2013a, pp. only two locations (Mozambique, in for many countries in Africa as 8–9). Niassa Reserve, and Zimbabwe) have evidenced by the rising rate of poaching (24) Comment: Several commenters fully implemented the process and are epidemics and corruption across the noted that current practices, unless completely transparent. However, many African continent. Enforcement of reformed according to best countries are still in the earliest stages trophy hunting regulations across the recommendations, should be considered of implementation, and their progress is range of the species is a critical issue. a potential threat to lion. Species still pending. After a review of this Currently, only two places within the experts recommend a maximum information, we concur that most range African continent have completely science-based offtake of no more than countries have multiple barriers (e.g. implemented the recommendations as <1 lion/2,000 km2 of hunting block until corruption and poverty) that will have set forth in this rule. Several other age restrictions are enforced. to be addressed concurrently with the countries have committed to Our Response: We have reexamined establishment of a transparent and implementing this strategy, but their information provided during the scientific-based, adaptive management progress is currently pending. We must formation of the proposed rule and have system. This information has been note here that enforcement is complex; reviewed new literature submitted incorporated into the rule. Import of it is only one component of a multi- during the public comment period sport-hunted trophies of Panthera leo tiered regulatory system. Successful regarding the best scientific information melanochaita, will require issuance of a enforcement will rely on a variety of available regarding quota setting for threatened species import permit under other factors related to management. lions. We agree and have incorporated 50 CFR 17.32, which will require an Countries will have to address this information in our rule as enhancement finding (see 4(d) Rule for corruption in order to ensure their appropriate. Panthera leo melanochaita, above). monitoring and management systems (25) Comment: Three commenters (27) Comment: One commenter noted are transparent. provided additional information on the that recent scientific knowledge has (30) Comment: During the public biological impacts of trophy hunting. established that hunting males aged five comment period, several commenters New information was provided and older does not affect lion expressed concern that local regarding (1) the evolutionary impacts population dynamics. communities do not actually benefit of selective removal of specimens Our Response: We have reviewed the from the revenue derived from trophy displaying key traits; (2) biological and literature provided and have hunting. Specifically, comments were genetic results of infanticide as it relates incorporated the recommended strategy focused on three issues (see Potential to subadult dispersal and survival; and into our rule. Whitman et al. (2004, pp. Impacts of Trophy Hunting): (1) (3) the role of adult male range and 175–177) found that if offtake is Corruption of concession operators and dispersal requirements in genetic restricted to males older than 6 years of corrupt practices surrounding variation and isolated populations. age, then trophy hunting will likely concession allocation prevent local Our Response: We reexamined the have minimal impact on the pride’s communities from benefitting from information available to us during the social structure and young. Restricting trophy derived revenue; (2) financial drafting of the proposed rule and offtake to males over 6 years of age will contributions to local communities from reviewed the citations and peer review decrease the frequency of male- trophy hunting is often exaggerated and input provided during the public takeovers, and reduce the potential for bears little connection to conservation comment period. We agree that infanticide and delayed infanticide by of the species (local communities additional discussion was required allowing younger males a chance to sire receive only 3–5 percent of revenues); regarding the impacts of infanticide, and raise a cohort of young, and by and (3) that benefits have never been including a review of the studies the allowing the subadults to stay within independently evaluated and commenters submitted. We agree that their pride longer (thus allowing them communities involved in hunting infanticide and associated factors to mature prior to dispersal) (Elliot concessions have not been adequately relating to trophy hunting of males may 2014, p. 1054; Packer et al. 2006, p. 6). surveyed as to their satisfaction of land have additive impacts on the decline of (28) Comment: One commenter stated use for trophy hunting. certain populations. Therefore, we have that the validity of the so-called 6-year Our Response: Corruption occurs incorporated this information into our age approach has been questioned. throughout the range of the species, and final rule. Our Response: The 6-year approach is it likely has an impact on the actual (26) Comment: Several commenters a relatively new development based on benefits received by local communities. noted that many range countries are in research conducted by Whitman (2004, Although many countries have the process of reforming their lion p. 175–177). Like all new concepts, incorporated incentives into their

