[Federal Register Volume 80, Number 246 (Wednesday, December 23, 2015)]
[Rules and Regulations]
[Pages 79999-80056]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2015-31958]



[[Page 79999]]

Vol. 80

Wednesday,

No. 246

December 23, 2015

Part II





Department of the Interior





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Fish and Wildlife Service





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50 CFR Part 17





Endangered and Threatened Wildlife and Plants; Listing Two Lion 
Subspecies; Final Rule

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

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DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

[Docket No. FWS-R9-ES-2012-0025; 450 003 0115]
RIN 1018-BA29


Endangered and Threatened Wildlife and Plants; Listing Two Lion 
Subspecies

AGENCY: Fish and Wildlife Service, Interior.

ACTION: Final rule.

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SUMMARY: We, the U.S. Fish and Wildlife Service (Service), determine 
endangered status for the lion subspecies Panthera leo leo and 
threatened status for P. l. melanochaita under the Endangered Species 
Act of 1973, as amended (Act). We are also publishing a concurrent rule 
under section 4(d) of the Act. This rule provides for conservation 
measures for P. l. melanochaita.

DATES: This rule is effective January 22, 2016.

ADDRESSES: This final rule is available on the Internet at http://www.regulations.gov and comments and materials received, as well as 
supporting documentation used in the preparation of this rule, will be 
available for public inspection, by appointment, during normal business 
hours at: U.S. Fish and Wildlife Service; 5275 Leesburg Pike; Falls 
Church, VA 22041.

FOR FURTHER INFORMATION CONTACT: Branch of Foreign Species, Ecological 
Services, U.S. Fish and Wildlife Service, MS: ES, 5275 Leesburg Pike, 
Falls Church, VA 22041-3803; telephone, 703-358-2171; facsimile, 703-
358-1735. If you use a telecommunications device for the deaf (TDD), 
call the Federal Information Relay Service (FIRS) at 800-877-8339.

SUPPLEMENTARY INFORMATION:

Executive Summary

I. Purpose of the Regulatory Action

    We are listing two subspecies of lion, Panthera leo leo and P. l. 
melanochaita, under the Endangered Species Act of 1973, as amended 
(Act). We are listing the P. l. leo subspecies as an endangered species 
and the P. l. melanochaita subspecies as a threatened species under the 
Act. We are also finalizing a rule under section 4(d) of the Act that 
will provide for conservation measures for P. l. melanochaita.

II. Major Provision of the Regulatory Action

    This action revises the taxonomic classification of the Asiatic 
lion (currently classified as P. l. persica and listed as an endangered 
species under the Act) to P. l. leo based on a taxonomic change. The P. 
l. leo subspecies will be listed as an endangered species and the P. l. 
melanochaita subspecies will be listed as a threatened species in the 
List of Endangered and Threatened Wildlife in title 50 of the Code of 
Federal Regulations (CFR) at 50 CFR 17.11(h). This action will also add 
a rule under section 4(d) of the Act for P. l. melanochaita which is 
set forth at 50 CFR 17.40(r).

Background

    The Endangered Species Act of 1973, as amended (Act) (16 U.S.C. 
1531 et seq.), is a law that was passed to prevent extinction of 
species by providing measures to help alleviate the loss of species and 
their habitats. Before a plant or animal species can receive the 
protection provided by the Act, it must first be added to the Federal 
List of Endangered and Threatened Wildlife or the Federal List of 
Endangered and Threatened Plants in part 17 of title 50 of the Code of 
Federal Regulations (CFR). Section 4 of the Act and its implementing 
regulations at 50 CFR part 424 set forth the procedures for adding 
species to these lists.

Previous Federal Actions

    In a final rule published in the Federal Register on June 2, 1970 
(35 FR 8491), the Asiatic lion (currently listed under the Act as 
Panthera leo persica) was listed under the Act's precursor, the 
Endangered Species Conservation Act of 1969, as an endangered species 
and has remained listed as an endangered species under the Act.
    On March 1, 2011, we received a petition dated the same day from 
the International Fund for Animal Welfare, the Humane Society of the 
United States, Humane Society International, the Born Free Foundation/
Born Free USA, Defenders of Wildlife, and the Fund for Animals 
requesting that the African lion subspecies be listed as endangered 
under the Act. The petition identified itself as such and included the 
information as required by 50 CFR 424.14(a). On November 27, 2012, we 
published a ``positive'' 90-day finding (77 FR 70727) indicating that 
we would initiate a status review of the African lion.
    On October 29, 2014 (79 FR 64472) we published in the Federal 
Register a finding that listing the African lion subspecies (Panthera 
leo leo) as a threatened species was warranted and proposed to list the 
subspecies as a threatened species under the Act. We also proposed a 
rule under section 4(d) of the Act to provide conservation measures for 
the African lion.

Summary of Changes From the Proposed Rule

    We fully considered comments from the public and the peer reviewers 
on the proposed rule to determine our final listing status of lion. 
This final rule incorporates changes to our proposed rule based on the 
comments we received that are discussed under Summary of Comments and 
Responses and newly available scientific and commercial information 
that became available after the close of the comment period. We accept 
the taxonomy as recommended by the International Union for Conservation 
of Nature (IUCN) Species Survival Commission Cat Classification Task 
Force: P. l. leo (Asia and western, central, and northern Africa) and 
P. l. melanochaita (southern and eastern Africa). Here we evaluate the 
status of the lion species (P. leo), which includes the previously 
unreviewed population of P. l. leo in India (formerly P. l. persica). 
Additionally, we have incorporated new population estimates and 
population trends for the lion into our Species Information section.
    Based on comments by peer reviewers and others, we revised the 
section on trophy hunting, providing additional information on the 
practices that experts have identified as undermining the 
sustainability of trophy hunting, recommended best practices and 
reforms, biological impacts of trophy hunting on lion populations, and 
corruption in range countries, and expanded our assessment of the level 
of threat that trophy hunting presents to the species. Additionally, we 
have incorporated information on infanticide, corruption, traditional 
use of lion parts and products, disease, and climate change. Under the 
discussion of the 4(d) rule in the preamble, we further clarify factors 
we will consider when making an enhancement finding for importation of 
sport-hunted trophies of P. l. melanochaita.
    Based on the information we received and our assessment of that 
information, we have altered our finding. Some of the information we 
received indicated threats may be worse than previously indicated. Due 
to significant differences in the impacts of threats within the 
species, we found that P. l. leo and P. l. melanochaita qualify for 
different statuses under the Act.

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Species Information

Taxonomy

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

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[GRAPHIC] [TIFF OMITTED] TR23DE15.000

    As required by the Act, and as explained in our proposed rule, we 
base our listing determinations on the best available scientific and 
commercial information. We accept the taxonomy as recommended by the 
IUCN Cat Classification Task Force, which is supported by mtDNA 
analysis, as well as analysis of nDNA sequence and microsatellite 
variation: P. l. leo (Asia and western, central, and northern Africa) 
and P. l. melanochaita (southern and eastern Africa) (Figure 1) as the 
best available scientific and commercial information. Because this new 
classification for lion includes subspecies whose ranges span two 
continents, we assessed the status of the entire lion species (P. leo).
    Currently, the Asiatic lion (P. l. persica) is listed as an 
endangered species under the Act. Based on the new taxonomic 
classification for lions, we are revising the List of Endangered and 
Threatened Wildlife at 50 CFR 17.11(h). In the Regulation Promulgation 
section of this document, we implement a taxonomic change by removing 
the invalid subspecies P. l. persica. This entity is now included in 
the assessment of the lion species (P. leo).

Species Description

    The lion is the second-largest extant cat species (second in size 
only to the tiger) and the largest carnivore in Africa (Ray et al. 
2005, p. 67). As with other widely distributed large cats, there is 
considerable morphological variation within the species as a result of 
sexual selection, regional environmental adaptations, and gene flow 
(Mazak 2010, p. 194). These include, among others, variation in size, 
coat color and thickness, mane color and form, and skull 
characteristics (Mazak 2010, p. 194, citing several sources; Hollister 
1917, in Dubach 2005, p. 15). They are described in the Convention on 
International Trade in Endangered Species of Wild Fauna and Flora 
(CITES) Periodic Review of the Status of African Lion Across Its Range 
(CITES 2014, p. 3) as follows:

    Characteristics include sharp, retractile claws, a short neck, a 
broad face with prominent whiskers, rounded ears and a muscular 
body. Lions are typically a tawny color with black on the backs of 
the ears and white on the abdomen and inner legs. Males usually have 
a mane around the head, neck and chest. Lions are sexually 
dimorphic, with males weighing about 20-27 percent more than 
females. Adult males, on average, weigh about 188 kilograms (kg) 
(414 pounds (lbs)) with the heaviest male on record weighing 272 kg 
(600 lbs). Females are smaller, weighing, on average, 126 kg (278 
lbs). The male body length, not including the tail, ranges from 1.7 
meters (m) to 2.5 m (5.6 feet (ft to 8.2 ft) with a tail from 0.9 m 
to 1 m (3 ft to 3.2 ft) (Nowell and Jackson, 1996).

    Lions in India tend to be smaller than those in Africa. Adult males 
weigh between 160-190 kg (353-419 lb), while females weigh between 110-
120 kg (243-265 lb) (Chellam in litt. in Nowell and Jackson 1996, p. 
37). The record total length for a male lion in India, including the 
tail, is 2.92 m (9.6 ft) (Sinha 1987 in Nowell and Jackson 1996, p. 
37). One characteristic unique to lions in India is a longitudinal fold

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of skin that runs along the belly (O'Brien et al. 1987, p. 100). 
Additionally, male lions in India do not have as large and full a mane 
as those in Africa, allowing their ears to always be visible, whereas 
the manes of male lions in Africa completely hide the ears (Nowell and 
Jackson 1996, p. 37; O'Brien et al. 1987, p. 100).

Habitat

    Historically, the species occurred in all habitats in Africa, 
except rainforest and the hyper-arid interior of the Sahara (Ray et al. 
2005, p. 66). Today they are found primarily in savannas, although 
there are some remnant populations in other habitat types (Riggio et 
al. 2013, p. 19). According to Nowell and Jackson (1996, p. 19), 
optimal habitat appears to be open woodlands and thick bush, scrub, and 
grass complexes, where sufficient cover is provided for hunting and 
denning. The highest lion densities are reached in savanna woodlands 
plains mosaics of southern and eastern Africa (Ray et al. 2005, p. 66). 
The species is intolerant of anthropogenic (human-caused) habitat 
conversion, such as farming or overgrazing by livestock (Ray et al. 
2005, p. 66). In India, the lion occurs in dry deciduous forests (Meena 
et al. 2014, p. 121). Moist mixed and mixed forest habitats are 
critical to lions as they seek moist shady habitats that provide 
shelter from the heat and cover to hide during peak times of human 
activities (Jhala et al. 2009, p. 3391).

General Biology

    Lions are well studied. Much information exists on habits, 
behavior, and ecology of lions in Africa. CITES (2014, p. 3) provides a 
general overview as follows:

    Lions are generalist, cooperative hunters, with foraging 
preferences changing with season and with lion group size. Lions 
live in groups called ``prides,'' which are ``fission-fusion'' 
social units with a stable membership that sometimes divide into 
small groups throughout the range. Lions have no fixed breeding 
season. Females give birth every 20 months if they raise their cubs 
to maturity, but the interval can be as short as 4-6 months if their 
litter is lost. Gestation lasts 110 days, litter size ranges 1-4 
cubs, and sex ratio at birth is 1:1. At about 4 years of age, 
females will have their first litter and males will become resident 
in a pride. Pride takeovers by male lions and subsequent infanticide 
of cubs sired by the ousted male lions greatly influences 
reproductive success. Lionesses defending their cubs from the 
victorious males are sometimes killed during the takeover. 
Infanticide accounts for 27 percent of cub mortality. Adult 
mortality is typically caused by humans, starvation, disease, or 
attacks from other lions. Injury and death can also occur during 
hunting attempts on some of their larger prey.

Haas et al. (2005, entire) provide a summary of information on lion, 
including the following:

    Prides vary in size and structure, but typically contain 5-9 
adult females (range, 1-18), their dependent offspring, and a 
coalition of 2-6 immigrant males (Heinsohn and Packer 1995; Packer 
et al. 1991). . . . Pride sizes are smallest in arid environments 
with limited prey species (Elliott and Cowan 1977; Hanby and Bygott 
1979; Ruggiero 1991; Schaller 1972; Stander 1992b; Wright 1960). . . 
. Males reside in a pride for [approximately] 2 years before being 
replaced by another group of males (Packer et al. 1988). . . . In 
the absence of a pride takeover, males generally leave their natal 
pride when 2-4 years old (Bertram 1975b; Pusey and Packer 1987). 
Most females are incorporated into their natal prides (Pusey and 
Packer 1987; Van Orsdol et al. 1985). . . . A small proportion of 
lions is nomadic, including young and adult males without a pride. 
Nomadic lions follow the migrations of prey and hunt and scavenge 
cooperatively (Bertram 1975a; Bygott et al. 1979; Schaller 1968, 
1969; Van Orsdol et al. 1985).
    . . . Lion productivity (measured as number of surviving cubs) 
is limited by food. . . . Cub mortality is high in lions and is 
linked to periods of prey scarcity and infanticide by male lions 
during pride takeovers (Packer and Pusey 1983b; Schaller 1972; Van 
Orsdol et al. 1985; Whitman and Packer 1997).
    . . . Lions are mainly active at night. . . . [They] usually 
hunt in groups; males hunt less frequently than do females, but 
males are stronger and can gain access to kills made by females 
(Bertram 1975a; Scheel and Packer 1991). Prey selection is related 
to seasonal weather patterns and the migration of large herbivores 
in some parts of Africa (Hanby et al. 1995). . . . Lions exhibit 
individual preferences in prey selection within and between prides 
in the same area (Rudnai 1973b; Van Orsdol 1984).

    Lion prides in India tend to be smaller than those in Africa; most 
prides in India contain an average of two females, with the largest 
having five. Coalitions of males will defend home ranges that contain 
one or more groups of females, but unlike lions in Africa, in India 
male lions only associate with pride females when mating or on a large 
kill (Meena 2009, p. 7; Nowell and Jackson 1996, p. 37). Females are 
approximately 4 years old at first reproduction, males 5-8 years 
(Banerjee and Jhala 2012, p. 1424; Nowell and Jackson 1996, p. 37). 
Banerjee and Jhala (2012, p. 1424) found that mating occurred 
throughout the year, but mostly in winter. Gestation lasts 110 days; 
births peaked in the summer (April-May). Average litter size is 2.5 
cubs, but as many as 5 have been observed (Banerjee and Jhala 2012, pp. 
1424, 1427; Nowell and Jackson 1996, p. 37). Lion reproduction in India 
appears to coincide with the fawning peak of chital deer (Axis axis) 
between December and January or with the rutting season of chital and 
peak fawning for sambar deer (Cervus unicolor) between May and June. 
Breeding lionesses may cue into these times of increased availability 
of food sources to time births for maximum survival of cubs (Banerjee 
and Jhala 2012, p. 1427). Average interbirth interval is estimated to 
be 1.37 years; however, if cubs of the previous litter survived to 
independence, it could be higher. After territorial takeovers and 
infanticides, females mated within an average 4.8 months (Banerjee and 
Jhala 2012, p. 1424). Banjeree and Jhala (2012, p. 1424) found that the 
major cause of cub mortality is infanticide due to territorial 
takeovers by adult males. Most observed adult mortalities (54.5 
percent) were due to natural causes and 43 percent were due to human 
causes; remaining mortalities were due to unknown causes.

Diet and Prey

    Lions are opportunistic hunters and scavengers. As scavengers, 
lions are dominant and can usually readily displace other predators 
from their kills (Packer 1986, Schaller 1972, in Haas et al. 2005, pp. 
4-5). As hunters, they are known to take a variety of prey. However, 
they are also the largest carnivore in Africa and, as a result, require 
large prey to survive. Ray et al. (2005, pp. 66-67) summarizes lion 
prey in Africa as follows:

    Lions are generalists and have been recorded to consume 
virtually every mammal species larger than 1 kg in their range, as 
well as a wide variety of larger reptiles and birds (Nowell & 
Jackson 1996; Sunquist & Sunquist 2002). The constraints of large 
physical size and extended social groups, however, bind them to 
large-bodied prey, and their diet is dominated by medium-large 
ungulates. In fact, only a few species of large ungulates comprise a 
majority of their diet wherever they occur (Schaller 1972; Stander 
1992; Packer et al. 1995), and they are unable to persist in areas 
without large-bodied prey. The threshold of this requirement is 
perhaps represented at Etosha National Park, Namibia, where Stander 
(1992) showed that lions hunting in pairs met their minimum 
requirements hunting springboks which, at < 50 kg, are the smallest 
preferred prey species recorded.

