64472 Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules 

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 the African Lion 

Subspecies as Threatened With a Rule 

Under Section 4(d) of the ESA 

AGENCY: Fish and Wildlife Service, 

Interior. 

ACTION: Proposed rule and 12-month 

finding. 

SUMMARY: We, the U.S. Fish and 

Wildlife Service (Service), announce a 

proposed rule and a 12-month finding 

on a petition to list the African lion 

(Panthera leo leo) as endangered under 

the Endangered Species Act of 1973, as 

amended (Act). After review of the best 

available scientific and commercial 

information, we find that listing the 

subspecies Panthera leo leo as 

threatened is warranted, and we 

propose to list the subspecies as 

threatened. We are also proposing a rule 

under section 4(d) of the Act to provide 

for conservation measures for the 

African lion. To ensure that subsequent 

rulemaking resulting from this proposed 

rule is as accurate and effective as 

possible, we are soliciting information 

from the scientific community; other 

governmental agencies, including those 

within the range of the African lion; 

nongovernmental organizations; the 

public; and any other interested parties. 

DATES: We will accept comments 

received or postmarked on or before 

January 27, 2015. We must receive 

requests for public hearings, in writing, 

at the address shown in FOR FURTHER

INFORMATION CONTACT by December 15, 

2014. 

ADDRESSES: You may submit comments 

by one of the following methods: 

(1) Electronically: Go to the Federal 

eRulemaking Portal: http://

www.regulations.gov. In the Search 

field, enter FWS–R9–ES–2012–0025, 

which is the docket number for this 

rulemaking. Then, click the Search 

button. You may submit a comment by 

clicking on ‘‘Comment Now!’’ 

(2) By hard copy: Submit by U.S. mail 

or hand-delivery to: Public Comments 

Processing, Attn: FWS–R9–ES–2012– 

0025, Division of Policy and Directives 

Management; U.S. Fish and Wildlife 

Service; MS: BPHC, 5275 Leesburg Pike, 

Falls Church, VA 22041–3803. 

FOR FURTHER INFORMATION CONTACT: 

Janine Van Norman, Chief, 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 

Under the Act, a species may warrant 

protection through listing if it is found 

to be an endangered or threatened 

species throughout all or a significant 

portion of its range. Under the Act, if a 

species is determined to be endangered 

or threatened we are required to publish 

in the Federal Register a proposed rule 

to list the species. The purpose of this 

proposed listing determination is to 

publish and seek comments on our 12- 

month finding on a petition to add the 

African lion to the list of threatened and 

endangered species. 

II. Major Provision of the Regulatory 

Action 

After review of the best available 

scientific and commercial information, 

we find that listing the African lion as 

threatened is warranted, and we 

announce a proposed rule to list the 

subspecies as threatened. We are also 

proposing a 4(d) rule to provide for 

conservation measures for the African 

lion. 

III. Costs and Benefits 

We have not analyzed the costs or 

benefits of this rulemaking action 

because the Act precludes consideration 

of such impacts on listing and delisting 

determinations. Instead, listing and 

delisting decisions are based solely on 

the best scientific and commercial 

information available regarding the 

status of the subject species. 

Information Requested 

Section 4(b)(1)(A) of the Act directs 

that determinations as to whether any 

species is an endangered or threatened 

species must be made solely on the 

basis of the best scientific and 

commercial data available. Therefore, 

we request comments or information 

from other concerned governmental 

agencies, the scientific community, 

industry, and any other interested 

parties concerning this proposed rule. 

We particularly seek comments 

concerning: 

(1) The subspecies’ biology, range, 

and population trends, including: 

(a) Genetics and taxonomy; 

(b) Historical and current range, 

including distribution; 

(c) Historical and current population 

levels; 

(d) Information pertaining to range 

countries’ regulatory mechanisms, 

including specific laws and regulations 

pertaining to loss of habitat, loss of prey 

base, and human-lion conflict. 

(e) Information pertaining to range 

countries’ management plans, including 

information on management and 

implementation of hunting concessions, 

conservation measures in place for this 

subspecies and its habitat, community 

education and outreach programs that 

address lion conservation, revenue 

gained from trophy hunting and how it 

is allocated, and any information 

pertaining to long-term conservation of 

lions and their habitat and prey base; 

and 

(f) Potential threats not already 

identified, such as extractive activities. 

(2) The factors that are the basis for 

making a listing determination for a 

species or subspecies under section 

4(a)(1) of the Act (16 U.S.C. 1531 et 

seq.), which are: 

(A) The present or threatened 

destruction, modification, or 

curtailment of its habitat or range; 

(B) Overutilization for commercial, 

recreational, scientific, or educational 

purposes; 

(C) Disease or predation; 

(D) The inadequacy of existing 

regulatory mechanisms; or 

(E) Other natural or manmade factors 

affecting its continued existence. 

(3) The potential effects of climate 

change on the subspecies and its 

habitat. 

Please include sufficient information 

with your submission (such as scientific 

journal articles or other publications) to 

allow us to verify any scientific or 

commercial information you include. 

Submissions merely stating support for 

or opposition to the action under 

consideration without providing 

supporting information, although noted, 

will not be considered in making a 

determination. 

We request that you send comments 

only by the methods described above in 

ADDRESSES. We will post all comments 

on http://www.regulations.gov. If you 

submit information via http://

www.regulations.gov, your entire 

submission—including any personal 

identifying information—will be posted 

on the Web site. If your submission is 

made via a hardcopy that includes 

personal identifying information, you 

may request at the top of your document 

that we withhold this information from 

public review. However, we cannot 

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Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules 64473 

guarantee that we will be able to do so. 