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trophy hunting policies, land USD annually and employed 38 remote areas lacking infrastructure, management policies, and national lion community members. It is important to attractive scenery, or high densities of action strategies, most countries are still note, however, that the authors are viewable wildlife). Additionally, the in the earliest stages of implementing uncertain regarding the sustainability of commenters state that trophy hunting the strategies discussed in the rule. long-term payments and questioned revenue provides a means of preserving Therefore, we have incorporated this what would happen if the compensation natural habitat despite strong pressure information into our final rule. stopped. In other countries within the to convert habitat into agriculture or (31) Comment: One commenter stated range of lion, systems like these are not rangelands. that there is no evidence to support that necessarily in place. Experts believe the Our Response: We agree that trophy trophy hunting might provide sufficient revenue from trophy hunting, if well hunting revenue provides conservation money to motivate communities in managed in a transparent way, could value at many levels, especially in terms hunting regions to protect lions against potentially fund similar programs of lion habitat, conservation programs, other threats such as retaliatory killings throughout the species’ range, thus anti-poaching programs, equipment, and for livestock losses. reducing retaliatory killings and poaching patrols. However, lion experts Our Response: Although there is benefitting the local population have documented the decline of many limited data on the motivations of simultaneously. populations of lion resulting from individuals who kill lions (see Hazzah (32) Comment: One commenter mismanagement of trophy hunting 2013), we recognize that human-lion suggested non-consumptive uses such (Rosenblatt et al. 2014, p. entire; conflict resulting in retaliatory killing is as eco-tourism could provide the Sogbohossou et al. 2014, entire; Becker a major threat. Although not the only promise of sustainable enterprise. et al. 2013, entire; Lindsey et al. 2013, mechanism for increasing tolerance, Our Response: We agree in part, but entire; Croes et al. 2011, entire; Packer incentives are an important aspect of ecotourism and the trophy hunting 2011, entire; Loveridge et al. 2007, changing individuals’ perceptions of community need to come together to entire). Additionally, the high revenue lions, especially for communities who support the African countries in lion potential associated with trophy live close to lion populations. conservation. Non-consumptive uses of hunting makes it a target for corruption. According to Packer et al. (2011, p. 152, wildlife such as eco-tourism have been As a result, we have reviewed the citing e.g., Baker 1997, Hurt and Ravn practiced in many regions throughout recommended best practices as 2000, Child 2004, Lindsey et al. 2006, Africa. Lindsey et al. (2007, entire) provided by species experts to and Dickson et al. 2009), ‘‘trophy studied viewing preferences among encourage countries to establish a hunting has been considered essential visitors in protected areas in South transparent, science-based, adaptive for providing economic incentives to Africa. Most tourists, especially first- quota management system. Import of conserve large carnivores.’’ For time and foreign visitors, were generally sport-hunted trophies of Panthera leo example, Kenya banned trophy hunting focused on charismatic mega-species leo will require issuance of an in 1977 due to questionable ethics and that are generally confined to protected endangered species import permit under poor management. Since then, ‘‘wildlife areas; African visitors had more interest 50 CFR 17.22, which will require an populations outside of parks have in bird and plant diversity, scenery, and enhancement finding. Import of sport- declined by at least 60%, due partly to other rare species. Lindsey et al. (2007) hunted trophies of Panthera leo the inability of local people to benefit acknowledge that ecotourism may align melanochaita will require issuance of a from wildlife’’ (Lindsey et al. 2006, with conservation objectives and threatened species import permit under citing Child, 2000, 2005). provide incentives for the development 50 CFR 17.32, which will require an Recently, Hazzah et al. (2014, entire) of tour operations geared away from the enhancement finding (see 4(d) Rule for conducted research in Kenya in the ‘big five.’ However, ecotourism as a Panthera leo melanochaita, above). Amboseli ecosystem, where it was replacement to trophy hunting will have (34) Comment: One commenter noted estimated that 55 percent of lion killings to be researched further. Information that that the estimates of revenue from were retaliatory in nature. In this area, provided by Hunter et al. (2013, trophy hunting presented in the two programs are used to provide unpaginated citing Norton-Griffiths proposed rule were not believed to be incentives to locals to prevent these 2007) indicates that ‘‘a significant the best scientific information available. types of killing. First, there is a Predator portion of the land where trophy Specifically, they questioned the Compensation Fund (PCF) wherein hunting occurs is unlikely to be viable objectivity of one source (Jackson 2013) local people are compensated for for alternate wildlife-based land uses and provided additional information depredated livestock and the system is such as photo- or ecotourism due to analyzing Lindsey et al. (2012a). carefully designed with a system of remoteness, lack of infrastructure Our Response: The new literature verification processes, payments, and including integration in established provided by the commenter (Campbell violation penalties (2014, p. 852). tourism circuits, lack of spectacular 2012, entire) identifies some analysis Second, the Lion Guardians (LG) scenery or lack of high densities of and data flaws in Lindsey (2012a). We program uses traditional techniques to viewable wildlife.’’ Additionally, have reviewed the information incorporate community value and belief according to Hunter et al. (2013, presented and updated this rule using systems to improve local perceptions. unpaginated citing Packer et al. 2007; the best available scientific information. According to Hazzah et al. (2014, pp. Groom 2013, pp. 2–3) ecotourism is We have removed information we used 857–858), compensation alone showed a highly dependent on political stability. from Jackson (2013) and Lindsey et al. 73 percent reduction in lion killing. As a result, ecotourism is unlikely to be (2012) and rely upon information from Combining this with the LG program (in able to provide the revenue potential Groom (2013) and Barnett and Patterson 2007) further reduced the decline by 91 that is currently associated with trophy (2005), which was also presented in the percent (less than one killed per year). hunting, although we agree there is proposed rule. Hazzah et al. estimated that the PFC potential for growth in this industry. (35) Comment: One commenter noted program cost an estimated $250,000 (33) Comment: Several commenters that the discussion as presented in the USD annually and employed 30 state that hunting is able to generate proposed rule was biased toward the community members. The LG program revenues for a larger proportion of areas hunting industry and did not discuss was estimated to have cost $140,000 that are unsuitable for ecotourism (e.g., the body of research documenting the

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potential negative impacts of trophy lions lost to diseases. Some populations stated that African lions, because they hunting. A peer reviewer requested a were able to recover to pre-outbreak are currently listed in CITES Appendix more thorough discussion be included levels, but for others, factors such as an II, would be covered by the presumption to address (1) the major flaws in current inbred population prevented provided by section 9(c)(2) if they are management practices, and (2) populations from recovering to pre- listed as threatened. One of the recommendations for how these issues outbreak levels. We found no commenters noted a disparity between can be addressed to account for information indicating the loss of lions the 4(d) rule for lions and a 4(d) rule for sustainability. to disease is a significant driver of the another species that was commonly Our Response: We reexamined the status to the species. However, we hunted. This commenter felt that information available to us during the acknowledge that diseases can debilitate because both species are listed in drafting of the proposed rule and rather than cause mortality, but Appendix II of CITES that their reviewed the citations and peer review debilitation may cause an individual to treatment under the Act should be input provided during the public succumb to other factors. Furthermore, similar. comment period. As a result, we have due to the prevalence of some diseases Our Response: While there has been incorporated this information into the in lion populations and current stressors question as to whether section 9(c)(2) of rule. on lions, it is likely that disease the Act might automatically require (36) Comment: Three range countries contributes to lion mortality. The allowing the importation of a species provided information on the occurrence information provided by the commenter that is both listed as threatened and in of human-lion conflict. All three did not alter our finding that disease is Appendix II, and preclude the issuance countries indicated that human-lion not a significant threat to the species; of more restrictive 4(d) rules covering conflict is a serious problem. however, we have altered the discussion importation, the Service has concluded Our Response: We incorporated this of disease to clarify that disease is a that such 4(d) rules may be issued to information into our discussion of secondary factor that is exacerbated by provide for the conservation of the human-lion conflict. The information other threats the lion faces. involved species. Section 9(c)(2) does further supported our conclusion that (39) Comment: Several commenters not expressly refer to threatened species human-lion conflict constitutes a threat stated that climate change has a or prevent the issuance of appropriate to lion persistence. detrimental impact on the species and 4(d) rules and could not logically have (37) Comment: One commenter agrees that the Service did not incorporate been intended to allow for an that human-lion conflict is a threat to recent climate trend data into our international convention to override remaining lion populations, but asserts analysis. U.S. law, where there is reliable that it does not constitute a level of Our Response: We have incorporated evidence to affect the presumption of threat in eastern and southern Africa to climate change data and its effect on the validity. Finally, the term ‘‘presumed’’ warrant a listing under the Act. The species into our analysis. implies that the established commenter further asserts that the lion (40) Comment: One commenter presumption is rebuttable under certain has been secured from the negative specifically commented that the 4(d) circumstances, including through the impacts of human-lion conflict where rule is appropriate and needed for the promulgation of a protective regulation 90 percent of its population exists and conservation of the species. A second pursuant to section 4(d) of the Act. that human-lion conflict can be commenter applauded the Service for (42) Comment: Two commenters controlled and reduced. recognizing the importance of regulated stated that, even if the Service had the Our Response: We agree that there are hunting and the conservation of the authority to promulgate a regulation that populations of lions where adequate African lion and the need for a system establishes the manner in which African management has reduced the that allows U.S. hunters to import lions are imported, it cannot use the occurrence and impacts of human-lion trophies. regulation to essentially shift