    In India, the lion's diet is comprised of both small and medium 
prey, as well as vulnerable livestock (Meena et al. 2011, p. 61; Singh 
and Gibson 2011, p. 1753; Meena 2009, p. 8). The most commonly taken 
species is chital, which weighs approximately 50 kg (110 lb), and a 
larger species, the sambar deer

[[Page 80004]]

(Meena et al. 2011, p. 63; Nowell and Jackson 1996, p. 37). The smaller 
size of the prey available in India may be responsible for the smaller 
lion group sizes and less interaction between male and female groups 
(Meena 2009, p. 8; Nowell and Jackson 1996, p. 37). Historically, 
domestic cattle also constituted a major portion of the lion's diet 
(Nowell and Jackson 1996, p. 37) and remains a significant portion 
today (Meena et al. 2011, pp. 63, 64; Singh and Gibson 2011, pp. 1753-
1754). The proportion of wild prey and domestic livestock in a lion's 
diet may vary by season and between protected areas and peripheral 
areas (Meena et al. 2011, pp. 64, 65).
    Prey availability affects the reproduction, recruitment, and 
foraging behavior of lions and, as a result, strongly influences lion 
movements, abundance, and population viability (Winterbach et al. 2012, 
p. 7, citing several sources). Lion densities are directly dependent on 
prey biomass (Van Orsdol et al. 1985, in Packer et al. 2013, p. 636; 
Hayward et al. 2007, entire). In Africa, lion densities range from 8-13 
lions per 100 square kilometers (km\2\) in Selous Game Reserve and up 
to 18 per 100 km\2\ in protected areas of eastern Africa and South 
Africa (Creel and Creel 1997, Nowell and Jackson 1996, in Haas et al. 
2005, p. 4). In India, densities are estimated to be 15 lions per 100 
km\2\ in Gir Protected Area, 6 per 100 km\2\ in Girnar Wildlife 
Sanctuary, and 2 per 100 km\2\ in the surrounding agro-pastoral land 
(Banerjee and Jhala 2012, p. 1421; Banerjee et al. 2010, p. 249). Aside 
from human-related mortality, prey availability is likely the primary 
determinant of lion density in Africa (Fuller and Sievert 2001, in 
Winterbach et al. 2012, p. 7). In areas of low natural prey density, or 
high human contact, lions may prey on livestock (see Human-Lion 
Conflict).

Movements/Home Range

    Availability of prey is perhaps the primary factor that determines 
the ranging behavior of large carnivores (Gittleman & Harvey 1982, Van 
Orsdol et al. 1985, Grant et al. 2005, Hayward et al. 2009, in 
Winterbach et al. 2012, p. 4). Home-range sizes of lion prides 
correlate with lean-season prey biomass (Van Orsdol et al. 1985, in 
Haas et al. 2005, p. 4) and, therefore, vary widely among habitats. 
Average range sizes of lion prides in Africa are 26-226 km\2\, but can 
be considerably larger (Stander 1992b; Van Orsdol et al. 1985; Viljoen 
1993, in Haas et al. 2005, p. 4). In areas of low or variable prey 
biomass, annual range requirements for a single lion pride can exceed 
1,000 km\2\ (Packer et al. 2013, p. 636). Funston (2011, p. 5) found 
the home ranges of lion prides in the dune-savanna habitat of Kgalagadi 
Transfrontier Park to range from 1,762 to 4,532 km\2\. In India, 
however, Jhala et al. (2009, p. 3391) found the average home range of a 
breeding group of lionesses to be 33 km\2\. Similarly, Meena (2009, pp. 
7-8) found home ranges of females and males to be 35 km\2\ and 85 
km\2\, respectively.

Range

    The historical range of the lion included most current continental 
African countries (Chardonnet 2002, pp. 25-28) and extended from Greece 
through eastern Europe, southwest Asia (the Middle East), and India 
(Bauer et al. 2015a, unpaginated; Nowell and Jackson 1996, p. 38). 
Lions have undergone dramatic range retraction from this historical 
distribution (Ray et al. 2005, p. 67). Extirpation of lions in Europe 
occurred almost 2,000 years ago. The species was extirpated from 
southwest Asia within the last 150 years and northern Africa in the 
1940s (Bauer et al. 2015a, unpaginated; Black et al. 2013, p. 1; Nowell 
and Jackson 1996, p. 38). Today, lions occur only in Asia and sub-
Saharan Africa (Table 1). In Asia, P. l. leo only remains in the Gir 
Forests of India. Within sub-Saharan Africa, P. l. leo and P. l. 
melanochaita remain in 34 range countries (35 with South Sudan, which 
gained its independence as a country in July 2011) and have been 
recently extirpated from 12 African range countries and potentially 
extirpated from another 4 (Bauer et al. 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\,
                                     C[ocirc]te 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, Botswana, Burundi \2\,
                                     Djibouti \2\, Eritrea \2\,
                                     Ethiopia, Kenya, Lesotho \2\,
                                     Malawi, Mozambique, Namibia, Rwanda
                                     \3\, Somalia, South Africa, Sudan/
                                     South Sudan, Swaziland, Tanzania,
                                     Uganda, Zambia, Zimbabwe.
------------------------------------------------------------------------
\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 Africa (the total size of 
protected areas where lions were confirmed) is estimated at 49,000 
km\2\, or 1.1 percent of the historic range (Henschel et al. 2014, p. 
5). The most recent estimate of the lion's range throughout Africa 
comes from Bauer et al. (2015a, unpaginated) who estimate the extant 
lion range (areas reasonably confident that lions persist based on 
recent records) to be approximately 1.6 million km\2\ (617,763 mi\2\), 
or 8 percent of the historical range in Africa. The areas classified by 
Bauer et al. (2015, unpaginted) as possibly extinct total approximately 
1.8 million km\2\ (694,984 mi\2\), which is over half (52 percent) of 
the range classified as extant by the previous estimate conducted by 
Riggio et al. (2013, p. 26), which was based on estimates of savanna 
habitat. The lion's range in Asia is estimated to be approximately 
10,500 km\2\ (4,054 mi\2\), which occurs within the Gir National Park 
and Wildlife Sanctuary (Gir Protected Area), Girnar Wildlife Sanctuary, 
and surrounding agro-pastoral land (Bauer et al. 2015a, unpaginated; 
Banerjee and Jhala 2012, p. 1421; Jhala et al. 2009, pp. 3384, 3385; 
Nowell and Jackson 1996, p. 38).

Distribution and Abundance

    The general distribution of lions in Africa is summarized by Ray et 
al. (2005, p. 67) as follows:

    Currently, lions are restricted mainly to protected areas and 
surrounding conservancies or `game management areas,' with the 
largest populations in East and southern Africa. Where protection is 
poor, particularly outside protected areas, range

[[Page 80005]]

loss or population decreases can be significant. Declines have been 
most severe in West and Central Africa, with only small, isolated 
populations scattered chiefly through the Sahel. Lions in the region 
are declining in some protected areas and, with the exception of 
southern Chad and northern Central African Republic, are virtually 
absent from unprotected areas (Bauer 2003).

    Estimates of lion abundance on a large geographical scale are few 
in number. For a variety of reasons--including low densities, large 
ranges, cryptic coloration, nocturnal and wary habits--lions are 
difficult to count (Riggio et al. 2013, p. 31; Bauer et al. 2005, p. 
6). There are large areas of the species' range in which no data are 
available on lion occurrence or abundance (IUCN 2006b, pp. 12-13). 
Species experts recognize that estimating the size of the lion 
population in Africa is an ambitious task, involving many uncertainties 
(Bauer et al. 2012, unpaginated). Estimates, particularly throughout 
Africa or broad region-wide estimates tend to rely to a considerable 
extent on expert opinion or inference (Riggio et al. 2013, p. 21; 
Chardonnet 2002, p. 19). Consequently, there is a large degree of 
uncertainty in these estimates. In addition, to date all efforts to 
estimate the number of lions in Africa have used different methods; 
therefore, the results of earlier estimates cannot be directly compared 
to those of later estimates to determine population trend.
    The earliest estimates of lion abundance in Africa were educated 
guesses made during the latter half of the 20th Century. Bauer et al. 
(2008, unpaginated) summarize the information as follows:

    There have been few efforts in the past to estimate the number 
of lions in Africa. Myers (1975) wrote, ``Since 1950, their [lion] 
numbers may well have been cut in half, perhaps to as low as 200,000 
in all or even less.'' Later, Myers (1986) wrote, ``In light of 
evidence from all the main countries of its range, the lion has been 
undergoing decline in both range and numbers, often an accelerating 
decline, during the past two decades.'' In the early 1990s, IUCN SSC 
Cat Specialist Group members made educated ``guesstimates'' of 
30,000 to 100,000 for the African Lion population (Nowell and 
Jackson 1996).

    Ferreras and Cousins (1996, entire) provided the first 
quantitatively derived estimate of lion abundance in Africa using a 
GIS-based model calibrated with information obtained from lion experts. 
Ferreras and Cousins predicted lion abundance in Africa in 1980 to be 
75,800. Later, four additional efforts--Chardonnet (2002), Bauer and 
Van Der Merwe (2004), IUCN (2006a, 2006b), and Riggio et al. (2013)--
estimated lion population sizes ranging from 23,000 to 40,000 (Table 
2).
    Between 2006 and 2012, Henschel et al. (2014, p. 2) conducted field 
surveys in protected areas within designated Lion Conservation Units 
(LCUs) of western Africa to confirm lion presence where evidence of 
occurrence was lacking and to establish population estimates where 
lions occurred. Lions were absent from protected areas in 5 of the 10 
countries in western Africa where lions were considered to be present 
(Henschel et al. 2014, p. 4). Henschel et al. (2014, p. 5) estimated 
only 400 lions remain in the entire western region, with most (about 
350, or 88 percent) concentrated in a single population.
    Bauer et al. (2015a, unpaginated) attempted to correct for outdated 
sources in Riggio et al. (2013) by applying regional trends (discussed 
below) to 2002 population estimates for central, eastern, and southern 
Africa from Bauer and Van Der Merwe (2004) and Chardonnet (2002); 
estimates for western Africa were taken from Henschel et al. (2014) 
because of the greater precision of their estimate. Applying regional 
trends to Bauer and Van Der Merwe (2004) lion populations estimates, 
Bauer et al. (2015a, unpaginated; supporting information, Table 7) 
estimated lions in central Africa to be 590, eastern Africa to be 
7,345, and southern Africa to be 10,385 (Table 2). When regional trends 
were applied to Chardonnet (2002) lion estimates, Bauer et al. (2015, 
unpaginated; supporting information, Table 7) estimated lions in 
central Africa to be 1,748, eastern Africa to be 13,316, and in 
southern Africa to be 15,925 (Table 2). In total, Bauer et al. (2015, 
unpaginated) estimate the lion population in Africa to be between 
18,841 and 31,394. However, the authors found that the study by Bauer 
and Van Der Merwe (2004) was more conservative and stricter on data 
quality; therefore they have a greater confidence in an estimate closer 
to 20,000 lions in Africa. Additionally, the lion population in India 
was estimated to be 445 by Bauer et al. (2015a, unpaginated). In 2015, 
the Government of Gujarat completed its latest census, estimating 523 
lions in India (BBC 2015, unpaginated) (Table 2).

                                                          Table 2--Estimates of Lion Abundance
                                                          [Rows may not tally due to rounding]
--------------------------------------------------------------------------------------------------------------------------------------------------------
                                      Western Africa        Central Africa       Eastern Africa      Southern Africa
              Source                (percent of total)    (percent of total)   (percent of total)   (percent of total)     India            Total
--------------------------------------------------------------------------------------------------------------------------------------------------------
Ferreras & Cousins 1996 (estimate  ....................  ...................  ...................  ...................  ...........  75,800 (18,600 in
 for lion abundance in 1980).                                                                                                         protected areas).
Chardonnet 2002..................  1,163 (3 percent)...  2,815 (7 percent)..  15,744 (40 percent)  19,651 (50 percent)  ...........  39,373
Bauer & Van Der Merwe 2004.......  850 (4 percent).....  950 (4 percent)....  11,000 (48 percent)  10,000 (44 percent)  ...........  23,000
IUCN 2006 \1\ (as calculated by    1,640 (5 percent)...  2,410 (7 percent)..  17,290 (52 percent)  11,820 (37 percent)  ...........  33,160
 Riggio et al. 2013).
Riggio 2013 (based on estimates    480 (1 percent).....  2,419 (7 percent)..  19,972 (57 percent)  12,036 (34 percent)  ...........  34,907
 of savanna habitat).
Henschel et al. 2014.............  406 (n/a)...........  ...................  ...................  ...................  ...........  ...................
Bauer et al. 2015a (trends         ....................  590 (3 percent)....  7,345 (39 percent).  10,385 (55 percent)  ...........  18,726 *
 applied to Bauer and Van Der
 Merwe 2004).
Bauer et al. 2015a (trends         ....................  1,748 (6 percent)..  13,316 (42 percent)  15,925 (51 percent)  ...........  31,394 *
 applied to Chardonnet 2002).
Bauer et al. 2015a...............  ....................  ...................  ...................  ...................          445  ...................

[[Page 80006]]

 
Government of Gujarat 2015 **....  ....................  ...................  ...................  ...................          523  ...................
--------------------------------------------------------------------------------------------------------------------------------------------------------
\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 populations are limited to 
protected areas. These populations are largely isolated and many are 
small. P. l. leo (totaling approximately 1,500 lions), is divided into 
15 populations in and around protected areas; of these, 14 are 
remaining populations from a total of 38 historical occurrences in 
western and central Africa, while one occurs in India (Bauer et al. 
2015a, unpaginated; Henschel et al. 2015b, unpaginated; Brugi[eacute]re 
et al. 2015, p. 515; Henschel et al. 2014, pp. 4-5; Jhala et al. 2009, 
p. 3384). Nearly 90 percent of the lions in western Africa persist in a 
single population, the W-Arly-Pendjari (WAP) Complex (Henschel et al. 
2014, p. 5). Based on Bauer et al. (2015a, unpaginated; Supporting 
Information, Table 3) and Bauer and Van Der Merwe (2004, pp. 28-30), 
most P. l. melanochaita occur in approximately 68 protected areas 
throughout southern and eastern Africa, with larger populations 
occurring in Botswana, Kenya, Namibia, South Africa, Tanzania, Zambia, 
and Zimbabwe.

Population Trends

    Based on the best available information, lion range and numbers 
have clearly declined over the past several decades. However, not all 
lion populations have declined--some have increased or remained stable, 
and some have been restored to areas from which they were previously 
extirpated (Bauer et al. 2015a, unpaginated; Packer et al. 2013, p. 
636; Funston 2011, p. 3; Ferreira and Funston 2010, pp. 201, 203).
    Bauer et al. (2015a, unpaginated), using a time trend analysis of 
census data, determined the trend of lion populations from 1993 to 
2014. Overall, these lion populations decreased by 43 percent in 21 
years (Table 3). However, the authors found significant regional 
differences. In Asia, the single population increased by 55 percent 
(Bauer et al. 2015a, unpaginated). The population inside the protected 
area has stabilized and expanded into surrounding agro-pastoral land 
(Bauer et al. 2015b, p. 2; Breitenmoser et al. 2008, unpaginated). 
Additionally, the 2015 census of Gir Sanctuary and surrounding forest 
areas showed a 27 percent increase from the 2010 census (The Guardian 
2015, unpaginated). In southern Africa, the sample populations overall 
increased by 8 percent (Bauer et al. 2015a, unpaginated). However, one 
of the largest populations, Okavango, and populations of 6 unfenced 
reserves are declining (Bauer et al. 2015a, unpaginated, supporting 
information Table 3; Bauer et al. 2015b, p. 1). Fifteen of the 23 
sample populations in southern Africa were fenced; none experienced 
sharp declines and many small fenced populations are increasing (Bauer 
et al. 2015a, unpaginated, supporting information Table 3; Bauer et al. 
2015b, p. 1). South Africa was the only African country with growth in 
every population. However, these were all fenced populations, and most 
were reestablished in the past 20 years and quickly reached capacity 
(Bauer et al. 2015b, pp. 1-2). Populations in eastern Africa decreased 
overall by 59 percent (Bauer et al. 2015a, unpaginated). The Serengeti 
population was the only large population surveyed that did not 
decrease. Katavi National Park experienced complete loss of lions from 
an estimated 1,118 in 1993 to zero in 2014 (Bauer et al. 2015a, 
unpaginated, supporting information Table 3; Bauer et al. 2015b, p. 1). 
Western and central Africa (combined) experienced the largest decline 
at 66 percent (Table 3). All populations are declining, except the 
population in Pendjari; populations in Como[eacute] and Mole are now 
likely extinct (Bauer et al. 2015a, unpaginated, supporting information 
Table 3; Bauer et al. 2015b, p. 1). Furthermore, almost all lion 
populations in Africa that historically exceeded 500 individuals, the 
minimum number estimated to constitute a viable population (according 
to Riggio et al. 2013, p. 32 and Bj[ouml]rklund in Riggio et al. 2013, 
p. 32), are declining (Bauer et al. 2015b, p. 1).
    Although these trends are based on 47 sample populations, they 
comprise a substantial portion of the total remaining lion populations; 
therefore, the authors are confident in applying the observed trends to 
regions and the species as a whole (Bauer et al. 2015a, 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
                             Region                              -------------------------------- Percent 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%
----------------------------------------------------------------------------------------------------------------


[[Page 80007]]

    Using these rates of change, the authors calculated that the 
population in 5 countries (Botswana, India, Namibia, South Africa, and 
Zimbabwe), or 25 percent of the lion's range, increased by 12 percent, 
while the population in the remaining 75 percent of the range decreased 
by 60 percent (Bauer et al. 2015a, unpaginated), resulting in a 43 
percent population decrease of the entire lion species between 1993 and 
2014.
    The growth rate estimates discussed above are the best available 
information on global trends for lion populations, although Bauer et 
al. (2015b, p. 2) caution that these numbers are rough estimates. 
However, it is unlikely that regional declines are a product of 
differences in methodological shortcomings. Sample populations are all 
monitored with at least partial protection. Research sites are known to 
be generally avoided by poachers and encroachers. Therefore, the 
estimated growth rates may be less optimistic. It is likely that 
unmonitored, unfenced populations will have suffered greater rates of 
decline than reported since lack of management generally means a lack 
of conservation effort (Bauer et al. 2015b, p. 3).
    The work of Packer et al. (2013a, pp. 639-640) predicts future 
declines within a number of protected areas. Bauer et al. (2015b, p. 2) 
found that if regional trends remain unchanged in the future, lions in 
western and central Africa would likely lose a third of their 
population in 5 years and half of their population in 10 years. The 
population in eastern Africa is likely to decline by a third in 20 
years and half in 30 years. The Okavago population, Botswana, will also 
likely decline by a third in 20 years (Bauer et al. 2015b, p. 2). Many 
lion populations are expected to disappear within the next few decades 
such that the intensely managed populations in southern Africa will 
replace savanna landscapes as sites for the most successful 
conservation of lions.