Please include sufficient information 

with your comments to allow us to 

verify any scientific or commercial 

information you include. 

Comments and materials we receive, 

as well as supporting documentation we 

used in preparing this proposed rule, 

will be available for public inspection 

on http://www.regulations.gov, or by 

appointment, during normal business 

hours, at the U.S. Fish and Wildlife 

Service, Ecological Services, Branch of 

Foreign Species (see FOR FURTHER

INFORMATION CONTACT). 

Public Hearing 

At this time, we do not have a public 

hearing scheduled for this proposed 

rule. The main purpose of most public 

hearings is to obtain public testimony or 

comment. In most cases, it is sufficient 

to submit comments through the Federal 

eRulemaking Portal, described above in 

ADDRESSES. If you would like to request 

a public hearing for this proposed rule, 

you must submit your request, in 

writing, to the person listed in FOR

FURTHER INFORMATION CONTACT by the 

date specified in DATES. 

Peer Review 

In accordance with our policy 

published on July 1, 1994 (59 FR 

34270), we will solicit the expert 

opinions of at least three appropriate 

and independent specialists for peer 

review of this proposed rule. The 

purpose of such review is to ensure that 

decisions are based on scientifically 

sound data, assumptions, and analysis. 

We will send peer reviewers copies of 

this proposed rule immediately 

following publication in the Federal 

Register. We will invite peer reviewers 

to comment, during the public comment 

period, on the specific assumptions and 

conclusions regarding the proposed 

listing status of threatened for the 

African lion subspecies. We will 

summarize the opinions of these 

reviewers in the final decision 

document, and we will consider their 

input and any additional information 

we receive, as part of our process of 

making a final decision on the proposal. 

Peer review is an important tool at our 

disposal to help evaluate the quality of 

the data and analyses we rely on in our 

decision making processes. The 1994 

peer review policy commits us to 

soliciting the expert opinions of 

‘‘appropriate and independent 

specialists regarding pertinent scientific 

or commercial data and assumptions 

relating to taxonomy . . . for species 

under consideration for listing.’’ The 

policy also requires that our final 

decision must document the opinions of 

all the independent peer reviewers, and 

that all information regarding peer 

review be included in the 

administrative record. All proposed 

listing rules must be peer reviewed 

according to this policy and to 

applicable standards under the Service’s 

guidelines for implementing the 

Information Quality Act and the 

December 15, 2004, Office of 

Management and Budget Final 

Information Quality Bulletin for Peer 

Review. 

Petition History and Previous Federal 

Action(s) 

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. This document consists of our 

proposed rule and our determination on 

the status review for the African lion 

and publishes our finding. Our status 

review may be obtained at http://

www.regulations.gov under Docket No. 

FWS–R9–ES–2012–0025. 

Conservation Status of the African Lion 

U.S. Endangered Species Act 

The African lion (Panthera leo leo) is 

currently not listed as either endangered 

or threatened under the Act, although 

the Asiatic lion (Panthera leo persica) 

has been listed as endangered since 

1970 under the Act and its precursor, 

the Endangered Species Conservation 

Act of 1969. 

International Union for the 

Conservation of Nature 

In 2008, the International Union for 

the Conservation of Nature (IUCN) 

classified the African lion as vulnerable 

with a declining population trend, 

which means the species is considered 

to be facing a high risk of extinction in 

the wild (Bauer et al. 2008, 

unpaginated). This classification is 

based on a suspected reduction in its 

population of approximately 30 percent 

over the previous two decades (Bauer et 

al. 2008, unpaginated). Because the 

regional lion population in western 

Africa is isolated and estimated to 

number well below the IUCN 

endangered criterion level of 2,500 

individuals, it is classified by the IUCN 

as regionally endangered (Bauer and 

Nowell 2004, entire). In the assessment 

for this classification, western Africa is 

defined as consisting of Benin, Burkina 

Faso, Cote d’Ivoire, Gambia (identified 

as ‘‘Regionally Extinct’’ (RE)), Ghana, 

Guinea, Guinea Bissau, Liberia (RE), 

Mali, Mauritania (RE), Niger, Nigeria, 

Senegal, Sierra Leone (RE), and Togo 

(Bauer and Nowell 2004, p. 35). 

Convention on International Trade in 

Endangered Species of Wild Fauna and 

Flora (CITES) 

The African lion is listed in Appendix 

II of the Convention on International 

Trade in Endangered Species of Wild 

Fauna and Flora (CITES). CITES (see 

http://www.cites.org) is an international 

agreement through which member 

countries work together to protect 

against over-exploitation of animal and 

plant species found in international 

trade. Parties regulate and monitor 

international trade in CITES-listed 

species—that is, their import, export, 

and reexport, and introduction from the 

sea—through a system of permits and 

certificates. CITES lists species in one of 

three appendices—Appendix I, II, or III. 

Species such as the African lion that are 

listed in Appendix II of CITES may be 

commercially traded, subject to several 

restrictions. CITES Appendix II includes 

species that are less vulnerable to 

extinction than species listed in 

Appendix I, and ‘‘although not 

necessarily now threatened with 

extinction, may become so unless trade 

in specimens of such species is subject 

to strict regulation in order to avoid 

utilization incompatible with their 

survival.’’ The status of the African lion 

with respect to CITES and how it is 

affected by international trade is 

discussed in more detail below, in the 

section titled Import/Export of Lion 

Parts and Products. 