to the conflict. However, the best available Our Response: The Service agrees that hunter/importer the burden of proving information indicates that retaliatory the 4(d) rule is necessary and advisable enhancement or survival of the species killing is a rangewide occurrence, and for the conservation of the subspecies P. criteria. given the limited number of lions l. melanochaita. The Service has Our Response: The burden of showing remaining, any loss of lions to recognized that a well-managed, that an ‘‘otherwise prohibited activity’’ retaliatory killing, or other actions, can scientifically based hunting program meets the issuance criteria under 50 have a detrimental impact on the can provide for the conservation of a CFR 17.32 is on the applicant. In some species. species and benefit local communities. cases for imports, such as sport-hunted (38) Comment: One commenter By establishing the 4(d) rule that trophies, it is not always possible for the disagreed with our conclusion that encourages range countries to effectively applicant to provide all of the necessary disease was not a significant threat to manage their lion populations, U.S. information needed by the Service to the lion and provided additional hunters can continue to contribute to make a positive determination under the information on FIV, bTB, and CDV and the long-term conservation of the Act to authorize the activity. For the discussed difficulties in determining the subspecies. import of sport-hunted trophies of P. l. role of disease in lion mortality. The (41) Comment: Four commenters melanochaita, the Service will typically commenter requested that we reconsider stated that the Service lacks the consult with the range country to the our determination based on authority to rebut the Act’s section extent practicable and other interested consequences of diseases to the immune 9(c)(2) with a blanket finding applicable parties to obtain necessary information. system. to lions throughout Africa, for an The Service has the discretion to make Our Response: As mentioned in their indefinite time period. Section 9(c)(2) the required findings on sport-hunted comment, the role of disease in lion states that any importation shall ‘‘be trophy imports of P. l. melanochaita on mortality and reproductive potential is presumed to be an importation not in a country-wide basis, although almost completely unknown in lion violation’’ of any provision of the Act or individual import permits will be populations. Except for a few implementing regulation for species not evaluated and issued or denied for each populations that have been studied, listed as endangered but listed on applicant. While the Service may make there are no estimates of the number of Appendix II of CITES. The commenters enhancement findings for sport-hunted

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trophy imports of P. l. melanochaita on threats that are facing lions within that information, obtained by the Service a country-wide basis, the Service country. during the public comment period, encourages the submission of (44) Comment: Three commenters, a indicates that not all hunting programs information from individual applicants. peer reviewer and comments from a are well managed or provide We would rely on the information consortium of seven range countries felt enhancement to survival of the available to the Service and may rely on that the proposed 4(d) rule did not subspecies. Delaying the information from sources other than the adequately explain the criteria used by implementation of a 4(d) rule may result applicant when making a permitting the Service to determine whether the in U.S. hunters participating in poorly decision. importation of any sport-hunted lion managed hunting programs, which (43) Comment: Two commenters would enhance the survival of the would be counter to the purposes of the stated the Service has offered nothing to species. The commenter expressed Act. We do not agree that such a delay demonstrate why limitations on the concern that because the Service has not would be appropriate for the importation of sport-hunted African adequately explained the criteria for conservation of the subspecies. lions from throughout the subspecies’ enhancement or made an enhancement Regarding the potential loss of deposits range is necessary and advisable to finding for lions in Africa, U.S. hunters for previously booked trophy hunts, provide for the conservation of the will be barred from importing their lion hunters were notified of a potential subspecies or sufficient to overcome the trophy. The peer reviewer expressed a regulatory change when the proposed Congressional conclusion that such need for the Service to elaborate rule with a 4(d) rule was published on imports would normally (i.e., concrete requirements to which October 29, 2014 (79 FR 64472). The presumptively) benefit the conservation countries must adhere as a minimum availability of the proposed rule would of the species. Further, these standard in order for imports of sport- have given hunters the opportunity to commenters did not feel that the hunted lion trophies from a country to use that information to minimize Service’s proposed rule for African lion qualify for the export of lion trophies, financial losses. supported a conclusion that a 4(d) rule including quotas of less than one male (46) Comment: One commenter urged requiring import permits for trophies per 2000 km2 with a minimum age the Service to adjust the rule to ensure was necessary and advisable for the requirement. that imports are not stopped, and that conservation of the subspecies. Our Response: We recognize that the the benefits generated by U.S. hunters in Our Response: For the import of preambular language of the proposed foreign countries continue while the sport-hunted trophies, while there is 4(d) rule was general, and we have Service is making determinations evidence that