Summary of Threats

    Today, lions are mainly restricted to protected areas; however, 
they still face serious threats that stem from inadequate management of 
those areas and increasing pressure on natural resources to meet the 
needs of a growing human population. Habitat loss has been extensive 
throughout the range of the lion, resulting in local and regional lion 
population extirpations and a dramatically reduced range with isolated 
lion populations that are increasingly limited to protected areas. As 
the human population increases, the protected areas where lions occur 
will be under increased pressure as more land is needed to satisfy the 
agricultural needs of the human population.
    Inadequate management and law enforcement has led to poaching of 
the lion's prey base in Africa for bushmeat, which has been critically 
depleted. Additionally, human population growth in Africa has led to 
human-lion conflict, particularly on the edge of protected areas, when 
pastoralists invade protected areas to allow their herds to graze or 
when lions move out of protected areas in search of prey, often preying 
on domestic livestock. Human-lion conflict leads to indiscriminate 
killing of lions, primarily as a result of retaliatory or preemptive 
actions to protect livestock and human lives. The close proximity of 
lions to humans and domestic livestock throughout their range exposes 
them to diseases, mainly transmitted through livestock and domestic 
dogs, which can impact general fitness, reproduction, and lifespan. 
These are in addition to diseases that naturally occur in lion 
populations in Africa. Furthermore, in some areas of Africa improper 
management has resulted in reduced lion numbers due to excessive lion 
harvests from trophy hunting. Subsequently, some lion populations are 
negatively impacted by infanticide following pride takeovers by new 
males.
    Because habitat loss has resulted in small, isolated populations 
across its range, lions face threats from stochastic events, such as a 
disease epidemic and inbreeding depression. An emerging threat to lions 
is trade in bones and other body parts for traditional medicine. These 
causes of lion population declines are widespread and likely to 
continue. The impacts of these threats are likely to be exacerbated by 
climate change. Projected changes indicate negative impacts to 
available habitat and, therefore, the range of the lion, prey 
availability, and the number of disease outbreaks as well as 
susceptibility to those diseases.

Habitat Loss

    Habitat destruction and degradation have been extensive throughout 
the range of the lion, resulting in local and regional lion population 
extirpations, reduced lion densities, a dramatically reduced range (see 
Range), and small, fragmented, and isolated lion populations that are 
increasingly limited to protected areas (see Distribution and 
Abundance) (Singh 2007, in Jhala et al. 2009, p. 3384; Ray et al. 2005, 
p. 69; Bauer and Van der Merwe 2004, pp. 29-30; Nowell and Jackson 
1996, pp. 20-21). In India, habitat loss is partly responsible for the 
decline of lions to a single population in a protected area. However, 
due to good protection and management, lions have dispersed to forested 
areas outside the protected area, extending their range from an initial 
1,883 km\2\ to 10,500 km\2\ (Johningh et al. 2007, Singh 2007, and 
Divyabhanusinh 2005, in Banerjee et al. 2010, p. 248; Singh 2007, in 
Jhala et al. 2009, p. 3384). Farming has been encouraged in the area 
and has flourished. Cultivated areas have created refuge areas and 
corridors for lion movement (Vijayan and Pati 2001 in Meena et al. 
2014, p. 124). At this time, no information indicates habitat loss is 
currently threatening the lion population in India. In Africa, however, 
despite lions being mainly found in protected areas, habitat loss and 
degradation continue to be among the main threats to lions (IUCN 2006a, 
p. 18; Ray et al. 2005, pp. 68-69).
    The main cause of lion habitat loss and degradation is expansion of 
human settlements and activities, particularly due to agriculture and 
intensive livestock grazing (IUCN 2006a, p. 18; IUCN 2006b, p. 23; Ray 
et al. 2005, pp. 68-69; Chardonnet 2002, pp. 103-106). From 1970 to 
2000, the human population in sub-Saharan Africa increased by 126 
percent (from 282 million to 639 million) (United Nations (UN) 2013, p. 
9), while at about the same time (1975 to 2000), agriculture area 
increased by 57 percent (from just over 200 million ha to almost 340 
million ha) and natural vegetation in the region decreased by 21 
percent (Brink and Eva 2009, p. 507). In 2009, approximately 1.2 
billion ha, or 40 percent, of Africa's land area was in permanent 
pasture or crops, with the vast majority (31 percent) in pasture (UNEP 
2012b, p. 68). Riggio et al. (2013, p. 29) estimate the original extent 
of savanna habitat in Africa to be approximately 13.5 million km\2\. 
Based on an analysis of land-use conversion and human population 
densities, Riggio et al. (2013, p. 29) found current savanna habitat 
that is suitable for lions to be fragmented and to total about 3.4 
million km\2\ (or 25 percent of African savanna habitat). This 
indicates a substantial decrease in lion habitat over the past 50 years 
and explains, in part, why lions are limited to protected areas.
    Based on a comparison of land-use and human population data, Riggio 
et al. (2013, p. 23) determined that a density of 25 or more people per 
km\2\ served as a proxy for the extent of land-use conversion that 
would render habitat unsuitable for lions. Woodroffe (2000, p. 167) 
analyzed the impact of people on predators by relating local

[[Page 80008]]

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

  Table 4--Human Population Projections in Countries Containing the 47 Sample Lion Populations Used by Bauer et
             al. (2015), Except C[ocirc]te D'ivoire and Ghana Where Lions Are Considered Extirpated
                                        [Population data is from UN 2013]
----------------------------------------------------------------------------------------------------------------
                                                        UN Population estimate, in thousands (people/km\2\)
          Subspecies                Country      ---------------------------------------------------------------
                                                       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

[[Page 80009]]

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

[[Page 80010]]

habitats along the Shetrunji River northeast of Gir (Meena 2014, p. 
27). Additionally, the size of the Gir Protected Area implies that 
dispersing lions will inevitably cross the protected area boundaries 
(Meena 2010, p. 212). When lions move, they must cross heavily 
populated human settlements and agricultural fields (Meena 2010, p. 
209). Traditional land uses are quickly changing in the region due to 
limestone mine and infrastructure development (Banerjee et al. 2010, p. 
250). Additionally, tourist activities (safaris to see the lions and 
religious pilgrimages to visit temples located within and on the border 
of protected areas) can have detrimental impacts to wildlife if not 
carefully planned. For example, construction of a road has been 
proposed to circle the outside of the whole Gir Protected Area System 
(Meena 2014, p. 28). Altering this habitat would result in land-use 
changes, promoting rapid development and urbanization and thereby 
disconnecting corridors for lion movement (Meena 2014, p. 28; Banerjee 
et al. 2010, p. 250). Furthermore, crossing these areas renders lions 
more vulnerable to disease transmission (See Disease below) and 
conflict with humans (see Human-Lion Conflict below). Because lions are 
social and territorial, they need adequate space to survive. Lack of 
adequate habitat will have a bearing on the lion's ecology, behavior, 
and population structure (Meena 2014, p. 28).
    Growing human populations have been associated with declines in 
large carnivore populations all over the world, and high human density 
is strongly associated with local extirpation of large carnivores 
(Linnell et al. 2001, Woodroffe 2001, in Woodroffe and Frank 2005, p. 
91; Woodroffe 2000, entire). Chardonnet et al. (2002, p.103) indicate 
that the distribution maps of lion subpopulations tend to confirm a 
direct inverse correlation of lion density and numbers with human 
activity and presence. Further, Packer et al. (2013a, entire) found 
that lions in unfenced reserves are highly sensitive to human 
population densities in surrounding communities.

Loss of Prey Base

    One of the most important requirements for carnivore survival, 
including lion, is prey availability, as it affects reproduction, 
recruitment, and foraging behavior and, therefore, also impacts lion 
movement, abundance, and population viability (Winterbach et al. 2012, 
p. 7, citing several sources). In India, prey abundance does not appear 
to be a concern for the lion population as conservation initiatives 
have ensured availability of ample prey (Banerjee et al. 2010, p. 249; 
Khan et al. 1996 and Singh and Kamboj 1996 in Meena 2010, p. 209; Jhala 
et al. 2009, p. 3384). The semi-nomadic pastoral communities that 
inhabit the Gir Forests are primarily vegetarian (Banerjee et al. 2013, 
p. 2); therefore, there is no great demand for bushmeat. However, in 
most African countries, large carnivores such as lions are under 
serious threat through decreased prey abundance (Bauer et al. 2014, p. 
97) due to unsustainable and increasingly commercialized bushmeat 
hunting in and around protected areas (Bauer et al. 2015a, unpaginated; 
Henschel et al. 2015, unpaginated; Henschel et al. 2014, p. 5; Lindsey 
et al. 2013b, p. 84; Lindsey and Bento 2012, pp. 1-2, 61; Scholte 2011, 
p. 7; Bouch[eacute] et al. 2010, pp. 1000, 1001; Cragie et al. 2010, p. 
2227; Brashares et al. 2004, p. 1181; Fischer and Linsenmair 2001, pp. 
132, 133).
    Humans in Africa rely on protein obtained from bushmeat, resulting 
in direct competition for prey between humans and lions, and commercial 
poaching of wildlife is becoming a significant threat to many species, 
including those that lions rely upon for food. Subsistence hunting was 
traditionally carried out with the use of spears, which had minimal 
impact to wildlife populations. Spears have since been replaced by 
automatic weaponry (Chardonnet et al. 2010, p. 27) and snares, which 
are most commonly used (Lindsey et al. 2013b, p. 83). These methods 
allow for poaching of large numbers of animals for the bushmeat trade, 
particularly snares, which are cheap, difficult to detect, and 
unselective as they can kill nontarget animals ranging from rodents to 
elephants (Lindsey et al. 2013b, p. 83).
    The human population in a majority of African countries within the 
range of the lion has quadrupled since the 1960s (Riggio et al. 2013, 
p. 29; IUCN 2009, p. 15), increasing the demand for bushmeat. Bushmeat 
contributes significantly to food security, and is often the most 
important source of protein in rural areas (Nasi et al. 2008 in Lindsey 
et al. 2013b, p. 82). It comprises between 6 percent (southern Africa) 
and 55 percent (CAR) of a human's diet within the lion's range in 
Africa (Chardonnet et al. 2005, p. 9; IUCN 2006b, p. 19). In western 
Africa, bushmeat is a secondary source of protein, with fish being the 
primary source. However, when widespread loss of jobs and income occurs 
due to poor fish harvests, bushmeat becomes an important source of 
income and sustenance, leading to increased presence of hunters in 
protected areas and higher than average declines in wildlife (Brashares 
et al. 2004, pp. 1180-1181).
    The sale of bushmeat is an important livelihood in Africa 
(Chardonnet et al. 2010, p. 27; Mesochina et al. 2010a, p. 38; Abwe and 
Morgan 2008, p. 26; Bennett et al. 2007, p. 885; Fa et al. 2006, p. 
507). The little meat produced from domestic livestock is unaffordable 
for common people (Bouch[eacute] et al. 2010, p. 1001). Bushmeat 
hunting is rarely practiced solely for subsistence. It supplies meat 
for local consumption and trade, urban markets, and even international 
markets (Lindsey et al. 2013b, pp. 86-87). Outlets for the sale of 
bushmeat have arisen in some areas, and full-time commercial bushmeat 
traders occur in most southern and eastern African countries (Lindsey 
et al. 2013b, p. 86). Significant distribution of bushmeat to Europe 
and the United States, where it is sold at elevated prices, drives 
increasing commercialization of trade, a greater number of hunters, 
adoption of more efficient hunting methods, and an unprecedented 
pressure on wildlife populations (Stiles 2011 and Barnett 2000 in 
Lindsey et al. 2013b, p. 88). Many illegal hunters are poor (Barnett 
2000 in Lindsey et al. 2013b, p. 88; Lindsey and Bento 2012, p. 37; 
Scholte 2011, p. 7). Bushmeat trade can provide a quick income to 
purchase other food and essentials (Lindsey et al. 2013b, p. 82; 
Lindsey and Bento 2012, p. 62). Hunters are wealthier than non-hunters 
(Knapp 2007 in Lindsey et al. 2013b, p. 86) and enjoy elevated social 
status.
    This growing demand and the availability of modern weapons have led 
to many African wildlife species being hunted at unsustainable levels 
and the lion prey base becoming depleted in many areas (Hoppe-Dominik 
et al. 2011, p. 452; Chardonnet et al. 2010, pp. 6, 13-14, 27; Packer 
et al. 2010, p. 8; Frank et al. 2006, p. 12). Because wildlife has been 
depleted in non-protected areas, illegal bushmeat hunters are 
increasingly focusing efforts on protected areas (Lindsey et al. 2013b, 
p. 84). Weak management effectiveness and inadequate law enforcement 
have facilitated poaching for bushmeat in protected areas and resulted 
in a widespread decrease in large mammal populations, including lion 
prey, in these areas (Henschel et al. 2015b, unpaginated; Henschel et 
al. 2014, pp. 5, 7; Lindsey et al. 2013b, pp. 84, 88; Lindsey and Bento 
2012, p. 61; Scholte 2011, p. 7; Bouch[eacute] et al. 2010, pp. 99, 
1001; Brashares et al. 2004 in Craigie et

[[Page 80011]]

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

Human-Lion Conflict

    The lion population in and around the Gir Protected Area, India, 
lives among and is surrounded by many pastoral and forest settlements 
(Banerjee and Jhala 2012, p. 1421; Singh and Gibson 2011 in Banerjee 
and Jhala 2012, p. 1421; Banerjee et al. 2010, p. 249; Singh 2007 in 
Jhala et al. 2009, p. 3385). The lion population of Gir has increased 
and dispersed into the large agro-pastoral area adjacent to the 
protected area. Only 10 percent of lions in India occur in the human-
free portion of Gir National Park (Banerjee et al. 2013, p. 8). 
Conflict there, like in Africa, arises from predation of livestock and 
associated threats to security of pastoral livelihoods (Karanth and 
Chellam in Banerjee et al. 2013, p. 1). The lion's diet there includes 
livestock (Banerjee et al. 2013, p. 6; Meena et al. 2011, pp. 63-65). 
Between 2001 and 2010 the number of villages reporting depredation of 
livestock increased (Meena et al. 2014, pp. 122-123). Additionally, 
Meena (2012, p. 36) found that in all Forest Divisions, except Gir 
West, annual livestock predation increased more than 100 percent in 5 
years. However, despite the lion's close occupation with human 
settlements and increased predation on livestock, human-lion conflict 
and associated retaliatory killing was not found to be a major source 
of lion mortality (Pathak et al. 2002 in Banerjee and Jhala 2012, p. 
1427), mainly due to low economic losses via certain husbandry 
practices and a compensation scheme (Meena et al. 2014, pp. 123, 124; 
Banerjee et al. 2013, pp. 6-7, 8), cultural ethics (Raval 1991 in 
Banerjee et al. 2013, p. 2; Banerjee et al. 2013, p. 8), and strict 
legal enforcement (Banerjee et al. 2013, p. 8).

