Endangered and Threatened Wildlife and Plants; Listing Two Lion Subspecies

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.

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.

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).

ER23DE15.000

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

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

(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

, Côte 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

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]

Source

Western Africa (percent of total)

Central Africa (percent of total)

Eastern Africa (percent of total)

Southern

Africa (percent of total)

India

Total

Ferreras & Cousins 1996 (estimate for lion abundance in 1980)

75,800 (18,600 in 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 Riggio

et al.

2013)

1,640 (5 percent)

2,410 (7 percent)

17,290 (52 percent)

11,820 (37 percent)

33,160

Riggio 2013 (based on estimates of savanna habitat)

480 (1 percent)

2,419 (7 percent)

19,972 (57 percent)

12,036 (34 percent)

34,907

Henschel

et al.

2014

406 (n/a)

Bauer

et al.

2015a (trends applied to Bauer and Van Der Merwe 2004)

590 (3 percent)

7,345 (39 percent)

10,385 (55 percent)

18,726 *

Bauer

et al.

2015a (trends applied to Chardonnet 2002)

1,748 (6 percent)

13,316 (42 percent)

15,925 (51 percent)

31,394 *

Bauer

et al.

2015a

445

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; Brugiére

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 Comoé 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 Björklund 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].

Region

Estimated lions in sample

populations

1993

2014

Percent change

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%

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

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 Côte D'ivoire and Ghana Where Lions Are Considered Extirpated

[Population data is from UN 2013]

Subspecies

Country

UN Population estimate, in thousands

(people/km

2

)

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

(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

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; Bouché

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 (Bouché

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; Bouché

et al.

2010, pp. 99, 1001; Brashares

et al.

2004 in Craigie

et

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 (Bouché

et al.

2010, pp. 99, 1000; Roulet 2004 in Bouché

et al.

2010, p. 1002); Chad (Potgieter

et al.

2009 in Bouché

et al.

2010, p. 1002); Côte d'Ivoire (Fischer and Linsenmair 2001, p. 134); DRC (Martin and Hillman-Smith 1999 in Bouché

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 Bouché

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). Bouché

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).

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 (Brugiére

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;

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

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 (Brugiére

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 (Brugiére

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 (Brugiére

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

(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

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, Côte 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*

Country

Annual lion trophy quotas

Year(s) of data

Annual offtakes

Year(s) of data

Panthera leo leo

Benin

5.0±0

2007-2009

2.0±0.4

2007-2009

Burkina Faso

20.0±0

2006-2009

13.3±1.45

2006-2009

Cameroon

29.2±2

2006-2010

6.9±1.0

2006-2010

CAR

31

2009

13.7±6.9

2008-2011

Panthera leo melanochaita

Mozambique

42-60

2013

19.2±7.3

2008-2011

Namibia

14.5

2010

14.0±3.2

2008-2011

Tanzania

315

2011-2012

85

2011-2012

Zambia (moratorium)

1

74(50

2

)

2012

47

2012

Zimbabwe

101(50

3

)

2011

42.5±7.5

2008-2011

* Source: Lindsey

et al.

2013a. p.6.

1

Zambia enacted a moratorium on sport hunting in 19 game management units. Sport hunting remained open in other game management units and on some private game ranches. Sport hunting of all cats is currently banned throughout Zambia (White 2015, pers. comm.).

2

Approximate average quota for Zambia in the few years prior to the moratorium placed on cat hunting in 2013. (White 2015, pers. comm.).

3

In 2014, Zimbabwe reduced its quota to 50 due to implementation of age restrictions (Henschel 2015, pers. comm., citing Lindsey pers. comm.)

Potential Benefits of Trophy Hunting

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 Savé Valley

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 Savé 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 Savé 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 Savé 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

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

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

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 a

l. 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

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 count

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