many of the range addressed this issue in this final rule. regarding the countries’ lion countries have lion management plans, However, we did not find that it was management program. This commenter we have little information indicating appropriate to establish specific criteria, suggested that the Service issue U.S. that the plans are being implemented, such as a set quota number, in this final import permits for all lion trophies until and we received new information rule because this may not allow for the such time as the Service deems that the during the public comment period countries to implement an adaptive import from a particular country would indicating that some hunting programs management strategy based on the not enhance the survival of the are not scientifically based or providing current status of the species within the subspecies. It is the commenter’s belief adequate conservation benefits to the country. During the public comment that there are beneficial aspects of species. We want to encourage U.S. period we received new information hunting (benefits to local communities, hunters to take advantage of one of the regarding infanticide and the effects of dollars coming into the country, etc.) conservation tools available, well- hunting younger male lions on pride that should not be interrupted while the regulated hunting programs, to improve structure. Therefore, we agree with the Service is making its determinations. the long-term survival of the subspecies. peer reviewer that the Service is in a The commenter expressed concern that The 4(d) rule will support implementing position to proactively engage with the Service has insufficient resources to well-managed plans by encouraging countries to ensure exported trophies make timely country-by-country countries that have insufficient lion fulfill minimum age requirements and determinations. management plans to develop plans that we will consider these factors in making Our Response: Import of sport-hunted are based on sound scientific our enhancement findings. trophies of Panthera leo leo will require information that would generate (45) Comment: Two commenters issuance of an endangered species revenue in support of communities and recommended that the Service should import permit under 50 CFR 17.22, conservation. As noted, the proposed not adopt a 4(d) rule until it makes which will require an enhancement 4(d) rule for African lion would provide specific enhancement-of-survival finding. Import of sport-hunted trophies for the importation into the United findings for each of the countries for of Panthera leo melanochaita will States of trophies taken legally in range which lions can be hunted, or delay the require issuance of a threatened species countries upon the issuance of a implementation of the 4(d) rule for 1 import permit under 50 CFR 17.32, threatened species import permit. While year. These two commenters, as well as which will require an enhancement the Service cannot control hunting of a third commenter, stated that finding (see 4(d) Rule for Panthera leo foreign species such as African lion, we implementing the 4(d) rule at this time melanochaita, above). We would be can regulate their importation and would impact hunters who had already unable to issue import permits until we thereby require that U.S. imports of booked trophy hunts months or even made such determinations. The Service sport-hunted African lion trophy years in advance, resulting in the loss of recognizes that making these findings specimens are obtained in a manner that money invested that could not be may be time consuming given our is consistent with the purposes of the recovered ‘‘in the event of a sudden current resources. We appreciate the Act and the conservation of the change in the rules governing the commenter’s willingness to use their subspecies in the wild, by allowing importation of sport-hunted trophies.’’ own resources to obtain information on importation from range countries that Our Response: In the proposed rule, the range countries’ management and have management plans that are based the Service found that hunting, if well assist the Service in making timely on scientifically sound data and are managed, may provide a benefit to the findings. We encourage the commenter being implemented to address the subspecies. However, the best available and others to work with us by

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submitting any information they may (49) Comment: Several lion range U.S. hunters will continue to support have to make these determinations. countries as well as two commenters the good efforts of the range countries, (47) Comment: One commenter stated expressed that successful conservation while encouraging those countries that that the Service should only apply a of African lion relies upon a thoughtful have not fully implemented a lion permitting requirement on lions taken strategy that includes sustainable use. management plan to do so in order to after the listing and 4(d) rule go into There was concern that the inability to receive business from U.S. hunters. effect. import lions into the United States (50) Comment: Several countries and Our Response: For lions held in would result in the increase of threats one commenter provided a combined captivity or a controlled environment on we identified in the proposed rule (e.g., comment expressing concern that the the date of the listing under the Act, no human-lion conflict and habitat loss). Service’s 4(d) rule surpasses the import permit will be required, if the The countries expressed that if U.S. regulatory requirements they are already lion meets all the requirements to be hunters are unable to import sport- following under CITES, and that such considered ‘‘pre-Act’’ (Section 9(b)(1) of hunted trophies, the economic value of restrictions undermine CITES and the Act). Accordingly, lions hunted after lions within