[[Page 80012]]

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

[[Page 80013]]

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

[[Page 80014]]

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

[[Page 80015]]

(Hazzah et al. 2014, p. 852). This situation further contributes to 
negative attitudes toward lion conservation programs (Moghari 2009, p. 
37).
    Lions are particularly vulnerable to retributive killing because 
they are often driven by a perceived level of lion predation on 
livestock rather than actual levels of conflict. In some locations, 
other predators (e.g., baboons (Papio ursinus), spotted hyenas (Crocuta 
crocuta), and leopards (Panthera pardus)) as well as disease are 
responsible for the majority of livestock losses and human casualties, 
yet it is lions that are sought and killed more often. In the Pendjari 
Biosphere Reserve, Sogbohossou et al. (2011, p. 74) found that just one 
case of a nonlethal attack on a human in a decade and mere rumors of 
attacks in other regions was enough to cause people to perceive lions 
as a threat. Negative perceptions of lions may be based on an over-
estimated number of lions in a community or protected area and an over-
estimated number of human-lion conflicts (Dickman 2013, p. 380; Begg 
and Begg 2010, p. 20; Chardonnet et al. 2010, pp. 12, 21-22; Hazzah et 
al. 2009, p. 2436; Maclennan et al. 2009 in Hazzah et al. 2009, p. 
2429; Moghari 2009, pp. 77-78, 107, 150; Holmern et al. 2007 in Moghari 
2009, p. 34; Butler 2001 in Moghari 2009, p. 34; Kissui 2008, pp. 426, 
428, 429; Hazzah 2006, pp. 18-19, 83-85, 96, 98, 107, 111; Patterson et 
al. 2004, pp. 514, 515). One cause for the disproportionate blame put 
on lions is that the lion is a highly visible species. It is a large-
bodied species that lives in groups and has cultural significance. 
Because of its physical presence, there is often a hyper-awareness of 
the potential risk for lion attacks and lions may be blamed simply 
because they have been seen in an area (Dickman 2013, pp. 380-381).
    Cultural beliefs and traditions can have a negative impact on 
lions. Because cattle are of great cultural significance to Maasai, 
their loss can impose social or cultural costs and incite greater 
resentment and higher levels of retributive killing (Dickman 2013, p. 
384; Kissui 2008, p. 429; Hazzah 2006, p. 99). Cultural beliefs still 
motivate ritual lion hunts for young Maasai warriors. Despite being 
outlawed, this practice persists due to community secrecy. However, it 
is easily disguised as retaliatory killings for livestock predation. 
The prohibition of ritual lion hunts provides a greater incentive for 
participating in retaliatory hunts (Hazzah et al. 2014, p. 852; Packer 
et al. 2010, p. 10; Moghari 2009, pp. 13-14, 28; Ikanda 2008, pp. 5, 6; 
Kissui 2008, p. 423; Frank et al. 2006, p. 10; Hazzah 2006, p. 99). In 
some areas of Africa, locals believe in ``spirit lions,'' a lion whose 
body is overtaken by evil to kill rivals or their livestock (West 2001 
in Dickman 2013, pp. 381-382). Because people believe spirit lions are 
created by their enemies, the number of perceived spirit lions, and 
killing of these lions, increases during times of social tension 
(Dickman 2013, p. 382.)
    Cultural beliefs can also have a positive impact on lions. An 
association with a totem is an important component of certain cultures 
and could explain why retaliatory killing is uncommon in some areas 
despite negative perceptions. However, the positive impact may not 
continue as cultural beliefs dwindle due to urbanization and 
modernization (Sogbohossou et al. 2011, pp. 73, 75).
    Social tensions within tribes and between local communities and 
other communities, the government, park officials, or tourists can lead 
to conflict and retributive killing of lions (Dickman 2013, p. 382; 
Hazzah 2006, p. 75). Locals often report that wildlife authorities do 
not react effectively when chronic livestock raiders are reported 
(Frank et al. 2006, p. 9). Significant numbers of lions have been 
killed when promised benefits were not received or adequate 
compensation was not provided for livestock and human losses (Dickman 
2013, p. 383; Hazzah 2006, p. 45).

Trophy Hunting

    Lions are a key species in sport hunting, or trophy hunting, as 
they are considered one of the ``big five'' African species (lion, 
leopard, elephant, rhino, and cape buffalo) touted to be the most 
challenging to hunt due to their nimbleness, speed, and behavioral 
unpredictability (Lindsey et al. 2012a, p. 2). However, with the 
documented decline in lion population numbers throughout Africa, sport 
hunting of lions for trophies has become a highly complex issue.
    Trophy hunting is carried out in a number of range countries and is 
considered an important management tool for conserving land and 
providing financial resources for lion conservation. However, 
management programs are not always sufficient to deter unsustainable 
off takes (harvests), which has occurred in many areas (Lindsey et al. 
2013a, pp. 8-9; Packer et al. 2006 in Bauer et al. 2015a, unpaginated). 
Documented declines in lion populations of Africa are a result, in 
part, of mismanaged trophy hunting (Rosenblatt et al. 2014, entire; 
Sogbohossou et al. 2014, entire; Becker et al. 2013, entire; Lindsey et 
al. 2013a, entire; Packer et al. 2013, p. 636; Croes et al. 2011, 
entire; Packer et al. 2011, entire; Loveridge et al. 2007, entire). 
Depending on how trophy hunting is regulated and managed, trophy 
hunting can be a tool for conservation, but may also have negative 
impacts on lions (Bauer et al. 2015a, unpaginated; Lindsey et al. 
2013a, p. 1; Whitman et al. 2004, pp. 176-177; Loveridge et al. 2007, 
p. 548).
    In response to growing international recognition of reduced 
population numbers, many countries began implementing moratoriums 
banning the sport hunting of lions. In this document we use the terms 
moratorium and ban interchangeably. A ban or moratorium can be 
permanent, long term, or temporary, and can occur in countries that 
have hunting quotas in place (e.g., Botswana and Zambia). Having both a 
moratorium and a quota in place at the same time means that, although 
the country may have a hunting quota, the country has halted 
authorization of trophy hunting pursuant to that quota until some later 
date or until some further action is taken, as prescribed by that 
country.
    Trophy hunting is currently banned in 12 range countries: Angola, 
Botswana, Congo, Gabon, Ghana, India, Kenya, Malawi, Mauritania, Niger, 
Nigeria, and Rwanda (CITES 2014, p. 14; Meena 2014, p. 26; Lindsey et 
al. 2013a, entire; Lindsey 2013, pers. comm.; Jackson 2013, pp. 7-8). 
In 1977, Kenya banned all sport hunting (Elliot and Mwangi 1998, p. 3). 
Botswana banned lion hunting between 2001 and 2004, and then again from 
2008 to the present (Davison et al. 2011, p. 114). Benin imposed a 2-
year moratorium, and CAR a 3-year moratorium, in the early 2000s 
(Lindsey et al. 2013a, p. 4). In January of 2013, Zambia placed a 
moratorium on sport hunting in 19 game management areas. While a few 
other game management areas and private game ranches in Zambia remain 
open for sport hunting for other species, the nationwide moratorium on 
sport hunting of cats remains in place (White 2015, pers. comm.; ABC 
News 2014, unpaginated; Flocken 2013, unpaginated). Trophy hunting is 
restricted to problem or dangerous animals in Ethiopia and Uganda 
(Lindsey 2008, p. 42). In our proposed rule, we had conflicting 
information regarding whether Cameroon had or has a lion hunting 
moratorium (CITES 2014, p. 14; Lindsey 2013, pers. comm.; Jackson 2013, 
p. 8). During the public comment period, a peer reviewer confirmed that 
Cameroon has not put a moratorium in place for lions, either in the 
past or present (Bauer 2015, pers. comm.). Additionally, Zimbabwe has

[[Page 80016]]

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

                 Table 5--Annual Trophy Quotas and Offtake by Country (Approximate) as of 2013*
----------------------------------------------------------------------------------------------------------------
                                                    Annual lion     Year(s) of        Annual        Year(s) of
                     Country                       trophy quotas       data          offtakes          data
----------------------------------------------------------------------------------------------------------------
Panthera leo leo
Benin...........................................       5.00                       minus>0.4
Burkina Faso....................................      20.00                      minus>1.45
Cameroon........................................      29.22                       minus>1.0
CAR.............................................              31            2009      13.76.9
Panthera leo melanochaita                         ..............  ..............  ..............  ..............
Mozambique......................................           42-60            2013      19.27.3
Namibia.........................................            14.5            2010      14.03.2
Tanzania........................................             315       2011-2012              85       2011-2012
Zambia (moratorium) \1\.........................      74(50 \2\)            2012              47            2012

[[Page 80017]]

 
Zimbabwe........................................      101(50\3\)            2011      42.57.5
----------------------------------------------------------------------------------------------------------------
* 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
    Proponents and most lion experts support trophy hunting as a 
conservation tool for the lion if it is practiced in a sustainable and 
scientifically based manner (Henschel 2015, pers. comm.; Hunter 2011, 
entire; van der Merwe 2013, entire; Hunter et al. 2013, entire) because 
it can provide: (1) Incentives for the conservation of large tracts of 
prime habitat, and (2) funding for park and reserve management, anti-
poaching activities, and security activities.
    As habitat loss has been identified as one of the primary threats 
to lion populations, it is notable that the total amount of land set 
aside for hunting throughout Africa, although not ameliorating threats 
to habitat loss, exceeds the total area of the national parks, 
accounting for approximately half of the amount of viable habitat 
currently available to lions (Chardonnet et al. 2010, p. 34; Packer et 
al. 2006, pp. 9-10). For example, in Tanzania, 25-33 percent of the 
total area, covering over 247,000 km\2\ and encompassing 190 hunting 
units, has been set aside for sport hunting purposes; this has resulted 
in an area 5.1 times greater than Tanzania's fully protected and 
gazetted parks (Jackson 2013, p. 6; Barnett and Patterson 2005, p. 61). 
Tanzania also has land set aside for sport hunting in the form of 
safari areas, communal land, and privately owned properties that make 
up 23.9 percent of the total land base (Barnett and Patterson 2005, pp. 
76-77).
    In Botswana, despite the current ban on lion hunting, the country 
currently has over 128,000 km\2\ of gazetted wildlife management areas 
and controlled hunting areas set aside for hunting purposes, which 
equates to 22.1 percent of the country's total area. This amount is in 
addition to 111,000 km\2\ (or 19.1 percent of the country's total area) 
set aside as habitat in the form of national parks, game reserves, and 
forest reserves (Barnett and Patterson 2005, p. 7). In 2000, five 
countries in southern Africa (Botswana, Namibia, South Africa, 
Tanzania, and Zimbabwe) set aside a combined 420,000 km\2\ of communal 
land, 188,000 km\2\ of commercial land, and 420,089 km\2\ of state land 
totaling over 1,028,000 km\2\ for sport hunting purposes (Barnett and 
Patterson 2005, p. iii).
    As a species with a considerable range (up to 1,000 km\2\) (Packer 
et al. 2013, p. 636; Haas et al. 2005, p. 4), suitable habitat is 
important to the survival of the species, and the marked decline in 
suitable habitat is a significant threat to the species (see Habitat 
Loss). The land currently designated in Africa for use in sport hunting 
has helped to reduce, but not eliminate, the impact of habitat loss on 
the lion.
    If trophy hunting is part of a scientifically based management 
program, it may provide direct economic benefits to the local 
communities and may potentially create incentives for local communities 
to conserve lions, reduce the pressure on lion habitat, and reduce 
retaliatory killing, primarily because lions are viewed as having 
value. Conversely, lack of incentives could cause declines in lion 
populations because lions are viewed as lacking value and they kill 
livestock, which are of great value to communities (see Human-lion 
Conflict).
    Over the last few decades, conservationists and range countries 
have realized the integral role local communities play in the 
conservation of lions and their habitat; when communities benefit from 
a species, they have incentive to protect it. Therefore, using wildlife 
as a source of income for rural populations has increasingly been 
employed throughout the lion's range countries in Africa. Many of these 
countries are classified as ``developing'' nations; specifically, seven 
of the ten countries (we include Cameroon here) where trophy hunting is 
permitted have 27-64 percent of their human populations living in 
severe poverty (United Nations Development Programme's (UNDP) 2014, 
unpaginated; Barnett and Patterson 2005, p. iii). These countries often 
have high population growth, high unemployment, limited industry, and a 
Gross Domestic Product (GDP) per capita lower than the poverty level 
(Barnett and Patterson 2005, p. iii). These combined challenges 
highlight the need for innovative solutions. Conservationists and range 
countries recognize the value of the wildlife sector; if managed 
sustainably, there is potential to contribute to rural economic 
development while simultaneously protecting the unique ecological 
habitats and species contained therein (Chardonnet et al. 2010, p. 33; 
Kiss 1990, pp. 1, 5-15).
    For species such as the lion to persist, local communities must 
benefit from or receive a percentage of funds generated from tourism 
such as wildlife viewing, photography, or trophy hunting (White 2013, 
p. 21; Martin 2012, p. 57; Kiss [editor] 1990, pp. 1, 5-15). The 
economic value of a species, such as lion, can encourage range 
countries to develop management and conservation programs that involve 
local communities and which would ultimately discourage indiscriminate 
killings by local communities (Groom 2013, pp. 3, 5; Hazzah et al. 
2013, p. 1; White 2013, p. 21; Martin 2012, p. 49). If local 
communities see no benefit of lions being present in their communal 
areas, sustainable use of lions becomes less competitive with other 
land-use options, such as grazing and livestock management, and local 
communities become unwilling and unable to manage their wildlife 
heritage (Barnett and Patterson 2005, p. iii). When the value of lions 
in areas outside national parks is diminished, those areas are likely 
to be converted to forms of land use less suitable for lions, such as 
agriculture, livestock pastures, or areas of resource extraction, 
making lions even more vulnerable to expanding human settlement (Van 
der Merwe 2013, p. 2).
    Community conservancies that benefit from trophy hunting have 
specifically been formed as a way to protect wildlife and habitat. As 
an example, in Namibia, 160,000 km\2\ of community conservancies were 
established in part due to revenue from trophy hunting. These 
conservancies benefit the local communities, which in turn protect lion 
habitat. In 2012, the Sav[eacute] Valley

[[Page 80018]]

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

 Arbitrary establishment of quotas and excessive harvest
 lack of age-restriction implementation
 fixed quotas
 hunting of females
 lack of minimum hunt lengths in some countries
 general problems associated with management of trophy hunting

    As discussed above, one of the primary practices experts identify 
as undermining sustainable trophy hunting is the use of non-scientific 
information underlying the development of quotas (Lindsey et al. 2013a, 
p. 8). The best available monitoring data should be used to set quotas 
if they are to be scientifically based and sustainable. However, 
monitoring data are often lacking (Barnett and Patterson 2005, p. 102). 
A limited number of independent, scientific population counts of lions

[[Page 80019]]

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

[[Page 80020]]

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

[[Page 80021]]

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

[[Page 80022]]

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

[[Page 80023]]

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

[[Page 80024]]

l. leo, due to a lack of lion populations in some of these countries, 
it is likely that fewer countries could conduct lion trophy hunts. A 
study found that Namibia, Tanzania, Zambia, and Zimbabwe all had quotas 
higher than the recommended level for sustainability; however, Zimbabwe 
has reduced their quota. Mozambique (Niassa National Reserve) is the 
only location found to have a quota below the recommended level. Age-
based strategies have been implemented and shown to reduce offtakes in 
Mozambique (only in Niassa National Reserve, excludes the rest of the 
country), Tanzania, and Zimbabwe. Furthermore, Zimbabwe and Niassa 
National Reserve are the only two locations that have fully implemented 
an age-based strategy with transparency, an element experts say is 
critical to a quota allocation system. Several other countries have 
made commitments to implement the age-restrictions strategy but their 
progress is pending. In South Africa, 99 percent of the lion trophies 
are captive bred, and, therefore, were not the result of removing lions 
from the wild.
    Unless reforms are made to the current management of trophy 
hunting, we expect the declines specifically documented from excessive 
offtakes in Benin, Cameroon, Tanzania, Zambia, and Zimbabwe to 
continue. Furthermore, we expect excessive harvests to further 
contribute to declines in the species across its African range.

Import/Export of Lion Trophies

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

[[Page 80025]]

trophies from seven African countries, as follows:

Botswana = 1 trophy (originated from Mozambique)
Burkino Faso = 3 trophies
Mozambique = 5 trophies
Namibia = 9 trophies
South Africa = 545 trophies (the majority of which are reported to be 
of captive-born origin; additionally 2 captive trophies originated in 
South Africa, imported to Canada, and then imported into the United 
States)
Tanzania = 3 trophies
Zambia = 17 trophies
Zimbabwe = 44 trophies

    Based on CITES trade data, lion trophy exports have decreased 
throughout most of the lion's range, including Tanzania, which has one 
of the largest lion populations. South Africa is the only country where 
exports have increased because most of these trophies are of captive 
origin.

Traditional Use of Lion Parts and Products

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

[[Page 80026]]

a market in Mong La, Myanmar (Oswell, 2010, p. 12). The town, known for 
incidents of wildlife trafficking, is less than 2km from the Chinese 
border. Up to 2006/2007, Williams et al. (2015, p. x, Table 11, Figure 
24) noted:

``The combined quantity of live lions and lion parts and derivatives 
exported to East-Southeast Asia from South Africa was minimal in the 
broader global trade. From 2008, however, the quantities exported 
increased almost six-fold from the previous year. Not only did the 
number of live lions exported to East-Southeast Asia reach record 
levels from this time, but also the first permits to export lion 
skeletons were issued. The demand for lion parts and derivatives 
appears to have coincided with the strengthened conservation 
measures adopted in 2006-2007 to protect tigers and Asian big cats. 
Accordingly, tiger parts were increasingly substituted with lion 
parts obtained from Africa. The trade in lion parts and derivatives 
to Lao PDR dominates the exports. Since 1998, but especially after 
2007, China, Viet Nam, Lao PDR, Myanmar and Thailand have imported 
increasing amounts of live lions, lion bodies and bones from South 
Africa.''