Periodic Review Under CITES 

In an attempt to increase CITES 

protections for the African lion, in 2004, 

Kenya submitted a proposal for 

consideration at the Thirteenth Meeting 

of the Conference of the Parties to CITES 

(CoP13) to change the listing of the 

African lion from Appendix II of CITES 

to Appendix I (CoP13 Prop. 6; http://

www.cites.org/eng/cop/13/prop/E13- 

P06.pdf). An Appendix-I listing 

includes species threatened with 

extinction whose trade is permitted only 

under exceptional circumstances, which 

generally precludes commercial trade. 

The import of specimens (both live and 

dead, as well as parts and products) of 

an Appendix-I species generally 

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requires the issuance of both an import 

and export permit under CITES. Import 

permits are issued only if findings are 

made that the import would be for 

purposes that are not detrimental to the 

survival of the species in the wild and 

that the specimen will not be used for 

primarily commercial purposes. For live 

specimens, a finding must also be made 

that the recipient must be suitably 

equipped to house and care for the 

specimens (CITES Article III(3)). Export 

permits are issued only if findings are 

made that the specimen was legally 

acquired and the export is not 

detrimental to the survival of the 

species in the wild, and that a living 

specimen will be so prepared and 

shipped as to minimize the risk of 

injury, damage to health, or cruel 

treatment. (CITES Article III(2)). 

Although Kenya had submitted its 

proposal to CoP13 for consideration, it 

withdrew its proposal due to the lack of 

regional consensus on the proposal. 

Furthermore, plans were under way at 

that time for convening a regional 

workshop on lion management in 2005, 

the results of which would be reported 

to the CITES Animals Committee 

(Animals Committee) (http://

www.cites.org/eng/cop/13/rep/E13- 

ComIRep13.pdf). 

Recognizing that lion workshops and 

other research had been completed, 

producing updated information on the 

conservation and status of this species, 

the Animals Committee, at its 25th 

Meeting (AC25) (Geneva, Switzerland, 

July 2011), agreed to include the African 

lion in the Periodic Review of Felidae 

[Decision 13.93 (Rev. CoP15)] (http://

www.cites.org/eng/dec/valid15/E15- 

Dec.pdf) under the Animals Committee 

periodic review of the appendices. 

Kenya and Namibia offered to lead the 

review as a high priority with range 

country consultation (http://

www.cites.org/eng/com/ac/25/sum/E25- 

SumRec.pdf). At CoP16 in March 2013, 

the Parties adopted a revised Decision 

[Decision 13.93 (Rev. CoP16); http://

www.cites.org/common/cop/16/sum/ECoP16-Plen-06.pdf; http://

www.cites.org/eng/dec/valid16/13_93_

CoP16.php], directing the Animals 

Committee to complete its Review of the 

Appendices for Felidae and to provide 

a report at CoP17 on the result of the 

review of all Felidae. Kenya and 

Namibia recently submitted a report of 

their work on the Periodic Review of the 

African lion for discussion at the 27th 

Meeting of the Animals Committee 

(AC27, Veracruz, Mexico, 28 April–3 

May 2014) (CITES 2014a, entire). During 

discussion of this document at AC27, a 

representative of the IUCN informed the 

committee that the IUCN would be 

completing an updated Red List 

Assessment of the lion in 2015. In 

addition, she suggested potential 

nomenclature changes to lion 

subspecies (see Taxonomy). The 

Animals Committee took note of the 

upcoming Red List Assessment and 

requested Namibia and Kenya to 

incorporate this information into their 

Periodic Review and prepare a revised 

document for consideration at the 28th 

Meeting of the Animals Committee. 

Further, the Animals Committee made 

plans to continue seeking information 

from lion range states that had not yet 

responded to requests for information 

on the species. Finally, the Animals 

Committee took note of the recent 

information concerning changes in the 

nomenclature of lion subspecies and 

requested that the nomenclature expert 

of the Animals Committee review the 

information (CITES 2014b, p. 3). 

Regions in Which African Lions Occur 

The literature on African lion often 

includes reference to the following 

broad geographic regions: northern, 

western, central, southern, and eastern 

Africa. The boundaries of these regions 

vary somewhat among authors, based on 

the nature and result of the studies 

undertaken. 

As reflected in the literature reviewed 

for this proposed rule, the lion 

conservation community generally 

works in the context of the regions of 

Africa as they are described in Table 1. 

The regions as described in Table 1 may 

vary somewhat from the descriptions of 

the regions that may be found in 

taxonomic and other research literature. 

TABLE 1—DESCRIPTIONS OF THE DIFFERENT REGIONS OF AFRICA AS GENERALLY USED BY THE CONSERVATION

COMMUNITY

[Information derived from Chardonnet 2012, IUCN 2006a and IUCN 2006b] 

Regions Countries 

North of Saharan Desert: 

North Africa 1 ........................................................ Algeria 1, Egypt 1, Libya 1, Morocco 1, Tunisia.1

Sub-Saharan Africa: 

Western Africa ...................................................... Benin, Burkina Faso, Cote d’Ivoire 3, Gambia 1, Ghana 3, Guinea, Guinea-Bissau 3, Mali 3, 

Mauritania 1, Niger, Nigeria, Senegal, Sierra Leone 1, Togo.2 3

Central Africa ........................................................ Cameroon, CAR, Chad, Congo, DRC, Gabon, Sudan/South Sudan. 

Eastern Africa ....................................................... Burundi 2, Djibouti 1, Eritrea 1, Ethiopia, Kenya, Rwanda, Somalia, Sudan/South Sudan, 

Tanzania, Uganda. 