the country would be increase the regulatory burden to lion the listing would require permits, and reduced or eliminated, resulting in range states by adding additional those hunters who have booked hunts, retaliatory killing of lions by local reporting requirements. These countries but have not yet hunted a lion, would communities because of real or noted that under CITES exports of require a U.S. import permit prior to perceived perceptions that lions kill trophies must not be detrimental to the importation. people and livestock. In addition, two survival of the species and expressed (48) Comment: Two commenters countries noted that, without an that proving their management stated their belief that most of the lion economic value, safari companies programs enhance the survival of the range countries do not have national would not support lions in hunting subspecies is an added administrative lion conservation plans in place, or have concessions because lions prey upon burden on their wildlife management authorities that are already limited on plans with quotas in place that are other valued trophy species, such as staff, resources, and time. Further, they based on inaccurate population and buffalo. One country felt the 4(d) rule would penalize numbers. One commenter spoke of lion noted that if hunting companies were countries that are already working hard conservation conferences in 2005 and unable to export to the United States, to achieve success in wildlife 2006 that established conference they would abandon their hunting areas to agro-pastoral uses, resulting in management. resolutions, very few of which have Our Response: As these countries been adequately addressed by the lion ‘‘unavoidable extinction of wildlife and collapse of ecosystem services.’’ These noted in their comments, CITES allows range states. This commenter felt there countries expressed that hunting zones for stricter domestic measures, such as is an urgent need to conduct often provide a buffer to protected areas the Act and our 4(d) rule for P. l. independent and scientifically valid as well as provide ecological corridors melanochaita promulgated under the lion population assessments throughout between protected areas. They Act. The Service recognizes that the 4(d) the range of the lion. This commenter expressed that the removal of lions from rule for P. l. melanochaita has stricter urged the Service to impose an import these hunting zones would decrease the requirements than CITES Appendix-II moratorium until these population range of the subspecies and result in requirements. We find that our 4(d) rule assessments have been conducted. The overall lion population declines. for P. l. melanochaita will support and second commenter recommended that Further, the loss of legal income from encourage countries to carry out strong prior to the import of trophies, there lion hunting, which supports anti- conservation programs for P. l. needs to be evidence of recovery and poaching efforts, will negatively affect melanochaita and ensure that U.S. stability, as well as clearly identified lion conservation and increase imports of sport-hunted lion trophy governmental reforms and their poaching. specimens are obtained in a manner that implementation in some of the range Our Response: The Service recognizes is consistent with the purposes of the states. the benefits that a well-managed trophy Act and the conservation of the P. l. Our Response: New information hunting program can provide by melanochaita in the wild. We do not received during the public comment increasing revenue for local anticipate a significant burden on the period raises questions about whether communities, providing jobs, and lion range countries to provide some of the range countries have supporting local microbusinesses. documentation that should already exist adequate management programs in Revenue is often used to build and for well-managed lion programs, and we place, and this information has been maintain fences, pay for security will work with the countries in order to incorporated in this final rule. The personnel, and provide resources for make our determinations under the Act Service is not imposing a moratorium; anti-poaching activities, habitat in a timely manner. The 4(d) rule is in however, permits will be required for all acquisition, and wildlife management. place to support countries that have imports. Import of sport-hunted trophies Our 4(d) rule for P. l. melanochaita achieved success in managing their of Panthera leo leo will require issuance will support and encourage lions. of an endangered species import permit conservation actions for this subspecies (51) Comment: Several countries and under 50 CFR 17.22, which will require and ensure that U.S. imports of sport- one commenter disagreed with how an enhancement finding. Import of hunted lion trophy specimens are trade in captive-bred lions would be sport-hunted trophies of Panthera leo obtained in a manner that is consistent subject to the prohibitions under the melanochaita will require issuance of a with the purposes of the Act and the Act. These countries expressed that threatened species import permit under conservation of the P. l. melanochaita in trade in captive-bred lion does not have 50 CFR 17.32, which will require an the wild. By ensuring that imports of an adverse effect on wild lion enhancement finding (see 4(d) Rule for lions occur only from range countries populations. They felt that the Act’s Panthera leo melanochaita, above). The that have management plans based on treatment of captive lions in the same import of lions hunted in countries that scientifically sound data which are manner as wild lions is inconsistent do not meet the criteria for being implemented to address the with CITES regulations and that the 4(d) enhancement will not be permitted. threats facing lions within that country, rule should exempt captive-bred lions.