    Evidence also indicates ``well established'' links between South 
Africa's legal lion bone trade and the Xaysavang Network, an 
international wildlife trafficking syndicate that is also involved in 
the illicit rhino horn trade in South Africa (Williams et al. 2015, pp. 
7-10, 59; Environmental Investigative Agency 2014, p. 13; U.S. 
Department of State 2013, unpaginated). The U.S. Department of State 
has issued a $1 million reward for information leading to the 
dismantling of this network. According to the U.S. Department of State, 
the Xaysavang Network facilitates the killing of endangered species in 
Africa and elsewhere and smuggles them to Laos for export to other 
Asian countries (U.S. Department of State 2013, unpaginated). During 
2008-2011, the vast majority (85%) of the permits issued by South 
Africa to export lion skeletons or carcasses were issued for exports to 
Laos (Williams et al. 2015, pp. x, 46) and, for the only 2 years for 
which data were available (2009 and 2010), over half of the 
consignments destined for Laos were listed as imported by Vixay 
Keosavang, believed by the U.S. Department of State to be the leader of 
the Xaysavang network (U.S. Department of State 2013, unpaginated; 
Williams et al. 2015, pp.8-10). The involvement of the Xaysavang 
Network in South Africa's lion bone trade indicates there are well-
established avenues for laundering of illegally obtained lion bones, 
such as those obtained from poached wild lions, into the legal trade.
    Lastly, evidence suggests incentive to poach wild lions for the 
bone trade may currently exist. According to Williams et al. (2015, p. 
x), the 2013 price paid to South African game farmers and landowners 
for lion bones was $1,260-2,100 USD per skeleton. In many lion range 
states this exceeds per capita GDP (gross domestic product) (World Bank 
2015, unpaginated). Thus, the current price paid for lion bone appears 
to provide incentive in some countries to poach wild lions.
    While the lion bone trade appears to currently be based primarily 
in South Africa's captive-bred lion hunting industry, the trade appears 
to be having little or no impact on wild lion populations in South 
Africa at this time--lion populations in South Africa are stable or 
increasing and there is little poaching of wild lions in the country 
(Funston and Levendal 2014, pp. 1, 26; Williams et al. 2015, pp. 79-
80). However, the impact of the lion bone trade on lion populations 
outside South Africa is unknown, and most wild lions occur outside 
South Africa (see Distribution and Abundance). Based on the effect of 
the tiger bone trade on tiger populations, if current conditions--for 
example, rapidly increasing demand and involvement of an international 
crime syndicate--continue unchanged, then there is considerable 
potential for extensive poaching of wild lions to occur in order to 
meet demand.

Disease

    Wild lions are known to be infected with various pathogens (Hunter 
et al. 2012, p. 2; Craft 2008, p. 6; Michel et al. 2006, p. 92; 
Hofmann-Lehmann et al. 1996, pp. 559-561). However, information on the 
extent of infections and impacts of diseases on lion populations is 
limited. We found one study documenting disease in a single wild lion 
in India that died from trypanosomiasis in 2007; analysis of tissue 
samples also detected peste des petits ruminants virus (PPRV), which is 
not known to cause disease in carnivores (LionAid 2013, unpaginated; 
Balamurugan et al. 2012, pp. 203, 205). Information on the presence of 
disease and impacts to lions come from a few long-term studies that 
have been conducted in Africa, including Serengeti National Park, 
Ngorongoro Crater, and Kruger National Park.
    As a result of human population expansion into lion habitat, lions 
are increasingly exposed to diseases from domestic animals (IUCN 2006b, 
p. 26). Because lions are a top predator, they are at a particularly 
high risk of exposure to pathogens (Keet et al. 2009, p. 11). Some 
pathogens are endemic, meaning they are constantly present, but often 
do not cause disease. Others are epidemic and cause a sudden severe 
outbreak with the potential to cause high mortality (Craft 2008, pp. 5, 
6). The association between disease, age, nutritional health and other 
factors that could predispose a lion to morbidity and, eventually, 
mortality is complex. It is often difficult to determine whether 
mortality was due to a single factor or a combination. Lions could be 
infected with and become debilitated by a disease, but the actual cause 
of death could be other factors, such as fighting with other lions or 
large predators (LionAid 2014a, p. 4).
    Feline calicivirus, feline herpesvirus, feline parvovirus, feline 
coronavirus, and feline leukemia virus are endemic viruses known to 
occur in lions of Serengeti National Park, Ngorongoro Crater, Lake 
Manyara National Park, Kruger National Park, and Etosha National Park 
(but not all viruses are known in all parks). However, these diseases 
are not known to affect lion survival (Hunter et al. 2012, p. 2; Craft 
2008, p. 6; Hofmann-Lehmann 1996, pp. 559, 561).
    Lions within Kruger National Park and Hluhluwe-iMfolozi Park, South 
Africa, and Serengeti National Park, Tanzania, are known to be infected 
with Mycobacterium bovis, a pathogen that causes bovine tuberculosis 
(bTB). This pathogen is not endemic to African wildlife and was likely 
introduced from cattle imported from Europe. M. bovis is transmitted to 
ungulates, such as African buffalo (Syncerus caffer) and wildebeest 
(Connochaetes taurinus), from domestic cattle located on the periphery 
of the parks (Maas et al. 2012, p. 4206; Keet et al. 2009, pp. 4, 11; 
Renwick et al. 2007, p. 532; Michel et al. 2006, pp. 92, 93; Cleaveland 
et al. 2005, pp. 446, 449, 450). Spillover of the disease from buffalo 
to other lion prey species, such as kudu (Tragelaphus strepsiceros) and 
warthog (Phacochoerus africanus), has also been documented (Keet et al. 
2009, pp. 4, 11; Renwick et al. 2007, p. 535; Cleaveland et al. 2005, 
p. 450). Because the lion's primary prey are infected with bTB, they 
are frequently exposed to large amounts of infected tissue and are at 
risk of infection (Keet et al. 2009, pp. 4, 6; Renwick et al. 2007, pp. 
532, 536; Michel et al. 2006, p. 93; Cleaveland et al. 2005, pp. 450, 
451). Furthermore, predators prey on weak animals and scavenge on 
carcasses, increasing their likelihood of being exposed to M. bovis 
(Renwick et al. 2007, p. 536; Michel et al. 2006, p. 93). Transmission 
may also occur among lions via scratching and biting (Keet et al. 2009, 
p. 7; Renwick

[[Page 80027]]

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

[[Page 80028]]

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

[[Page 80029]]

Reserve and Ngorongoro Crater lions following drought events.

Deleterious Effects Due to Small Population Sizes

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

[[Page 80030]]

2015, unpaginated; Duerr 2014, unpaginated; Meena 2014, p. 29).

Regulatory Mechanisms

    Regulatory mechanisms in place to provide protections to African 
lions vary substantially throughout Africa. The lion species (Panthera 
leo) is listed in Appendix II of CITES; however, the former Asiatic 
lion (P. l. persica) is listed in Appendix I. With the exception of 
South Sudan, all of the lion range states are Parties to CITES. 
According to the draft CITES Periodic Review of the Status of African 
Lions (CITES 2014, pp. 14-15) outside of CITES, lions have no legal 
protections in four countries: Burundi, Guinea Bissau, Lesotho, and 
Swaziland. However, CITES 2014 (p. 15) states that most of the southern 
and eastern lion range states have regulatory mechanisms in place to 
protect lions. We found that most of the range states have national 
environmental legislation to establish national parks and conservation 
areas, and to conserve and regulate the take, hunting, and trade of 
wildlife, including parts and products, but could find no legislation 
specific to lions, or to the main threats affecting lions: habitat 
loss, human-lion conflict, and loss of prey base (Ecolex \1\ 
information last accessed November 6, 2015).
---------------------------------------------------------------------------

    \1\ ECOLEX is a comprehensive database on environmental law, 
maintained by the International Union for Conservation of Nature 
(IUCN), the United Nations Environment Programme (UNEP), and the 
Food and Agriculture Organization of the United Nations (FAO). Our 
search terms used with respect to wildlife laws were ``African 
lion,'' ``Asiatic lion,'' ``Panthera leo leo,'' ``Panthera leo 
persica,'' and ``country,'' e.g., ``Angola,'' ``Benin,'' etc. 
Information accessed at http://ecolex.org.
---------------------------------------------------------------------------

    National and international conservation strategies rely on 
protected areas to protect natural resources from negative impacts of 
human populations (Craigie et al. 2010, p. 2221). The lion is largely 
limited to protected areas; therefore, effective management is crucial 
to the survival of the species. However, weak management of protected 
areas has been documented across its range, especially in western 
Africa where most protected areas are experiencing severe management 
deficiencies (Henschel et al. 2015, unpaginated; Henschel et al. 2014, 
pp. 5, 7; Brugi[eacute]re 2012 in Henschel et al. 2014, p. 7; Craigie 
et al. 2010, entire). The WAP complex in western Africa had received 
high scores for management effectiveness (Henschel et al. 2015, p. 7).
    Effective management requires adequate funding, resources, and 
staff. Packer et al. (2013a, pp. 638-639) found that lion densities 
were highest in protected areas with the highest management budgets. 
Cost estimates for maintaining lion populations in protected areas 
range from an annual budget of $500 USD per km\2\ in smaller fenced 
reserves to $2,000 USD per km\2\ for unfenced reserves (Packer et al. 
2013, p. 640). This includes but is not limited to costs associated 
with permanent and temporary staff, fencing installation and 
maintenance (fences can cost $3,000 USD per km to install), 
infrastructure maintenance, anti-poaching activities such as 
surveillance and snare/trap removal, wildlife restocking fees (both for 
lions killed by illegal poaching/snares as well as other trophy species 
killed by lions on the reserves), community outreach, and compensation 
for loss of livestock in surrounding communities. However, many 
management areas lack adequate funding (Packer et al. 2013, p. 640; 
Groom 2013, pp. 4-5; Barnett and Patterson 2005, p. 82).
    Of 12 protected areas assessed in western Africa, 6 had no budget 
for management activities or the budget was too low to conserve lion 
populations; nine reported having either no law enforcement activity or 
major deficiencies in staff and resources to conduct patrols. In 
Como[eacute] National Park, the staff was found to be too small for the 
size of the park (Henschel et al. 2014, p. 7). Protected areas in 
Guinea are essentially parks on paper only. They have no staff, 
management plan, or operating budget (Brugi[eacute]re 2012 in Henschel 
et al. 2014, p. 7). Although the WAP complex has received high scores 
for management effectiveness, the presence of 50,000 head of cattle 
inside W National Park indicates weak management. Livestock are rare in 
Arly-Pendjari, and lion density is higher; a higher management budget 
allocation is suspected to be the cause of the observed differences 
(Henschel et al. 2014, pp. 5-6). Across the lion's range, Africa's 
protected areas have generally failed to mitigate threats to large 
mammal populations, including the lion and its prey (Craigie et al. 
2010, entire).
    Poor management leads to many of the threats that lions face, 
including encroachment by pastoralists, increased poaching pressure, 
collapse of prey populations, and persecution by pastoralists 
(Brugi[eacute]re et al. 2015, pp. 519-520; Henschel et al. 2015, 
unpaginated; Henschel et al. 2014, pp. 5, 7; Henschel et al. 2010, p. 
38). Therefore, it can be said that management of protected areas that 
still harbor lions is inadequate to address the threats impacting 
lions, especially those in western Africa (Henschel 2015, unpaginated). 
Overall, investment in conservation activities is extremely low in 
western Africa, compared to central, eastern, and southern Africa. 
Countries in the former or current western Africa lion range are among 
the 50 poorest countries in the world, and six are classified as least 
developed countries. These countries will likely be unable to generate 
the resources required to secure their remaining lion populations 
(Henschel et al. 2014, pp. 7-8). Investment from the international 
community is needed to increase management effectiveness of these 
protected areas (Henschel et al. 2015, unpaginated).
    In India, most lions occur within five designated protected areas: 
Gir National Park and Gir Wildlife Sanctuary (Gir Protected Area) and 
Pania, Mitiyala, and Girnar sanctuaries (Bauer et al. 2015a, 
unpaginated; Banerjee and Jhala 2012, p. 1421; Singh and Gibson 2011, 
p. 1754; Jhala et al. 2009, pp. 3384, 3385; Nowell and Jackson 1996, p. 
38). Under India's Wild Life Protection Act of 1972 (Act No. 53 of 
1972; Chapter IV, sections 27, 28, 33, 35), entry into protected areas 
is regulated and certain activities are controlled and managed, 
including security of wild animals and grazing of livestock. In 2012, 
India's Ministry of Environment and Forests (2012, p. 22) declared the 
area 5 km from the boundary of Girnar Wildlife Sanctuary an Eco-
sensitive Zone for the long-term protection and conservation of the 
lion. This designation prohibits certain activities within the 
designated zone, such as mining, unregulated tourism, polluting 
industries, and unregulated felling of trees.
    Because of the protections afforded by the Government of Gujarat, 
threats that contributed to the decline of this population have been 
ameliorated and most threats faced by lions are not an immediate 
threat. Protections ensure food security, water availability, habitat 
suitability, and safety for these lions (Meena 2014, p. 26). However, 
because this population is small and isolated, it is vulnerable to 
extinction from stochastic events. Although a second location has been 
proposed to establish another free-ranging population geographically 
separate from Gir to reduce the risk of extinction of this population, 
translocation of lions from Gujarat are still pending (see Deleterious 
Effects Due to Small Population Sizes).

Climate Change

    Consideration of ongoing and projected climate change is a 
component of our analysis under the Act. The term ``climate change'' 
refers to a change in the mean, variability, or seasonality of climate 
variables over time periods of decades or hundreds of

[[Page 80031]]

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

[[Page 80032]]

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

[[Page 80033]]

could negatively affect reproduction. This species is also sensitive to 
rainfall due to its high water consumption rate (up to 30-40 liters per 
animal per day) (Du Troit 2005, as cited in The Heinz Center 2012, p. 
15; Whyte et al. 1995, pp. 84-85). Variation in the buffalo population 
then is tied to rainfall conditions year-to-year. Funston and Mills 
(2006, p. 20) observed that the buffalo population increases only 
during periods of average to above-average rainfall, which means that 
climate projections for a drier Africa will have detrimental impacts on 
the buffalo population. Lions are opportunistic predators that feed on 
a variety of prey. This flexibility in prey may aid lions in exhibiting 
some resiliency to changes in prey populations (The Heinz Center 2012, 
p. 12). However, as discussed under Loss of Prey Base and Human-Lion 
Conflict, the loss of prey species can result in lions shifting their 
diet towards livestock which may increase retaliatory killings by 
humans (Bauer and Kari 2001, as cited in Tumenta et al. 2013, p. 241; 
Whyte et al. 1995, p. 85).
    Variation in lion home ranges may have an impact on the frequency 
of human-lion conflict especially in situations where lion home ranges 
expand into areas inhabited by humans (Peterson et al. 2014, p. 562). 
The interplay between the types of climate, the density of prey, and 
seasonal variation in temperature and precipitation all affect lion 
home range. Areas with a more arid climate and small prey density are 
associated with larger home ranges, while temperate or tropical regions 
with higher prey density are associated with smaller home ranges. In 
addition, prey living in an arid climate tend to disperse, while prey 
in a wetter climate are more concentrated, leading to a larger and 
smaller home range, respectively (Tuqa et al. 2014, p. 2; Celesia et 
al. 2010, pp. 63, 67; Sogbohossou 2011, p. 17; Loveridge et al. 2009, 
p. 953). In southern Africa, where most of the lion populations are 
enclosed (fenced), variation in the species' home range may be more 
limited. Lion home ranges are also influenced by the season with ranges 
being smaller during the dry season and larger during the wet season. 
During the dry season, prey congregate around the few remaining water 
sources, concentrating prey species in a smaller area, shrinking the 
home range needed by the lion to find food. Conversely, home ranges 
expand during the wet season due to prey dispersal (Tuqa et al. 2014, 
p. 8).
    Climate projections point toward a drier climate for western, 
central, and southern Africa (Niang et al. 2014, p. 1209; Hijioka et 
al. 2014a, p. 1333; Carr et al. 2013, p. 14; Chidumayo et al. 2011, p. 
21). Drought in the western and central African regions is expected to 
increase by a rate of 5-8 percent by 2080 (UNEP 2007, p. 2). Although 
drier conditions might initially lead to the lion home range shrinking 
as prey congregate around remaining water sources (Sogbohoussou 2011, 
p. 133), Tuqa et al. (2014, p. 8) found that lion home ranges expand in 
the time after a drought. The reason for this expansion may be that, as 
prey populations around water sources are depleted, the lion has to 
travel greater distances to find prey. In addition, researchers found 
that lions move beyond reserve boundaries and into communal ranches 
where there will be greater conflict with humans (Tuqa et al. 2014, p. 
9). It is likely that lions prey on livestock, which will intensify 
human-lion conflict. To compound the issue, pastoralists in sub-Saharan 
Africa will often lead their herds into protected areas where lions 
occur during a drought in search of water, which increases the risk of 
lion predation (Tumenta et al. 2013, p. 240).
    When lion prey on livestock, they primarily focus on cattle 
(Patterson et al. 2004, p. 510). Out of all livestock that are 
domesticated in Africa, cattle have the highest monetary value, which 
means the loss of cattle to lion predation will have the most adverse 
effect on pastoralists (Tumenta et al. 2013, p. 240). Additionally, 
droughts affect the survival of livestock (Peterson et al. 2014, p. 
562). A study of the drought that occurred in Kenya in 2008-2009 found 
that mortality rates among the cattle population varied between 57 and 
64 percent in six districts (Dolrenry 2013, p. 47; Zwaagstra et al. 
2010, p. 21). Such high mortality may make pastoralists less tolerant 
of lion predation and may increase the frequency of retaliatory 
killings (Peterson et al. 2014, p. 562).
    Climate change may increase the number and intensity of disease 
outbreaks in lion prey species, as well as lions (The Heinz Center 
2012, p. 12; Baylis 2006, p. 4). Diseases can be directly and 
indirectly affected by climate change by impacting distribution, the 
timing of outbreaks, and the intensity of outbreaks (Baylis 2006, p. 
4). Higher temperatures may increase the rates of development of 
pathogens and parasites, shorten generation times, and increase the 
number of generations per year, increasing the population (Baylis 2006, 
p. 8; Thuiller et al. 2006, p. 435). Temperatures can have impacts on 
vectors (e.g., ticks and mosquitoes) and hosts that may further 
influence the spread of diseases (Baylis 2006, pp. 9, 11) and increase 
risks of extinctions (Thuiller et al. 2006, p. 435). Additionally, 
rainfall conditions also affect the susceptibility of animals to 
disease outbreaks (Thuiller et al. 2006, p. 435). Munson et al. (2008) 
concluded that severe climate change could synchronize temporal and 
spatial convergence of multiple infectious agents, triggering epidemics 
with greater mortality than infections from a single pathogen.