Southern Africa ..................................................... Angola, Botswana, Lesotho 1, Malawi, Mozambique, Namibia, South Africa, Swaziland, 

Zambia, Zimbabwe. 

1 Lions extirpated. 

2 Lions considered occasional or transient by Chardonnet 2002. 

3 Lions considered absent by Henschel et al. 2014. 

Species Description 

The lion is the second-largest extant 

cat species (second in size only to the 

tiger) and the largest carnivore in Africa. 

It has a broad geographical range, 

historically distributed throughout 

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 by 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 kg with the heaviest male 

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on record weighing 272 kg. Females are 

smaller, weighing, on average, 126 kg. The 

male body length, not including the tail, 

ranges from 1.7 m to 2.5 m with a tail from 

0.9 m to 1 m (Nowell & Jackson, 1996). 

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 Panthera leo is 

accepted within the scientific 

community, there is a lack of consensus 

regarding lion intraspecific taxonomy 

(Mazak 2010, p. 194; Barnett et. al. 

2006b, p. 2,120). 

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. 2,120), which is 

recognized by the Integrated Taxonomic 

Information System (ITIS) (ITIS 2013, 

www.itis.gov, accessed June 6, 2013). 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); 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. Results indicated that lions 

in India differed from lions in Africa, 

supporting a two-subspecies 

classification for extant lions: P. leo leo 

and P. leo persica, the African and 

Asian lion, respectively (Ellerman et al. 

1953, Meester and Setzer 1971, O’Brien 

et al. 1987, in Dubach 2005, p. 16). 

According to Dubach (2005, p. 16), most 

taxonomic authorities recognize this 

two-subspecies taxonomy. This 

taxonomy is also recognized by the 

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 represented by few samples. Results 

of analysis indicate that a major genetic 

subdivision among lions occurs in 

Africa, with lions in southern and 

eastern Africa being genetically distinct 

from and more genetically diverse than 

lions elsewhere (western and central 

western and central Africa and Asia). 

Evidence indicates that lions in western 

and central Africa (as well as nowextinct 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) 

contemporary range collapse and 

fragmentation is too recent a 

phenomenon to explain the lower 

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. 1,362– 

1,364). 

These genetic studies on lion 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 

have renewed debate on lion taxonomy 

among the experts. For this reason, the 

IUCN Species Survival Commission Cat 

Specialist Group has commissioned a 

Cat Classification Task Force from 

among its expert members to determine 

a consensus taxonomy for the group. 

Until then, we conclude that the 

taxonomy of the species is currently 

unresolved. However, as required by the 

Act, we base this status review on the 

best available scientific and commercial 

information, which is the most recent 

taxonomy that is the most widely 

recognized by taxonomic experts: P. leo 

leo (Africa) and P. leo persica (India). 

Consequently, in this document we 

review the status of the petitioned 

entity, the African lion, P. leo leo. 

Range 

Historically, lions occupied most of 

the African continent except the West 

African coastal rainforest zone, the 

Congo Basin rainforest zone, and the 

inner Sahara Desert (Bauer 2003, in Ray 

et al. 2005, p. 67; IUCN 2006a, p. 10; 

IUCN 2006b, p. 10). Ray et al. (2005, p. 

52) estimate lion historical range in 

Africa (at about 150 years prior to their 

study) to be roughly 22.2 million square 

kilometers (km2), while IUCN (2006a, p. 

12; 2006b, p. 13) estimates lion 

historical range in sub-Saharan Africa to 

be 19.3 million km2 (Table 2). 

Depending on the study and methods 

used, the species’ range is reported to 

currently cover between 3.0 million and 

5.0 million km2 (Table 2). The most 

recent range-wide study was based on a 

review of all of the most current 

available estimates of lion populations 

(up through 2012) (Riggio et al, p. 21), 

combined with satellite imagery of 

savannah habitat, and provided 

estimates of current lion range to be 3.4 

million km2 (Riggio et al. 2013, p. 26), 

or about 25 percent of the subspecies’ 

historic range in savannah habitat. 

According to Chardonnet (2002, pp. 24– 

25), about half the range of the African 

lion falls within protected areas. 

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1Current range includes occasional and probable 

range. 2Bauer et al (2008) provides a synthesis of the 

efforts from which the IUCN (2006a, b) estimates 

were generated, providing somewhat different 

numbers for southern and eastern Africa. Also, 

current range is range where lion occurrence is 

known, and in approximately 38 percent of 

historical range, the occurrence of lion is unknown. 3Riggio et al. (2013) calculate estimates for 

savannah habitat, defined as areas that receive 

between 300 and 1,500 mm of rain annually and 

which includes most of sub-Saharan Africa. 

The African lion is now believed to be 

extirpated from between 75 and 83 

percent of its former range (Table 2). 

The subspecies has been extirpated from 

all of its former range in northern Africa 

(Black et al. 2013, p. 1). In addition, 

according to IUCN (2006a,b; see Table 

2), the species’ range has declined by an 

estimated 91 percent in western Africa, 

79 percent in central Africa, and 68 

percent in eastern/southern Africa 

(Table 2), with lion occurrence 

unknown in an additional 38 percent of 

the historical range (Bauer et al 2008, p. 

16). More recently, Henschel et al. 

(2014, p. 5) estimate the confirmed lion 

range in western Africa, based on data 

collected between 2006 and 2012, to be 

49,000 km2, or an estimated 1.1 percent 

of the species’ former range in the 

region. 