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Our Response: In analyzing threats to addressed in the preambular language of Our Response: We have determined the species, we focused our analysis on the 4(d) rule. that we do not need to prepare an threats acting upon wild specimens (53) Comment: The petitioners also environmental assessment, as defined within the native range of the species, asserted that we should not authorize under the authority of the National because the goal of the Act is survival imports of lions from western Africa, Environmental Policy Act of 1969, in and recovery of the species within its Tanzania or Zimbabwe; imports of connection with regulations adopted native ecosystem. We did not separately trophies from females or males under 6 under section 4(a) of the Act. We analyze ‘‘threats’’ to captive-held years of age; or trophies obtained from published a notice outlining our reasons specimens because the statutory five captive-hunting facilities, or authorize for this determination in the Federal factors under section 4 (16 U.S.C. 1533) imports, interstate commerce or foreign Register on October 25, 1983 (48 FR are not well-suited to consideration of commerce in lion parts. 49244). Furthermore, under our 1983 specimens in captivity and captive-held Our Response: While the comments policy, we determined that we do not specimens are not eligible for separate are outside the scope of this rulemaking, need to prepare an environmental consideration for listing. However, we the Service must make a finding that an assessment in connection with did consider the extent to which ‘‘otherwise prohibited activity,’’ such as regulations adopted under section 4(a) specimens held in captivity create, import, export, interstate and foreign of the Act, including 4(d) rules that contribute to, reduce, or remove threats commerce, must meet the issuance accompany listings of threatened to the species. See the Captive Lions criteria under 50 CFR 17.32. We cannot species. and Traditional Use of Lion Parts and make any determination of whether a Because we are listing P. l. Products sections above. Under CITES, particular permit application can be melanochaita as threatened and are captive specimens are still listed the approved or denied until the finalizing this 4(d) rule simultaneously same as their wild counterparts; application is reviewed. with our final listing determination, we however, the Convention does allow for (54) Comment: One commenter called consider this 4(d) rule to be part of the different treatment of captive-bred on the Service to specifically prohibit listing determination for the purposes of specimens in regard to permitting. As the importation of sport-hunted lions in National Environmental Policy Act stated earlier, CITES also provides for the 4(d) rule, citing that there is no compliance. stricter domestic measures, and the documented evidence that trophy (56) Comment: One commenter stated protections afforded to all specimens of hunting supports conservation of the that lions do not lend themselves to the subspecies through listing under the subspecies. In addition, the commenter population surveying due to the boom ESA and the 4(d) rule would constitute felt that allowing for legal trade of sport- and bust nature and high fecundity of such a measure. hunted lions would support the illegal lion populations. The commenter felt (52) Comment: A joint comment from harvest of the subspecies. that population surveys have long been the petitioners asked us to scrutinize Our Response: We found no evidence considered impractical, and as such, applications for the import of lion that allowing legal import of lion quotas can never be set scientifically trophies or parts to ensure that they trophies would stimulate illegal trade and, therefore questioned how the were obtained within a scientifically into the United States. In evaluating the Service can make this a criteria for based management program that best available scientific and commercial determining enhancement. Finally, the promotes the conservation of the information, we concluded that a well- commenter was concerned that having subspecies and provided suggestions for managed, scientifically based lion countries have an understanding of lion criteria to consider when making an management program can provide a population numbers and developing enhancement finding. The comment benefit to the species. While we lion management plans would be cost included a number of suggestions for obtained new information indicating prohibitive to many of the range establishing a formal internal guidance that some hunting programs are not countries. on how we would evaluate each scientifically based or providing Our Response: We are not requiring application. Finally, the petitioners adequate conservation benefits to the an exact count of the lions within each called on the Service to publish the species, this 4(d) rule will support country before being able to make a receipt of threatened species permit implementing well-managed plans by determination of whether imports could applications in the Federal Register and encouraging countries that have occur. However, we need to consider allow for a 30-day comment period. insufficient lion management plans to what methods countries are using to Another commenter questioned develop plans that are based on sound establish quotas, such as population establishing findings on a country-wide scientific information that would trend data, in order to determine if the basis instead of specific regions/hunting generate revenue in support of offtake by U.S. hunters is sustainable programs within a country. communities and conservation. and meets the criteria under 50 CFR Our Response: We appreciate the Therefore, we are not prohibiting the 