Conservation Measures in Place To Protect Lions

    There has been awareness for several years that conservation 
strategies need to be implemented for the lion due to the apparent 
decrease in its population numbers (Hamunyela et al. 2013, p. 1; 
Henschel et al. 2010, p. 34; Gebresenbet et al. 2009, p. 5; IUCN 2006a, 
b, entire). Prior to 2006, institutional inconsistencies throughout the 
lion's African range resulted in poor lion conservation policies and 
little to no enforcement of existing laws (IUCN 2006b, p. 18). As 
mentioned, in 2005 and 2006, nongovernmental organizations (NGOs) and 
several governments at various levels organized two regional lion 
conservation workshops. Species specialists, wildlife managers, and 
government officials attended these regional workshops in order to 
provide range country governments with frameworks for developing their 
own national action plans for the conservation of lions. Over 50 lion 
specialists, representing all lion range countries, participated in 
these workshops (Henschel et al. 2010, p. 34). During the workshops, 
lion experts collectively assessed what they believed to be the then-
current status of African lions based on a variety of information, and 
subsequently identified 86 African LCUs. This information was then used 
as a framework to identify lion areas, strongholds, and potential 
strongholds by Riggio et al. (2013, p. 32).
    Many African countries with very small lion populations have 
developed or updated their conservation plans for the lion. Some of 
these include Benin, Cameroon, Uganda, and Malawi. Some range countries 
participate in transboundary conservation projects and are 
collaborating on transboundary lion conservation initiatives for shared 
lion populations. Most range countries have a national lion action plan 
or strategies in place, particularly if there are economic incentives 
for them to have viable lion populations (Groom 2013, p. 4; Namibia 
2013, pp. 11-12; Zambia Wildlife Authority 2012, p.3;

[[Page 80034]]

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

[[Page 80035]]

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

[[Page 80036]]

and 1982, eight lions were killed, whereas six lions were killed 
between 1998 and 2007 (Mesochina et al. 2010b, p. 35). While fewer 
lions are being killed than in the previous decades, problems remain, 
including lack of resources, lack of manpower, and corruption within 
the range countries.
    Current governmental management of lions in countries such as 
Malawi, Tanzania, and Zambia are managed by the Problem Animal Control 
units (Mesochina et al. 2010a, p. 41; Mesochina et al. 2010b, p. 36). 
When lion attack incidents occur, Problem Animal Control dispatches 
officials to investigate the problems. If the problem lion is located, 
it is either removed or eliminated. When properly funded, this program 
has helped in reducing not only conflicts between lions and humans but 
also has driven down the numbers of lions killed. Between 2005 and 
2009, there were 116 reported cases of lions killed, with the number of 
lions killed being less than 50 per year in Tanzania (Mesochina et al. 
2010a, p. 41). However, limitations of resources (including both 
manpower and funds) have hampered the effectiveness of these officials 
in responding to these incidents. In addition, many Problem Animal 
Control interventions resulted in the death of the lion (Mesochina et 
al. 2010a, p. 41; Chardonnet et al. 2009, p. 36). Even in cases of 
translocation, the lions that were being transported often end up 
injured or continue to pose problems to the community (Bauer et al. 
2007, p. 91).
    NGOs are also assisting in protecting lions. Intervention by NGOs 
often takes the form of interacting with the local community 
(Winterbach et al. 2010, p. 98). Lion Guardians, which operates in 
Kenya and Tanzania, recruits and educates local young men to monitor 
and track lion movement and warn herders of lion presence in the area, 
recover lost livestock, reinforce protective fencing, and intervene to 
stop lion hunting parties, thereby mitigating or preventing possible 
human-lion conflict (Hazzah et al. 2014, p. 853; Lion Guardians 2013, 
p. 7; Lion Guardians 2012, p. 3). From 2010 to 2013, Lion Guardians 
maintained a recovery rate of lost livestock of more than 85, totaling 
over $1.5 million USD; in 2014 alone, more than 20,000 livestock (93 
percent) were recovered (Lion Guardians 2014, p. 7; Lion Guardians 
2013, p. 6). Since 2010, 1,700 bomas have been reinforced to reduce 
depredation of livestock. End-of-year sampling shows that more than 90 
percent of reinforced bomas sampled did not experience further 
depredation (Lion Guardians 2014, p. 7; Lion Guardians 2013, p. 6). 
Additionally, 103 lion hunts were stopped or prevented between 2010 and 
2014 (Lion Guardians 2014, p. 6; Lion Guardians 2013, p. 5). Lastly, in 
the years of Lion Guardians operations, lion kills have decreased by 95 
percent and the number of lions has steadily increased; a total of 286 
lions have been documented in the Amboseli-Tsavo ecosystem (Lion 
Guardians 2014, p. 6; Lion Guardians 2013, p. 5).
    In addition, Lion Guardians work with tribal elders to dissuade 
young men from killing lions for ceremonial purposes. Historically, the 
killing of lions through ritualized lion hunts called ilmurran is 
rewarded with gifting of cows and other rewards (Lion Guardians 2012, 
p. 5; Goldman et al. 2010, p. 334). After introducing village elders to 
the Lion Guardians program first hand, many return home to their 
village and give their blessings to the project. This education led to 
significant results; on August 11, 2013, two Lion Guardians stopped a 
group of hunters who were planning to hunt a lion in retaliation for 
the lion preying on their livestock. The local village elders fined the 
potential hunters two cattle each for going on a lion hunt, marking a 
gradual but significant shift in the cultural attitudes regarding the 
lion (Hazzah et al. 2014, p. 858; Lion Guardians 2013, p. 20). Between 
2007 and 2014, only five lions had been killed in territories where 
Lion Guardians operates, in contrast to more than 100 lions killed in 
adjacent areas (Lion Guardians 2013, p. 5). Furthermore, reduced lion 
mortality was sustained across multiple years, resulting in the reserve 
having one of the highest lion densities in Africa (Hazzah et al. 2014, 
p. 857; Schuette et al. 2013, p. 149). Despite the success of this 
program, retaliatory as well as ceremonial killings of lions outside 
the program areas remain a threat to the species.
    We found that many of the lion range states are trying to address 
lion conservation through the establishment of protected areas, 
wildlife management areas, wildlife corridors, and reconnecting 
habitat. In some areas, creating incentives for lion conservation is 
occurring through community conservation programs in range countries. 
In other cases, participatory strategies have been implemented to 
enhance local tolerance for large carnivores in Africa. An increasing 
number of programs encourage local communities to solve problems that 
arise from human-lion conflict without killing lions. However, the 
effectiveness of these measures still ranges from successful to 
unsuccessful, due in part to lack of resources, political will, and 
infighting. It is imperative that range countries continue to recognize 
and support the role that local communities play in lion conservation. 
Greater support by countries to address the needs of local communities, 
and thereby address the needs of lions, may be the single-most 
important role these countries can play in changing the trajectory of 
lion declines.

Finding

    Section 4 of the Act (16 U.S.C. 1533) and implementing regulations 
(50 CFR part 424) set forth procedures for adding species to, removing 
species from, or reclassifying species on the Federal Lists of 
Endangered and Threatened Wildlife and Plants. Under section 4(a)(1) of 
the Act, a species may be determined to be an endangered species or a 
threatened species based on any of the following five factors:

    (A) The present or threatened destruction, modification, or 
curtailment of its habitat or range;
    (B) Overutilization for commercial, recreational, scientific, or 
educational purposes;
    (C) Disease or predation;
    (D) The inadequacy of existing regulatory mechanisms; or
    (E) Other natural or manmade factors affecting its continued 
existence.

    A species is ``endangered'' for purposes of the Act if it is in 
danger of extinction throughout all or a significant portion of its 
range and is ``threatened'' if it is likely to become endangered within 
the foreseeable future throughout all or a significant portion of its 
range. The ``foreseeable future'' is the period of time over which 
events or effects reasonably can or should be anticipated, or trends 
extrapolated.
    As required by the Act, we conducted a review of the status of the 
species and considered the five factors in assessing whether the lion 
is in danger of extinction throughout all or a significant portion of 
its range or likely to become endangered within the foreseeable future 
throughout all or a significant portion of its range. We examined the 
best scientific and commercial information available regarding the 
past, present, and future threats faced by the lion. We reviewed the 
petition, information available in our files, other available published 
and unpublished information, and comments received from peer reviewers 
and the general public.
    When considering what factors might constitute threats to a 
species, we must look beyond the mere exposure of the species to a 
factor to evaluate whether the species may respond to the factor in a 
way that causes actual impacts to the species. If there is exposure to 
a factor

[[Page 80037]]

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

[[Page 80038]]

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

[[Page 80039]]

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

[[Page 80040]]

across its range, especially in western Africa where most protected 
areas are experiencing severe management deficiencies.
    Based on the best scientific and commercial information, we find 
that several factors are negatively impacting the lion and contributing 
to the risk of extinction. However, we find there is a substantial 
difference in the magnitude of these threats to the risk of extinction 
between the subspecies P. l. leo and P. l. melanochaita. Based on 
current population estimates, projected population trends, and the 
threats described herein, we find that the subspecies P. l. leo and P. 
l. melanochaita qualify for different statuses under the Act.

Finding for Panthera leo leo

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

[[Page 80041]]

areas to protect natural resources from negative impacts of human 
populations. However, weak management of protected areas has been 
documented across the lion's range, especially in western Africa where 
most protected areas are experiencing severe management deficiencies, 
including the lack of a budget or a budget insufficient to carry out 
management activities.
    The lion population in India is one population of P. l. leo that is 
increasing and could potentially be considered a viable population 
based on the number of lions. However, intense management, including 
healthcare interventions, may interfere with natural selection 
processes by ensuring the survival of unfit lions, which facilitates 
the propagation of deleterious genes in the population. This population 
is also running out of area to expand. Being a small, isolated 
population and less genetically diverse, it is more vulnerable to the 
loss of any individuals due to environmental and stochastic events, and 
more prone to local extinction events.
    As previously stated, threats to the lion are expected to continue 
or increase in conjunction with predicted human population growth. The 
human population, and thus negative impacts to lions, as well as 
decreases in lion populations, associated with human population growth, 
is expected to increase substantially by 2050. If regional trends 
continue at their current rate, western and central Africa will likely 
lose a third of its population in 5 years and half the population in 10 
years. Lion bone may be increasingly used as a replacement for tiger 
bone in traditional Asian medicine and in Asian luxury products. 
Therefore, trade in lion bone could become lucrative, spur considerable 
demand from suppliers of the black market, result in extensive poaching 
of wild lions, and have significant impacts to lion populations. 
Additionally, future development in India could alter habitat vital for 
dispersal. Tolerance to loss of livestock may also wane as traditional 
beliefs and traditional value systems are rapidly changing under the 
influence of globalization. Furthermore, effects of climate change on 
lion habitat are projected to manifest as early as 2040. Under climate 
change scenarios, a broad swath of potential distributional area in 
western Africa is projected to become distinctly less suitable or even 
uninhabitable. Increases in rainfall predicted for India may not have 
detrimental impacts on lion habitat; however, increased risks of 
flooding could result in increased mortality, and post-flooding 
conditions could be conducive to disease outbreaks and are a serious 
concern to the persistence of the lion population as this population is 
more vulnerable to stochastic events and local extinction.
    Threats acting on P. l. leo have contributed to large reductions in 
the subspecies' range and suitable habitat, abundance, and number and 
connectivity of populations. The subspecies has reached critically low 
numbers of individuals and potentially viable populations. Furthermore, 
while one small population may be increasing, we are not aware of any 
information indicating that the overall trend of large declines in the 
subspecies range, abundance, and connectivity, will reverse course.
    Threats continue to act on this subspecies. Due to small population 
size and lack of connectivity between populations, most populations are 
not able to recover from the loss of suitable habitat or individuals. 
Furthermore, because all populations are small and isolated, the 
subspecies lacks resiliency to recover from stochastic or catastrophic 
events and is thus highly vulnerable to extirpation. Threats are 
currently affecting the subspecies and the impacts on the subspecies 
are expected to continue or even intensify over time as the human 
population increases and as climate change progresses, negatively 
impacting availability of suitable habitat, lion distribution, and lion 
numbers. Based on the current distribution and size of P. l. leo 
populations, the current threats acting on this subspecies, the impacts 
of those threats, and the impacts of future threats and climate change 
on lion distribution, lion numbers, habitat, prey availability, 
susceptibility to disease, loss of lions via human-lion conflict and 
trophy hunting, and resiliency to stochastic and catastrophic events, 
we find that the viability of this subspecies is compromised and will 
not be resistant or resilient to ongoing and future threats. Therefore, 
we find that P. l. leo is in danger of extinction throughout its range 
and list the subspecies as endangered.

Finding for Panthera leo melanochaita

    The range of P. l. melanochaita includes the southern and eastern 
African regions. Although this subspecies has experienced range 
reduction, a decline in the number of populations, and a decline in the 
number of lions, it remains relatively widespread. Currently, there are 
approximately 17,730 P. l. melanochaita lions distributed among 68 
protected areas, with larger populations in Botswana, Kenya, Namibia, 
South Africa, Tanzania, Zambia, and Zimbabwe. Between 1993 and 2014, 
the lion population in eastern Africa declined by 59 percent. In 
southern Africa the lion population increased by 8 percent during the 
same time period. Most of the increasing populations contributing to 
this trend are small, fenced reserves. However, one of the largest 
populations in southern Africa, Okavango, and populations in 6 unfenced 
reserves in Botswana, Namibia, and Zimbabwe declined. Although there 
are larger populations of P. l. melanochaita that may meet the 
suggested MVP, almost all lion populations in Africa that historically 
exceeded 500 individuals, are declining.
    Expansion of human settlements, agriculture, and/or livestock 
grazing is occurring in or on the major populations and is particularly 
a threat in eastern Africa and some parts of southern Africa. In 
particular, expansion of agriculture and livestock grazing is occurring 
in or around major populations in Kenya, Tanzania, and Zambia and both 
are major threats to lion survival in these countries. Expansion of 
human settlements and activities into lion habitat renders it 
unsuitable for lions, primarily because human expansion results in 
reduced availability of wild prey and lion mortality due to increases 
in human-lion conflict. Both of these factors influence the 
distribution and population viability of lions. However, in some parts 
of southern Africa, lions are repopulating areas where lions were 
recently extirpated due to adequate protection of habitat and prey.
    Significant decreases in prey abundance have occurred in protected 
areas throughout Africa, including Botswana, Kenya, Mozambique, Sudan, 
Zambia, and Zimbabwe. Herbivore populations have decreased by 52 
percent in eastern Africa, although they have increased by 24 percent 
in southern Africa. Protected areas in Ethiopia, Mozambique, Tanzania, 
and Zambia are increasingly settled; decreases in prey abundance in 
African protected areas are driven by human population growth, 
especially along the boundaries of protected areas where human 
population growth rates are high, encroachment and habitat loss occurs, 
and people are dependent on bushmeat. Additionally, many communities 
lack the rights over land and in most cases in Botswana, Tanzania, 
Zambia, and Zimbabwe, the government retains a significant portion of 
revenue from wildlife; therefore, those that bear the costs of wildlife 
do not receive benefits, and bushmeat hunting is the only way to 
benefit from

[[Page 80042]]

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

[[Page 80043]]

wild lions to meet demand, and have significant impacts to lion 
populations.
    Although there are laws in place in lion range countries that are 
meant to protect wildlife, including lions and their prey species, the 
drastic and continuing decline of the species and its prey in some 
parts of its range indicate these regulatory mechanisms are not 
adequate to ameliorate threats to the P. l. melanochaita throughout its 
range. Furthermore, national and international conservation strategies 
rely on protected areas to protect natural resources from negative 
impacts of human populations. However, weak management of protected 
areas has been documented across the lion's range.
    As indicated above, P. l. melanochaita remains relatively 
widespread and some remaining populations are large enough to be 
considered viable. Therefore, due to the size of some populations, the 
number of remaining populations, and the stability or increasing status 
of some populations, we find that P. l. melanochaita is not currently 
in danger of extinction. However, the overall population of the 
subspecies continues to decline and threats to the lion are expected to 
continue or increase in the future in conjunction with predicted human 
population growth. If regional trends in lion populations continue at 
the current rate, eastern Africa will lose a third of its lion 
population in 20 years and half the population in 30 years. Effects of 
climate change on lion habitat are projected to manifest as early as 
2040. Although climate change effects on potential lion distribution 
are projected to be more neutral in eastern Africa than across the 
entire range, southern Africa, where the broadest areas of suitable 
conditions occur, is projected to become less suitable because of 
climate change. Specifically, park areas, including the ``Etosha Pan, 
Lake Opnono, Cuvelai Drainage, Kalahari Gemsbok, and Kgalagadi 
Transfrontier Park areas'' are projected to decline substantially in 
suitability for lions. In addition, reforms to trophy hunting have been 
made to ensure sustainability of trophy hunting, but these reforms have 
been implemented in only a few places. Furthermore, demand for lion 
bone is expected to increase in the future and high prices for lion 
bone provide incentive to poach wild lions. As a result of the likely 
impacts of these threats, it is reasonable to conclude that the 
population of P. l. melanochaita is likely to be drastically reduced 
and fragmented in the foreseeable future, limiting the ability of the 
subspecies to recover from stochastic and catastrophic events. 
Therefore, we find that this subspecies is likely to become an 
endangered species within the foreseeable future and we are listing P. 
l. melanochaita as a threatened species.