TABLE 2—ESTIMATES OF THE AFRICAN LION RANGE

Source Region of Africa 

Historic 

range 

(km2) 

Current 

range 

(km2) 

Current range as 

percent of historic 

range 

(percent of historic 

range w/unknown 

lion presence) 

Ray et al. 2005: ........................................ Continent-wide .......................................... 22,200,000 3,800,000 17 percent. 

Chardonnet 2002: ..................................... Western .................................................... ........................ 121,980 

Central ...................................................... ........................ 651,970 

Eastern ..................................................... ........................ 1,137,205 

Southern ................................................... ........................ 1,039,212 

Total .......................................................... ........................ 2,950,367 

IUCN 2006a, b: 1 ....................................... Western .................................................... 3,814,576 331,749 9 percent. 

Central ...................................................... 3,392,241 715,482 21 percent. 

Western + Central .................................... 7,206,817 1,047,231 15 percent. 

Southern + Eastern .................................. 12,080,000 3,915,000 32 percent. 

Total .......................................................... 19,286,817 4,962,231 26 percent. 

Bauer et al. 2008: 1 thnsp;2 .................... Western + Central .................................... 7,206,817 1,047,231 15 percent. 

(0 percent). 

Southern + Eastern .................................. 13,010,000 3,564,000 23 percent. 

(58 percent). 

Total .......................................................... 20,216,817 4,611,231 22 percent. 

(38 percent). 

Riggio 2013 3 (based on estimates of savannah habitat):.Western ....................................................

Central ......................................................

........................ 133,784 

936,465 

Eastern ..................................................... ........................ 780,401 

Southern ................................................... ........................ 1,540,171 

Total .......................................................... 13,500,000 3,390,821 25 percent. 

Henschel et al. 2014: ................................ Western .................................................... ........................ 49,000 1 percent. 

The historical range of the African 

lion included most current continental 

African countries (Chardonnet 2002, pp. 

25–28). Currently, the subspecies occurs 

only in sub-Saharan Africa. Within this 

region, Chardonnet (2002, p. 27) 

described lions as present in 34 range 

states (35 with South Sudan, which 

gained its independence as a country in 

July 2011) and recently extirpated from 

6 range countries (Chardonnet 2002, p. 

27) (Table 1). The 34 sub-Saharan 

African range countries in which 

Chardonnet considered lions present 

included 10 in western Africa. More 

recently, during surveys of 21 large 

protected areas in western Africa, 

Henschel et al. (2014, p. 4) considered 

lions to be absent from protected areas 

in 5 of these 10 countries (Table 1). 

Distribution and Abundance 

The general distribution of lions in 

Africa is summarized by Ray et al. 

(2005, p. 67) as follows: 

Lions formerly occupied most of the 

African continent except for equatorial forest 

and the inner-Sahara. Today, they are extinct 

in North Africa and have undergone dramatic 

range retraction at the limits of their 

historical distribution. 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 (Bauer et al. 

2005, p. 6; Riggio et al. 2013, p. 31). 

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 

African lion population is an ambitious 

task, involving many uncertainties 

(IUCN 2012, p. 2). Estimates, 

particularly range-wide or broad regionwide 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 size of the African 

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4Estimates 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. 

(2013, p. 27). 

lion population have used different 

methods; 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 using a GIS-based model 

calibrated with information obtained 

from lion experts. Ferreras and Cousins 

predicted African lion abundance 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 3). 

Currently, about 90 percent of all 

African lions occur in southern and 

eastern Africa (Table 3). According to 

most studies, most African lions are in 

eastern Africa (Table 3). According to 

Riggio et al. (2013, p. 27), only nine 

countries contain resident populations 

of at least 1,000 free-ranging lions 

(Central African Republic, Kenya, 

Tanzania, Mozambique, Zambia, 

Zimbabwe, South Africa, Botswana, and 

possibly Angola). Approximately 40 

percent of all lions are found in 

Tanzania (Riggio et al. 2013, p. 27). 

Only about 10 percent of all lions occur 

in western and central Africa (Table 3). 

According to the most recent survey 

effort, numbers in western Africa are 

extremely low. Henschel et al. (2014, p. 

5) estimate that only 400 lions in the 

entire region, with most (about 350, or 

88 percent) concentrated in a single 

population. 

TABLE 3—ESTIMATES OF AFRICAN 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) 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 4 (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 

savannah 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).

In 2005–2006, in response to a 

growing concern that the African lion 

was in decline, IUCN and the Wildlife 

Conservation Society sponsored 

workshops to determine a lion 

conservation strategy. During these 

workshops, lion experts collectively 

assessed what they believed to be the 

then-current status of African lions 

based on a variety of information, 

including professional opinion. During 

the workshops, lion experts identified 

86 African lion Conservation Units 

(LCUs). They defined LCUs as areas of 

known, occasional, or possible lion 

range that can be considered an 

ecological unit of importance for lion 

conservation (IUCN 2006a, p. 14; IUCN 

2006b, p. 17). Of the 86 LCUs, 20 are in 

western and central Africa and 66 are in 

southern and eastern Africa (Table 4). 

Most (71 percent) have more than half 

their area under some form of legal 

protection (Bauer et al. 2008, p. 19). Few 

(16 percent) were estimated to contain 

large populations (Table 4). This was 

particularly the case for western and 

central Africa, where most (13, or 65 

percent) of LCUs were estimated to 

contain fewer than 50 lions (Table 4). 

The majority of those with large 

populations were in southern and 

eastern Africa (Table 4). Only 23 of 86 

LCUs (27 percent) were considered to 

contain viable populations, though more 

than half were thought to contain 

potentially viable populations (Table 4). 