17.32. input regarding publishing the receipt of import of sport-hunted trophies. Import (57) Comment: One commenter stated threatened species applications, of sport-hunted trophies of Panthera leo that lions have an extraordinary high establishing formal internal guidance on melanochaita will require issuance of a fecundity, which contributes to its boom how we will evaluate each application, threatened species import permit under or bust population characteristic and and consideration of making 50 CFR 17.32, which will require an helps ensure its long-term existence, enhancement findings on a specific enhancement finding (see 4(d) Rule for making it far less vulnerable to region/hunting program scale. We will Panthera leo melanochaita, above). The endangerment. consider these suggestions; however, import of lions hunted in countries that Our Response: We agree that lions this issue is outside the scope of this do not meet the criteria for have high fecundity and in absence of rulemaking process. In regard to the enhancement will not be permitted. stressors populations can rapidly suggested criteria for making (55) Comment: One commenter stated increase. However, across most of its enhancement findings, we have that the Service has failed to comply range, the lion is not without stressors, expanded the discussion of with the National Environmental Policy and given the threats the lion is enhancement within this final rule, and Act (NEPA) in regard to promulgating currently facing, natural fecundity is many of the suggestions have been the 4(d) rule. reduced. One of the greater stressors on

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lions, excessive harvests of lions for Required Determinations recordkeeping requirements, trophies, can negatively impact the National Environmental Policy Act (42 Transportation. reproduction of a lion such that it U.S.C. 4321 et seq.) causes local extirpations. Harvesting Regulation Promulgation We have determined that we do not males that are too young causes male Accordingly, we amend part 17, replacements, which results in need to prepare an environmental assessment, as defined under the subchapter B of chapter I, title 50 of the increased infanticide rates, death of the Code of Federal Regulations, as set forth surviving male coalition, and a 100 authority of the National Environmental below: percent fatality rate for males that are Policy Act of 1969, in connection with regulations adopted under section 4(a) prematurely forced to disperse. PART 17—[AMENDED] Furthermore, the population will be of the Act for the listing, delisting, or driven to extinction as female reclassification of species. We published ■ 1. The authority citation for part 17 populations collapse as they eventually a notice outlining our reasons for this continues to read as follows: are unable to mate. The species is determination in the Federal Register largely not able to rapidly recover from on October 25, 1983 (48 FR 49244). Authority: 16 U.S.C. 1361–1407; 1531– population declines. This is evidenced References Cited 1544; and 4201–4245; unless otherwise by long-term population trends that noted. A list of all references cited in this indicate an overall 43 percent decline in document is available at http:// lions over 21 years and higher regional ■ 2. Amend § 17.11(h), the List of www.regulations.gov at Docket No. rates of decline in western and eastern Endangered and Threatened Wildlife, FWS–R9–ES–2012–0025, or upon Africa. by: request from the U.S. Fish and Wildlife (58) Comment: One commenter stated Service, Endangered Species Program, ■ a. Removing the entry for ‘‘Lion, that the Service should use its power to Branch of Foreign Species (see FOR Asiatic (Panthera leo persica)’’; and list Distinct Population Segments FURTHER INFORMATION CONTACT). ■ b. Adding entries for ‘‘Lion (Panthera (DPSs), rather than the entire African leo leo)’’ and ‘‘Lion (Panthera leo lion subspecies in light of the recent Authors melanochaita)’’ in alphabetic order ruling in Humane Society of the United The primary authors of this rule are States v. Jewell, No. CV 13–186 (BAH), under MAMMALS to read as set forth staff of the Branch of Foreign Species, below: 2014 WL 7237702 (D.D.C. Dec. 19, Ecological Services, U.S. Fish and 2014).. Wildlife Service. § 17.11 Endangered and threatened Our Response: We disagree with this wildlife. conclusion. Pursuant to 50 CFR List of Subjects in 50 CFR Part 17 * * * * * 17.11(g), all populations are included in Endangered and threatened species, the listing. Exports, Imports, Reporting and (h) * * *

Species Vertebrate population where Critical Special Historic range endangered or Status When listed habitat rules Common name Scientific name threatened

MAMMALS

******* Lion ...... Panthera leo leo ..... Africa, Asia ...... Entire ...... E 862 NA NA Lion ...... Panthera leo Africa ...... Entire ...... T 862 NA 17.40(r) melanochaita.

*******

* * * * * (2) The import exemption found in (3) All applicable provisions of 50 ■ 3. Amend § 17.40 by adding paragraph § 17.8 for threatened wildlife listed in CFR parts 13, 14, 17, and 23 must be (r) to read as follows: Appendix II of the Convention on met. International Trade in Endangered Dated: December 10, 2015. § 17.40 Special rules—mammals. Species of Wild Fauna and Flora * * * * * (CITES) does not apply to this Daniel M. Ashe, (r) Lion (Panthera leo melanochaita). subspecies. A threatened species import Director, Fish and Wildlife Service. (1) General requirements. All permit under § 17.32 is required for the [FR Doc. 2015–31958 Filed 12–21–15; 4:15 pm] prohibitions and provisions of §§ 17.31 importation of all specimens of BILLING CODE 4333–15–P and 17.32 apply to this subspecies. Panthera leo melanochaita.

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