Significant Portion of Its Range

    Under the Act and our implementing regulations, a species may 
warrant listing if it is endangered or threatened throughout all or a 
significant portion of its range. The term ``species'' includes ``any 
subspecies of fish or wildlife or plants, and any distinct population 
segment [DPS] of any species of vertebrate fish or wildlife which 
interbreeds when mature.'' We published a final policy interpreting the 
phrase ``Significant Portion of its Range'' (SPR) (79 FR 37578, July 1, 
2014). The final policy states that (1) if a species is found to be 
endangered or threatened throughout a significant portion of its range, 
the entire species is listed as endangered or threatened, respectively, 
and the Act's protections apply to all individuals of the species 
wherever found; (2) a portion of the range of a species is 
``significant'' if the species is not currently endangered or 
threatened throughout all of its range, but the portion's contribution 
to the viability of the species is so important that, without the 
members in that portion, the species would be in danger of extinction, 
or likely to become so in the foreseeable future, throughout all of its 
range; (3) the range of a species is considered to be the general 
geographical area within which that species can be found at the time 
the Service or the National Marine Fisheries Service makes any 
particular status determination; and (4) if a vertebrate species is 
endangered or threatened throughout an SPR, and the population in that 
significant portion is a valid DPS, we will list the DPS rather than 
the entire taxonomic species or subspecies.
    We found the lion subspecies P. l. leo to be in danger of 
extinction throughout its range, and the subspecies P. l. melanochaita 
likely to become endangered within the foreseeable future throughout 
its range. Therefore, no portions of the species' range are 
``significant'' as defined in our SPR policy, and no additional SPR 
analysis is required.

4(d) Rule for Panthera leo melanochaita

    The purposes of the ESA are to provide a means whereby the 
ecosystems upon which endangered species and threatened species depend 
may be conserved, to provide a program for the conservation of such 
endangered species and threatened species, and to take such steps as 
may be appropriate to achieve the purposes of the treaties and 
conventions set forth in the ESA. When a species is listed as 
endangered, certain actions are prohibited under section 9 of the ESA 
and are implemented through our regulations in 50 CFR 17.21. These 
include, among others, prohibitions on take within the United States, 
within the territorial seas of the United States, or upon the high 
seas; import; export; and shipment in interstate or foreign commerce in 
the course of a commercial activity. Exceptions to the prohibitions for 
endangered species may be granted in accordance with section 10 of the 
ESA and our regulations at 50 CFR 17.22.
    The ESA does not specify particular prohibitions and exceptions to 
those prohibitions for threatened species. Instead, under section 4(d) 
of the ESA, the Secretary, as well as the Secretary of Commerce 
depending on the species, was given the discretion to issue such 
regulations as deemed necessary and advisable to provide for the 
conservation of such species. The Secretary also has the discretion to 
prohibit by regulation with respect to any threatened species any act 
prohibited under section 9(a)(1) of the ESA. Exercising this 
discretion, the Service has developed general prohibitions in the ESA 
regulations (50 CFR 17.31) and exceptions to those prohibitions (50 CFR 
17.32) that apply to most threatened species. Under 50 CFR 17.32, 
permits may be issued to allow persons to engage in otherwise 
prohibited acts for certain purposes.
    Under section 4(d) of the ESA, the Secretary, who has delegated 
this authority to the Service, may also develop specific prohibitions 
and exceptions tailored to the particular conservation needs of a 
threatened species. In such cases, the Service issues a 4(d) rule that 
may include some or all of the prohibitions and authorizations set out 
in 50 CFR 17.31 and 17.32, but which also may be more or less 
restrictive than the general provisions at 50 CFR 17.31 and 17.32. For 
P. l. melanochaita, the Service has determined that a 4(d) rule is 
necessary and advisable.
    We are adding a 4(d) (special) rule for P. l. melanochaita at 50 
CFR 17.40(r). This 4(d) rule maintains all of the prohibitions and 
exceptions codified in 50 CFR 17.31 and 17.32 with regard to this 
subspecies and supersedes the import exemption found in 50 CFR 17.8 for 
threatened wildlife listed in Appendix II of CITES, such that a 
threatened species import permit under 50 CFR 17.32 is now required for 
the importation of all P. l. melanochaita specimens. Therefore, through 
the promulgation of this 4(d) rule, the

[[Page 80044]]

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

[[Page 80045]]

under the Act, such as import, export, interstate and foreign commerce 
and take, including noncommercial imports of live lions, we can ensure 
that live lions are not used to supplement the trade in lion bones.
    Therefore, we find that regulation of the importation of all P. l. 
melanochaita parts and products, including live animals and sport-
hunted trophies, will ensure that imported specimens are obtained in a 
manner that is consistent with the purposes of the Act and the 
conservation of the subspecies in the wild.
    Our threatened species permitting regulations at 50 CFR 17.32 
provide issuance criteria for threatened species permits (50 CFR 
17.32(a)(2)), but do not specify what would constitute the enhancement 
of propagation or survival with regard to authorizing the import of 
parts or products of P. l. melanochaita, including sport-hunted 
trophies. Therefore, when making a determination of whether an 
otherwise prohibited activity enhances the propagation or survival of 
P. l. melanochaita, the Service will examine the overall conservation 
and management of the subspecies in the country where the specimen 
originated and whether that management of the subspecies addresses the 
threats to the subspecies (i.e., that it is based on sound scientific 
principles and that the management program is actively addressing the 
current and longer term threats to the subspecies). In that review, we 
will evaluate whether the import contributes to the overall 
conservation of the species by considering whether the biological, 
social, and economic aspects of a program from which the specimen was 
obtained provide a net benefit to the subspecies and its ecosystem.
    The Service will evaluate any application received that involves P. 
l. melanochaita in the context of enhancement of propagation or 
survival permitting in accordance with our threatened species 
permitting regulations at 50 CFR 17.32 and issuance criteria for 
threatened species permits (50 CFR 17.32(a)(2)). These include, in 
addition to the general permitting criteria in 50 CFR 13.21(b):

    (i) Whether the purpose for which the permit is required is 
adequate to justify removing from the wild or otherwise changing the 
status of the wildlife sought to be covered by the permit;
    (ii) The probable direct and indirect effect that issuing the 
permit would have on the wild populations of the wildlife sought to 
be covered by the permit;
    (iii) Whether the permit, if issued, would in any way, directly 
or indirectly, conflict with any known program intended to enhance 
the survival probabilities of the population from which the wildlife 
sought to be covered by the permit was or would be removed;
    (iv) Whether the purpose for which the permit is required would 
be likely to reduce the threat of extinction facing the species of 
wildlife sought to be covered by the permit;
    (v) The opinions or views of scientists or other persons or 
organizations having expertise concerning the wildlife or other 
matters germane to the application; and
    (vi) Whether the expertise, facilities, or other resources 
available to the applicant appear adequate to successfully 
accomplish the objectives stated in the application.

    In addition to these factors, particularly in relation to sport 
hunting, we find the IUCN Species Survival Commission (SSC) Guiding 
Principles on Trophy Hunting as a Tool for Creating Conservation 
Incentives, Ver. 1.0 (IUCN SSC 2012), to provide useful principles, 
which, considered in conjunction with our threatened species issuance 
criteria, will aid the Service when making an enhancement finding for 
importation of sport-hunted trophies of P. l. melanochaita. This 
document sets out guidance from experts in the field on the use of 
trophy hunting as a tool for ``creating incentives for the conservation 
of species and their habitats and for the equitable sharing of the 
benefits of use of natural resources'' (IUCN SSC 2012, p. 2) and 
recognizes that recreational hunting, particularly trophy hunting, can 
contribute to biodiversity conservation and more specifically, the 
conservation of the hunted species.
    The SSC document lays out five guiding principles that, considered 
in conjunction with our threatened species issuance criteria, will aid 
the Service when making an enhancement finding for importation of 
sport-hunted trophies of P. l. melanochaita:

    (a) Biological sustainability: The hunting program cannot 
contribute to the long-term decline of the hunted species. It should 
not alter natural selection and ecological function of the hunted 
species or any other species that share the habitat. The program 
should not inadvertently facilitate poaching or illegal trade in 
wildlife by acting as a cover for such illegal activities. The 
hunting program should also not manipulate the ecosystem or its 
component elements in a way that alters the native biodiversity.
    (b) Net Conservation Benefit: The biologically sustainable 
hunting program should be based on laws, regulations, and 
scientifically based quotas, established with local input, that are 
transparent and periodically reviewed. The program should produce 
income, employment, and other benefits to create incentives for 
reducing the pressure on the target species. The program should 
create benefits for local residents to co-exist with the target 
species and other species. It is also imperative that the program is 
part of a legally recognized governance system that supports 
conservation.
    (c) Socio-Economic-Cultural Benefit: A well-managed hunting 
program can serve as a conservation tool when it respects the local 
cultural values and practices. It should be accepted by most members 
of the community, involving and benefiting local residents in an 
equitable manner. The program should also adopt business practices 
that promote long-term economic sustainability.
    (d) Adaptive Management: Planning, Monitoring, and Reporting: 
Hunting can enhance the species when it is based on appropriate 
resource assessments and monitoring (e.g., population counts, trend 
data), upon which specific science-based quotas and hunting programs 
can be established. Resource assessments should be objective, well 
documented, and use the best science available. Adaptive management 
of quotas and programs based on the results of resource assessments 
and monitoring is essential. The program should monitor hunting 
activities to ensure that quotas and sex/age restrictions of 
harvested animals are met. The program should also generate reliable 
documentation of its biological sustainability and conservation 
benefits.
    (e) Accountable and Effective Governance: A biologically 
sustainable trophy-hunting program should be subject to a governance 
structure that clearly allocates management responsibilities. The 
program should account for revenues in a transparent manner and 
distribute net revenues to conservation and community beneficiaries 
according to properly agreed decisions. All necessary steps to 
eliminate corruption should be taken and to ensure compliance with 
all relevant national and international requirements and regulations 
by relevant bodies such as administrators, regulators and hunters.

    The Service's approach to enhancement findings for the importation 
of sport-hunted trophies of P. l. melanochaita is consistent with the 
purpose and intent of the Endangered Species Act. Before we will 
authorize the importation of a sport-hunted trophy, we must determine 
that the trophy hunting program is managed to ensure the long-term 
survival of the species. In many parts of the world, wildlife exists 
outside of protected areas and must share the same habitat and compete 
with humans living in these areas for space and resources. If 
communities that share these resources with wildlife do not perceive 
any benefits from the presence of wildlife, they may be less willing to 
tolerate the wildlife. However, under certain circumstances, trophy 
hunting can address this problem by making wildlife more valuable to 
the local communities and encourage community support for managing and 
conserving the hunted species, as well as other species.
    When evaluating whether the importation of a trophy of P. l.

[[Page 80046]]

melanochaita would be authorized pursuant to 50 CFR 17.32, in 
accordance with our threatened species issuance criteria, we will 
examine how a country's management program for lions addresses the 
three main threats that have led to the decline of the subspecies: 
Habitat loss, loss of prey base, and human-lion conflict. When 
examining a management program and whether trophies taken as part of 
that program meet the issuance criteria, we would study a number of 
factors. Some of the factors we would consider include whether the 
program is based on sound scientific information and identifies 
mechanisms that would arrest the loss of habitat or increase available 
habitat (i.e., by establishing protected areas and ensuring adequate 
protection from human encroachment). We would consider whether the 
management program actively address the loss of the lion's prey base by 
addressing poaching or unsustainable offtake within the country. A 
component of a management plan from which trophy imports would meet the 
issuance criteria would be whether there are government incentives in 
place that encourage habitat protection by private landowners and 
communities and incentives to local communities to reduce the incursion 
of livestock into protected areas or to actively manage livestock to 
reduce conflicts with lions. We would examine if the hunting component 
of the management program supports all of these efforts by looking at 
whether hunting concessions/tracts are managed to ensure the long-term 
survival of the lion, its prey base, and habitat. As stated previously, 
hunting can generate significant economic benefits if properly 
conducted. In looking at whether we would be able to authorize the 
import of a trophy under the issuance criteria of 50 CFR 17.32, we 
would examine if the trophy hunting provides financial assistance to 
the wildlife department to carry out elements of the management program 
and if there is a compensation scheme or other incentives to benefit 
local communities that may be impacted by lion predation. We would also 
consider how a U.S. hunter's participation in the hunting program 
contributes to the overall management of lions within a country.
    Management programs for P. l. melanochaita would be expected to 
address, but are not limited to, evaluating population levels and 
trends; the biological needs of the species; quotas; management 
practices; legal protection; local community involvement; and use of 
hunting fees for conservation. In evaluating these factors, we will 
work closely with the range countries and interested parties to obtain 
the information. By allowing entry into the United States of P. l. 
melanochaita trophies from range countries that have science-based 
management programs, we anticipate that other range countries would be 
encouraged to adopt and financially support the sustainable management 
of lions that benefits both the species and local communities. In 
addition to addressing the biological needs of the subspecies, a 
scientifically based management program would provide economic 
incentives for local communities to protect and expand P. l. 
melanochaita habitat.
    As stated, under this 4(d) rule any person wishing to conduct an 
otherwise prohibited activity, including all imports of P. l. 
melanochaita specimens, must first obtain a permit under 50 CFR 17.32. 
As with all permit applications submitted under 50 CFR 17.32, the 
individual requesting authorization to import a sport-hunted trophy of 
P. l. melanochaita bears the burden of providing information in their 
application showing that the activity meets the requirements for 
issuance criteria under 50 CFR 17.32. In some cases for imports, such 
as sport-hunted trophies, it is not always possible for the applicant 
to provide all of the necessary information needed by the Service to 
make a positive determination under the Act to authorize the activity. 
For the import of sport-hunted trophies of P. l. melanochaita, the 
Service will typically consult with the range country to the extent 
practicable and other interested parties to obtain necessary 
information. The Service has the discretion to make the required 
findings on sport-hunted trophy imports of P. l. melanochaita on a 
country-wide basis, although individual import permits will be 
evaluated and issued or denied for each applicant. While the Service 
may make enhancement findings for sport-hunted trophy imports of P. l. 
melanochaita on a country-wide basis, the Service encourages the 
submission of information from individual applicants. We would rely on 
the information available to the Service and may rely on information 
from sources other than the applicant when making a permitting 
decision.

Effects of This Rule

    This action revises the taxonomic classification of the Asiatic 
lion (currently classified as P. l. persica and listed as an endangered 
species under the Act) to P. l. leo based on a taxonomic change. This 
rule revises 50 CFR 17.11(h) to add P. l. leo subspecies and the P. l. 
melanochaita subspecies to the List of Endangered and Threatened 
Wildlife as an endangered species and a threatened species, 
respectively. This rule establishes a 4(d) rule for P. l. melanochaita, 
which implements all of the prohibitions and exceptions under 50 CFR 
17.31 and 17.32 and requires a threatened species import permit under 
50 CFR 17.32 for the importation of all P. l. melanochaita specimens. 
Under the 4(d) rule, the import exemption found in 50 CFR 17.8 for 
threatened wildlife listed in Appendix II of CITES does not apply to 
this subspecies. Therefore, through the promulgation of this 4(d) rule, 
the presumption of legality provided under section 9(c)(2) of the Act 
for the otherwise lawful importation of wildlife listed in Appendix II 
of CITES that is not an endangered species listed pursuant to section 4 
of the Act does not apply to this subspecies (See: 4(d) Rule for 
Panthera leo melanochaita).