Lion populations within 42 percent of 

the 86 LCUs were considered to be 

decreasing, whereas those in 9 percent 

were considered increasing. The 

remaining were considered stable or of 

unknown trend (Table 4). 

TABLE 4—LION CONSERVATION UNITS (LCUS) AS IDENTIFIED AND CHARACTERIZED IN IUCN 2006a AND IUCN 2006b 

Number of LCUs Western & 

Central Africa 

Eastern & 

Southern 

Africa 

All regions 

(percent) 

Total ................................................................................................................................. 20 66 86. 

Estimated to contain: 

>500 lions ................................................................................................................. 2 12 14 (16 percent). 

50–500 lions ............................................................................................................. 5 28 33 (38 percent). 

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TABLE 4—LION CONSERVATION UNITS (LCUS) AS IDENTIFIED AND CHARACTERIZED IN IUCN 2006a AND IUCN 2006b— 

Continued 

Number of LCUs Western & 

Central Africa 

Eastern & 

Southern 

Africa 

All regions 

(percent) 

<50 lions ................................................................................................................... 13 26 39 (45 percent). 

Considered: 

Viable ........................................................................................................................ 4 19 23 (27 percent). 

Potentially Viable ...................................................................................................... 12 34 46 (53 percent). 

Doubtful Viability ....................................................................................................... 4 13 17 (20 percent). 

With Populations Considered to be: 

Increasing ................................................................................................................. 3 5 8 (9 percent). 

Stable ........................................................................................................................ 5 21 26 (30 percent). 

Decreasing ................................................................................................................ 12 24 36 (42 percent). 

Unknown ................................................................................................................... 16 16 (19 percent). 

Riggio et al. (2013, entire) provide the 

most recent, most comprehensive 

estimates to date of free-ranging lion 

populations in Africa. They compiled 

all existing estimates of African lion 

populations since 2002, including data 

from Chardonnet (2002), Bauer and Van 

Der Merwe (2004), IUCN (2006a, 2006b), 

over 40 mainly country-specific reports, 

and their own experiences. They then 

combined these data with satellite 

imagery and information on habitat 

condition to estimate lion abundance 

and identify lion areas that they 

characterized as strongholds and 

potential strongholds. They conducted 

this within the context of savannah 

Africa, which they defined as areas that 

receive between 300 and 1,500 

millimeters (mm) of rain annually, and 

within which most of the present range 

of the African lion occurs. Also, they 

used the LCUs identified in the 2005– 

2006 lion workshops as the general 

framework within which to identify lion 

areas, strongholds, and potential 

strongholds. 

Riggio et al. (2013, p. 32) describe lion 

strongholds as areas meeting the 

necessary requirements for long-term 

viability; broadly, where management 

appears to be working. Potential 

strongholds are described, broadly, as 

areas where immediate interventions 

might create a viable population. 

Specifically defined, strongholds (1) 

contain at least 500 lions, (2) are within 

protected areas (including those that 

allow hunting), and (3) have stable or 

increasing lion numbers as assessed by 

IUCN (2006a, 2006b) (Riggio et al. 2013, 

p. 22). Potential strongholds contain at 

least 250 lions, but do not satisfy either 

requirement (2) or (3) above. The 

remaining lion areas—those not meeting 

the requirements of a stronghold or 

potential stronghold—are described as 

areas ‘‘where present management 

clearly isn’t working’’ (Riggio et al. 

2013, p. 32). Riggio et al. (2013, p. 32) 

derived the thresholds of 500 and 250 

using information in Bjo ̈rklund (2003) 

on the number of prides needed to avoid 

the risk of inbreeding in lion 

populations, and information in Bauer 

et al. (2008) on the average size of lion 

prides. Bjo ̈rklund (in Riggio et al. 2013, 

p. 32) assessed the risk of inbreeding 

due to habitat loss and determined that, 

‘‘. . . to sustain a large out-bred 

population of lions, a continuous 

population of at least 50 prides, but 

preferably 100 prides, with no limits to 

dispersal is required.’’ Bauer et al. 2008 

(in Riggio et al. 2013, p. 32) indicate the 

average lion pride as containing 

approximately five adults. 

The results of Riggio et al. indicate the 

size of the African lion population to be 

about 35,000, which falls within the 

range of the other recent estimates 

(Table 3). However, they state that 

‘‘Although these numbers are similar to 

previous estimates, they are 

geographically more comprehensive. 

There is abundant evidence of 

widespread declines and local 

extinctions’’ (Riggio et al. 2013, p. 18). 

Riggio et al. identified lions as 

occurring in 67 areas (Table 5). While a 

small portion (22 percent) of lion areas 

identified by Riggio et al. contain large 

populations, the majority are small and 

isolated (Riggio et al. 2013, p. 30; Table 

5). Most (69 percent) contain fewer than 

250 lions. A considerable portion (39 

percent) contains very small 

populations of fewer than 50 lions. 

These include 63 percent of the lion 

areas in western and central Africa, and 

31 percent of those in e/s Africa. 

TABLE 5—NUMBER OF LION AREAS AND NUMBER OF AREAS CONTAINING LION POPULATION CLASSES ACCORDING TO

RIGGIO ET AL. 2013 

Number of lion areas Western Central Eastern Southern All regions 

(percent) 

Total ......................................................................... 8 8 28 23 67. 

# Estimated to contain: 

≥500 lions ......................................................... 0 1 7 7 15 (22 percent). 

250–499 lions ................................................... 1 2 1 2 6 (9 percent). 

50–249 lions ..................................................... 0 2 12 6 20 (30 percent). 

<50 .................................................................... 7 3 8 8 26 (39 percent). 

Riggio et al. identify 10 lion 

strongholds (viable populations) and 7 

potential strongholds (Table 6). 

According to Riggio et al. (2013, p. 29), 

the 10 strongholds contain 

approximately 24,000 lions, or about 70 

percent of the current African lion 

population. Of those, most (about 

19,000 lions) are in protected areas. 

Potential strongholds contain about 

4,000 lions. More than 6,000 lions are 

located in areas not considered 

strongholds or potential strongholds and 

have a very high risk of being extirpated 

(Riggio et al. 2013, p. 33). 

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5Two lion areas in central Africa make up one 

potential stronghold. 

6Riggio et al. make one exception to the 

requirement that lion strongholds contain 

populations that are stable or increasing. IUCN 2006 

indicate lion numbers in the Tsavo/Mkomazi lion 

area are decreasing in numbers, but Riggio et al. 

believe that, while lion numbers are declining 

outside of protected areas, lions within the parks 

are usually well protected and in sufficient 

numbers to meet the criteria. 

TABLE 6—LION STRONGHOLDS AND POTENTIAL STRONGHOLDS IDENTIFIED BY RIGGIO ET AL. 2013 

Lion area Country Area 

(km2) Stronghold 

Lion 

population 

size 

Population 

size in 

protected 

areas 

IUCN 

(2006a, b) 

Trend 

Western Africa 

W-Arly-Pendjari .................... Benin, Burkina Faso, Niger .. 29,403 Potential ....... 350 350 Stable. 

Central Africa 

SE Chad ............................... Chad ..................................... 133,408 Potential 5 ..... 400 140 Stable. 

E CAR .................................. Central African Republic ...... 328,721 Potential 6 ..... 1,244 148 Stable. 

Eastern Africa 

Boma-Gambella ................... Ethiopia, South Sudan ......... 106,941 Potential ....... 500 ∼ 500 Unknown. 

Laikipia-Samburu ................. Kenya ................................... 35,511 Potential ....... 271 46 Stable. 

Tarangire .............................. Tanzania .............................. 28,771 Potential ....... 731 208 Decreasing. 

Ruaha-Rungwa .................... Tanzania .............................. 195,993 Stronghold .... 3,779 2,235 Stable. 

Selous .................................. Tanzania .............................. 138,035 Stronghold .... 7,644 4,953 Stable. 

Serengeti-Mara ..................... Kenya, Tanzania .................. 35,852 Stronghold .... 3,673 3,516 Increasing. 

Tsavo-Mkomazi .................... Kenya, Tanzania .................. 39,216 Stronghold .... 880 820 Decreasing. 

Southern Africa 

Etosha-Kunene ..................... Angola, Namibia ................... 123,800 Potential ....... 455 ∼ 315–595 Increasing. 

Kafue .................................... Zambia ................................. 58,898 Potential ....... 386 386 Stable. 

Great Limpopo ..................... Mozambique, South Africa, 

Zimbabwe.

150,347 Stronghold .... 2,311 2,179 Increasing. 

Kgalagadi ............................. Botswana, South Africa ....... 163,329 Stronghold .... 800 ∼ 800 Stable. 

Luangwa ............................... Malawi, Zambia .................... 72,992 Stronghold .... 574 574 Stable. 

Mid-Zambezi ......................... Mozambique, Zambia, 

Zimbabwe.

64,672 Stronghold .... 755 ∼ 350–650 Stable. 

Niassa .................................. Mozambique, Tanzania ........ 177,559 Stronghold .... 1,573 1,080 Increasing. 

Okavango-Hwange ............... Botswana, Zimbabwe ........... 99,552 Stronghold .... 2,300 ∼ 2,300 Stable. 

Most of the strongholds and potential 

strongholds identified by Riggio et al. 

are trans-boundary areas. The vast 

majority, including all 10 strongholds, 

are located in southern and eastern 

Africa. Of the 17 strongholds and 

potential strongholds, only two 

potential strongholds are located in 

western and central Africa, one each in 

western Africa and central Africa. Only 

a small portion of the lions in the 

central Africa potential stronghold are 

within protected areas. The western 

Africa potential stronghold has one of 

the smallest lion populations of the 17 

strongholds/potential strongholds and, 

according to Herschel et al. (2014, p. 5), 

contains 88–90 percent of all lions in 

the western Africa region. 

By definition, all 10 strongholds 

identified by Riggio et al. include 

protected areas. Packer et al. (2013a, 

entire; 2013b, entire) looked at the 

relationship between lion densities, 

population trends, management 

practices, and several other variables 

(human population densities, 

governance, sport hunting, private 

management, and reserve size) from 42 

sites in 11 countries in Africa. Results 

of modeling indicate that by 2050 about 

43 percent of lion populations in 

unfenced reserves may decline to less 

than 10 percent of the carrying 

capacities of the unfenced reserves, 

including those in Botswana, Kenya, 

Cameroon, Ghana, Tanzania, and 

Uganda. According to the same 

modeling results lion populations in 

fenced reserves are expected to remain 

at or above the carrying capacity of the 

fenced reserves for the next 100 years, 

although most are small protected areas 

with small lion populations (Creel et al. 

2013, entire). 

Trends 

Based on the best available 

information, as discussed above, African 

lion range and numbers have clearly 

declined over the past several decades. 