Available Conservation Measures

    Conservation measures provided to species listed as endangered or 
threatened under the Act include recognition of conservation status, 
requirements for Federal protection, and prohibitions against certain 
practices. Recognition through listing encourages and results in public 
awareness and conservation actions by Federal and State governments in 
the United States, foreign governments, private agencies and groups, 
and individuals.
    Section 7(a) of the Act, as amended, and as implemented by 
regulations at 50 CFR part 402, requires Federal agencies to evaluate 
their actions that are to be conducted within the United States or upon 
the high seas, with respect to any species that is proposed to be 
listed or is listed as endangered or threatened. Because P. l. leo and 
P. l. melanochaita are not native to the United States, no critical 
habitat is being proposed for designation with this rule. Regulations 
implementing the interagency cooperation provision of the Act are 
codified at 50 CFR part 402. Section 7(a)(2) of the Act requires 
Federal agencies to ensure that activities they authorize, fund, or 
carry out are not likely to jeopardize the continued existence of a 
listed species or to destroy or adversely modify its critical habitat. 
If a proposed Federal action may adversely affect a listed species, the 
responsible Federal agency must enter into formal consultation with the 
Service. Currently, with respect to the lion, no Federal activities are 
known that would require consultation.

[[Page 80047]]

    Section 8(a) of the Act authorizes the provision of limited 
financial assistance for the development and management of programs 
that the Secretary of the Interior determines to be necessary or useful 
for the conservation of endangered or threatened species in foreign 
countries. Sections 8(b) and 8(c) of the Act authorize the Secretary to 
encourage conservation programs for foreign listed species, and to 
provide assistance for such programs, in the form of personnel and the 
training of personnel.
    Section 9 of the Act and our implementing regulations at 50 CFR 
17.21 and 50 CFR 17.31 set forth a series of general prohibitions that 
apply to all endangered and threatened wildlife, respectively, except 
where a 4(d) rule applies to threatened wildlife, in which case the 
4(d) rule contains all the applicable prohibitions and exceptions. 
Under the 4(d) rule for P. l. melanochaita, all of the prohibitions 
under 50 CFR 17.31 apply to P. l. melanochaita specimens. These 
prohibitions, at 50 CFR 17.21 and 17.31, in part, make it illegal for 
any person subject to the jurisdiction of the United States to ``take'' 
(includes harass, harm, pursue, hunt, shoot, wound, kill, trap, 
capture, or to attempt any of these) within the United States or upon 
the high seas; import or export; deliver, receive, carry, transport, or 
ship in interstate or foreign commerce, by any means whatsoever, in the 
course of commercial activity; or sell or offer for sale in interstate 
or foreign commerce any lion specimens. It also is illegal to possess, 
sell, deliver, carry, transport, or ship any such wildlife that has 
been taken in violation of the Act. Permits may be issued to carry out 
otherwise prohibited activities involving endangered and threatened 
wildlife species under certain circumstances. Regulations governing 
permits for endangered species, such as P. l. leo, are codified at 50 
CFR 17.22. Regulations governing permits for threatened species, such 
as P. l. melanochaita, are codified at 50 CFR 17.32. Certain exceptions 
apply to agents of the Service and State conservation agencies.

Summary of Comments and Recommendations

    We based this action on a review of the best scientific and 
commercial information available, including all information received 
during the public comment period. In the October 2014 proposed rule, we 
requested that all interested parties submit information that might 
contribute to development of a final rule. We also contacted 
appropriate scientific experts and organizations and invited them to 
comment on the proposed listing. We received tens of thousands of 
comments.
    We reviewed all comments we received from the public for 
substantive issues and new information regarding the proposed listing 
of this species, and we address those comments below. Overall, most 
commenters supported the proposed listing, but did not provide 
additional scientific or commercial data for consideration. We have not 
included responses to comments that supported the listing decision but 
did not provide specific information for consideration. Most of the 
commenters that did not support the proposed listing were affiliated 
with the trophy hunting industry and opposed the rule due to potential 
impacts on importing trophies. These comments are addressed below.

Peer Review

    In accordance with our policy published on July 1, 1994 (59 FR 
34270), we solicited expert opinions from ten individuals with 
scientific expertise that included familiarity with the species, the 
geographic region in which wild members of the species occur, and 
conservation biology principles. We received responses from five of the 
peer reviewers from whom we requested comments. The peer reviewers 
generally supported our rule; however, they provided updated 
information on taxonomy, current population estimates, and population 
trends. They also found our analysis of some of the threats to be 
inaccurate. Specifically, they provided comments and additional 
information on loss of prey base, trophy hunting, infanticide, 
corruption, and trade in lion bones. In some cases, a correction is 
indicated in the citations by ``personal communication'' (pers. comm.), 
which could indicate either an email or telephone conversation; in 
other cases, the research citation is provided.

Peer Reviewer Comments

    (1) Comment: Several peer reviewers commented on our section of the 
proposed rule regarding the taxonomic classification of lion. These 
peer reviewers confirmed that the IUCN Cat Specialist Group recommended 
a two-subspecies classification: Panthera leo leo for lions of India 
and western and central Africa, and P. l. melanochaita for lions in 
eastern and southern Africa.
    Our Response: We have reviewed the 2015 IUCN Red List Assessment 
for the lion, which proposes the new classification as recommended by 
the IUCN Cat Specialist Group, and the genetic studies supporting this 
classification. We found this information to be the best available 
scientific and commercial information; therefore, we have accepted this 
taxonomic change and incorporated this decision into this document 
under the Taxonomy section of this document. As a result, our 
assessment is of the status of the lion species (both P. l. leo and P. 
l. melanochaita), including the lion population in India.
    (2) Comment: Several peer reviewers provided updated information on 
population estimates and trends. Based on a time trend analysis of 
scientific census data for 46 well-monitored populations, an overall 43 
percent decline in lion populations across Africa was inferred. 
Furthermore, regional trends emerged, showing that, while populations 
in southern African increased by 22 percent, populations in eastern and 
western and central Africa combined decreased by 57 percent and 66 
percent, respectively. The peer reviewers also indicated that the 
actual number of lions in Africa is much lower than previous estimates. 
Application of regional trends to lion estimates made in 2002 resulted 
in an estimate of fewer than 20,000 lions, a significant difference 
from the previous estimate of 32,000.
    Our Response: We considered this information and note that this 
information was also included in the IUCN Red List Assessment for the 
lion. Information on population estimates and trends was incorporated 
into the Species Information section of this document. Assessment of 
this information led us, in part, to conclude that the status of the 
lion is more serious than previously indicated, especially in the 
western and central regions of Africa (P. l. leo).
    (3) Comment: One peer reviewer commented that the section on prey 
loss does not address the issue of prey loss in protected areas where 
most lions occur.
    Our Response: The peer reviewer provided a list of literature on 
the patterns and trends of prey loss in protected areas that were 
recently or are currently occupied by lions. We have reviewed these 
articles and have incorporated the findings in this document (under 
Loss of Prey Base). This information did not change our determination, 
but rather further supported our determination that prey loss has 
occurred throughout the African range countries and is one of the major 
threats to lion.
    (4) Comment: One peer reviewer stated that although most lions in 
Africa persist inside protected areas, the majority of the protected 
areas should be uninhabited by humans; therefore,

[[Page 80048]]

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

[[Page 80049]]

drafting of the proposed rule and reviewed the citations provided 
during the public comment period. We have incorporated the new 
information accordingly.
    (16) Comment: One peer reviewer commented that our review of 
infanticide as a result of trophy hunting was incomplete and provided 
additional literature and citation on the subject for our 
consideration.
    Our Response: We agree that additional discussion was appropriate 
regarding the impacts of infanticide, including a review of the new 
studies provided on evolutionary adaptions and impacts of subadult 
early dispersal on the species. We agree that infanticide and 
associated factors relating to trophy hunting of males may have 
additive impacts on the decline of certain populations. Therefore, we 
have incorporated this information into our final rule.

Public Comments

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

[[Page 80050]]

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

[[Page 80051]]

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

[[Page 80052]]

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

[[Page 80053]]

trophy imports of P. l. melanochaita on a country-wide basis, the 
Service encourages the submission of information from individual 
applicants. We would rely on the information available to the Service 
and may rely on information from sources other than the applicant when 
making a permitting decision.
    (43) Comment: Two commenters stated the Service has offered nothing 
to demonstrate why limitations on the importation of sport-hunted 
African lions from throughout the subspecies' range is necessary and 
advisable to provide for the conservation of the subspecies or 
sufficient to overcome the Congressional conclusion that such imports 
would normally (i.e., presumptively) benefit the conservation of the 
species. Further, these commenters did not feel that the Service's 
proposed rule for African lion supported a conclusion that a 4(d) rule 
requiring import permits for trophies was necessary and advisable for 
the conservation of the subspecies.
    Our Response: For the import of sport-hunted trophies, while there 
is evidence that many of the range countries have lion management 
plans, we have little information indicating that the plans are being 
implemented, and we received new information during the public comment 
period indicating that some hunting programs are not scientifically 
based or providing adequate conservation benefits to the species. We 
want to encourage U.S. hunters to take advantage of one of the 
conservation tools available, well-regulated hunting programs, to 
improve the long-term survival of the subspecies. The 4(d) rule will 
support implementing well-managed plans by encouraging countries that 
have insufficient lion management plans to develop plans that are based 
on sound scientific information that would generate revenue in support 
of communities and conservation. As noted, the proposed 4(d) rule for 
African lion would provide for the importation into the United States 
of trophies taken legally in range countries upon the issuance of a 
threatened species import permit. While the Service cannot control 
hunting of foreign species such as African lion, we can regulate their 
importation and thereby require that U.S. imports of sport-hunted 
African lion trophy specimens are obtained in a manner that is 
consistent with the purposes of the Act and the conservation of the 
subspecies in the wild, by allowing importation from range countries 
that have management plans that are based on scientifically sound data 
and are being implemented to address the threats that are facing lions 
within that country.
    (44) Comment: Three commenters, a peer reviewer and comments from a 
consortium of seven range countries felt that the proposed 4(d) rule 
did not adequately explain the criteria used by the Service to 
determine whether the importation of any sport-hunted lion would 
enhance the survival of the species. The commenter expressed concern 
that because the Service has not adequately explained the criteria for 
enhancement or made an enhancement finding for lions in Africa, U.S. 
hunters will be barred from importing their lion trophy. The peer 
reviewer expressed a need for the Service to elaborate concrete 
requirements to which countries must adhere as a minimum standard in 
order for imports of sport-hunted lion trophies from a country to 
qualify for the export of lion trophies, including quotas of less than 
one male per 2000 km\2\ with a minimum age requirement.
    Our Response: We recognize that the preambular language of the 
proposed 4(d) rule was general, and we have addressed this issue in 
this final rule. However, we did not find that it was appropriate to 
establish specific criteria, such as a set quota number, in this final 
rule because this may not allow for the countries to implement an 
adaptive management strategy based on the current status of the species 
within the country. During the public comment period we received new 
information regarding infanticide and the effects of hunting younger 
male lions on pride structure. Therefore, we agree with the peer 
reviewer that the Service is in a position to proactively engage with 
countries to ensure exported trophies fulfill minimum age requirements 
and we will consider these factors in making our enhancement findings.
    (45) Comment: Two commenters recommended that the Service should 
not adopt a 4(d) rule until it makes specific enhancement-of-survival 
findings for each of the countries for which lions can be hunted, or 
delay the implementation of the 4(d) rule for 1 year. These two 
commenters, as well as a third commenter, stated that implementing the 
4(d) rule at this time would impact hunters who had already booked 
trophy hunts months or even years in advance, resulting in the loss of 
money invested that could not be recovered ``in the event of a sudden 
change in the rules governing the importation of sport-hunted 
trophies.''
    Our Response: In the proposed rule, the Service found that hunting, 
if well managed, may provide a benefit to the subspecies. However, the 
best available information, obtained by the Service during the public 
comment period, indicates that not all hunting programs are well 
managed or provide enhancement to survival of the subspecies. Delaying 
the implementation of a 4(d) rule may result in U.S. hunters 
participating in poorly managed hunting programs, which would be 
counter to the purposes of the Act. We do not agree that such a delay 
would be appropriate for the conservation of the subspecies. Regarding 
the potential loss of deposits for previously booked trophy hunts, 
hunters were notified of a potential regulatory change when the 
proposed rule with a 4(d) rule was published on October 29, 2014 (79 FR 
64472). The availability of the proposed rule would have given hunters 
the opportunity to use that information to minimize financial losses.
    (46) Comment: One commenter urged the Service to adjust the rule to 
ensure that imports are not stopped, and that the benefits generated by 
U.S. hunters in foreign countries continue while the Service is making 
determinations regarding the countries' lion management program. This 
commenter suggested that the Service issue U.S. import permits for all 
lion trophies until such time as the Service deems that the import from 
a particular country would not enhance the survival of the subspecies. 
It is the commenter's belief that there are beneficial aspects of 
hunting (benefits to local communities, dollars coming into the 
country, etc.) that should not be interrupted while the Service is 
making its determinations. The commenter expressed concern that the 
Service has insufficient resources to make timely country-by-country 
determinations.
    Our Response: Import of sport-hunted trophies of Panthera leo leo 
will require issuance of an endangered species import permit under 50 
CFR 17.22, which will require an enhancement finding. Import of sport-
hunted trophies of Panthera leo melanochaita will require issuance of a 
threatened species import permit under 50 CFR 17.32, which will require 
an enhancement finding (see 4(d) Rule for Panthera leo melanochaita, 
above). We would be unable to issue import permits until we made such 
determinations. The Service recognizes that making these findings may 
be time consuming given our current resources. We appreciate the 
commenter's willingness to use their own resources to obtain 
information on the range countries' management and assist the Service 
in making timely findings. We encourage the commenter and others to 
work with us by

[[Page 80054]]

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

[[Page 80055]]

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

[[Page 80056]]

lions, excessive harvests of lions for trophies, can negatively impact 
the reproduction of a lion such that it causes local extirpations. 
Harvesting males that are too young causes male replacements, which 
results in increased infanticide rates, death of the surviving male 
coalition, and a 100 percent fatality rate for males that are 
prematurely forced to disperse. Furthermore, the population will be 
driven to extinction as female populations collapse as they eventually 
are unable to mate. The species is largely not able to rapidly recover 
from population declines. This is evidenced by long-term population 
trends that indicate an overall 43 percent decline in lions over 21 
years and higher regional rates of decline in western and eastern 
Africa.
    (58) Comment: One commenter stated that the Service should use its 
power to list Distinct Population Segments (DPSs), rather than the 
entire African lion subspecies in light of the recent ruling in Humane 
Society of the United States v. Jewell, No. CV 13-186 (BAH), 2014 WL 
7237702 (D.D.C. Dec. 19, 2014)..
    Our Response: We disagree with this conclusion. Pursuant to 50 CFR 
17.11(g), all populations are included in the listing.

Required Determinations

National Environmental Policy Act (42 U.S.C. 4321 et seq.)

    We have determined that we do not need to prepare an environmental 
assessment, as defined under the authority of the National 
Environmental Policy Act of 1969, in connection with regulations 
adopted under section 4(a) of the Act for the listing, delisting, or 
reclassification of species. We published a notice outlining our 
reasons for this determination in the Federal Register on October 25, 
1983 (48 FR 49244).

References Cited

    A list of all references cited in this document is available at 
http://www.regulations.gov at Docket No. FWS-R9-ES-2012-0025, or upon 
request from the U.S. Fish and Wildlife Service, Endangered Species 
Program, Branch of Foreign Species (see FOR FURTHER INFORMATION 
CONTACT).

Authors

    The primary authors of this rule are staff of the Branch of Foreign 
Species, Ecological Services, U.S. Fish and Wildlife Service.

List of Subjects in 50 CFR Part 17

    Endangered and threatened species, Exports, Imports, Reporting and 
recordkeeping requirements, Transportation.

Regulation Promulgation

    Accordingly, we amend part 17, subchapter B of chapter I, title 50 
of the Code of Federal Regulations, as set forth below:

PART 17--[AMENDED]

0
1. The authority citation for part 17 continues to read as follows:

    Authority:  16 U.S.C. 1361-1407; 1531-1544; and 4201-4245; 
unless otherwise noted.


0
2. Amend Sec.  17.11(h), the List of Endangered and Threatened 
Wildlife, by:
0
a. Removing the entry for ``Lion, Asiatic (Panthera leo persica)''; and
0
b. Adding entries for ``Lion (Panthera leo leo)'' and ``Lion (Panthera 
leo melanochaita)'' in alphabetic order under Mammals to read as set 
forth below:


Sec.  17.11  Endangered and threatened wildlife.

* * * * *
    (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.
 
                                                                      * * * * * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------

* * * * *

0
3. Amend Sec.  17.40 by adding paragraph (r) to read as follows:


Sec.  17.40  Special rules--mammals.

* * * * *
    (r) Lion (Panthera leo melanochaita).
    (1) General requirements. All prohibitions and provisions of 
Sec. Sec.  17.31 and 17.32 apply to this subspecies.
    (2) The import exemption found in Sec.  17.8 for threatened 
wildlife listed in Appendix II of the Convention on International Trade 
in Endangered Species of Wild Fauna and Flora (CITES) does not apply to 
this subspecies. A threatened species import permit under Sec.  17.32 
is required for the importation of all specimens of Panthera leo 
melanochaita.
    (3) All applicable provisions of 50 CFR parts 13, 14, 17, and 23 
must be met.

    Dated: December 10, 2015.
Daniel M. Ashe,
Director, Fish and Wildlife Service.
[FR Doc. 2015-31958 Filed 12-21-15; 4:15 pm]
BILLING CODE 4333-15-P