However, not all African lion 

populations have declined—some have 

increased or remained stable (see 

Distribution and Abundance), and some 

have been restored to areas from which 

they were previously extirpated (Packer 

et al. 2013, p. 636). Reports from the 

IUCN Species Survival Commission Cat 

Specialist Group (IUN 2006a, b) 

characterize the population as 

increasing in 3 of the lion strongholds 

identified by Riggio et al. (Table 6), as 

stable in 6 of the strongholds, and as 

decreasing in 1 stronghold. While four 

of the lion strongholds or potential 

strongholds identified by Riggio et al. 

(Table 6) are considered to be 

increasing, several African lion 

populations, containing a total of more 

than 6,000 individuals, have a very high 

risk of local extinction (Riggio et al. 

2013, p. 33). During the 2005–2006 

African lion workshops, lion experts 

characterized lion populations in 36 (42 

percent) of the 86 LCUs as decreasing. 

In extensive surveys recently conducted 

within 15 of the 20 LCUs in western and 

central Africa, Henschel et al. (2010, 

entire) were able to confirm lion 

presence in only four. The work of 

Packer et al. (2013) suggests future 

declines within a number of protected 

areas. Craigie et al. (2010, entire) 

provide evidence of declining large 

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mammal populations in Africa’s 

protected areas, indicating that 

protected areas in Africa have generally 

failed to mitigate threats to large 

mammal populations, including African 

lion. Although Craigie et al. (2010, p. 

2,225) found large regional differences 

(from large declines in western Africa to 

positive rates of change in southern 

Africa), they found overall populations 

decreased steadily from 1970 to 2005. 

Biology/Ecology 

Habitat 

Historically, the species occurred in 

all habitats in Africa, except rainforest 

and the hyper-arid interior of the Sahara 

(Ray et al. 2006, p. 66). Today they are 

found primarily in savannah, 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 savannah woodlands 

plains mosaics of eastern and southern 

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). 

General Biology 

Lions are well studied. Much 

information exists on African lion 

habits, behavior, and ecology. CITES 

(2014a, 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 ‘‘fissionfusion’’ 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 weeks 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 four 

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). 

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 

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. 

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. 

2013a, p. 636; Hayward et al. 2007, 

entire), and range from 0.08–0.13 adults 

and subadults per 100 km2 in Selous 

Game Reserve up to 18 per km2 in 

protected areas of eastern Africa and 

South Africa (Creel and Creel 1997, 

Nowell and Jackson 1996, in Haas et al. 

2005, p. 4). Aside from human-related 

mortality, prey availability is likely the 

primary determinant of lion density 

(Fuller & 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 

African lion prides are 26–226 km2, 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 km2 (Packer et 

al. 2013, p. 636). Funston (2011, p. 5) 

found the home ranges of lion prides in 

the dune-savannah habitat of Kgalagadi 

Transfrontier Park to range from 1,762 

to 4,532 km2. 

Because lion home ranges can be very 

large, many protected areas are not large 

enough to sustain them (Winterbach et 

al. 2014, p. 1; Funston 2011, p. 1, citing 

several sources). Where lion ranges 

approximate protected area size, lions 

roam near or beyond the protected area 

boundary, increasing human-lion 

contact and human-caused lion 

mortality. In these situations, local or 

regional extirpation probability is high 

due to the population sink created 

around the boundary of the protected 

area (Davidson et al. 2011, in 

Winterbach et al. 2012, p. 5; Funston 

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2011, p. 1, citing several sources; 

Brashares et al. 2001, entire). This ‘‘edge 

effect’’ is a major threat to carnivore 

populations inside protected areas 

throughout the world (Woodroffe 2001, 

in Winterbach et al. 2012, p. 5) (also see 

Human-Lion Conflict). 

Habitat Loss 

Habitat loss and degradation is 

reported to be among the main threats 

to African 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 agriculture and 

intensive livestock grazing in lion 

habitat (IUCN 2006a, p. 18; IUCN 2006b, 

p. 23; Ray et al. 2005, pp. 68–69; 

Chardonnet 2002, pp. 103–106). 

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). 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. Lions are sensitive 

to loss of cover or prey. Riggio et al. 

(2013, p. 18) state that dense human 

populations and widespread conversion 

of land to human use preclude use by 

lions. 

Habitat destruction and degradation 

has been extensive throughout the range 

of the African lion, resulting in local 

and regional lion population 

extirpations, reduced lion densities, a 

dramatically reduced subspecies range 

(see Range), and small, fragmented, and 

isolated lion populations that are 

increasingly limited to protected areas 

(see Distribution and Abundance) (Ray 

et al. 2005, p. 69; Bauer and Van der 

Merwe 2004, pp. 29–30; Nowell and 

Jackson 1996, pp. 20–21). Lions appear 

to have one of the lowest levels of 

ecological resilience to human-caused 

habitat fragmentation; they are the least 

successful large African carnivore 

outside conservation areas (Woodroffe 

2001, in Winterbach et al. 2012, p. 6). 

Large carnivores with low ecological 

resilience have a high risk of local 

extinction. In order to survive, they 

require larger contiguous habitats with 

lower 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). 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), there was a 57 percent increase 

in agriculture area (from just over 200 

million ha to almost 340 million ha) and 

21 percent decrease in natural 

vegetation in the region (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). 

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. (2013, 

entire) found that lions in unfenced 

reserves are highly sensitive to human 

population densities in surrounding 

communities. 

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 km2

served as a proxy for the extent of landuse 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 km2 as the 

mean human density at which lions 

went locally or regionally extinct. Riggio 

et al. (2013, p. 29) estimate that there 

were originally approximately 13.5 

million km2 of savannah habitat in 

Africa. In 1960, 11.9 million km2 of 

these habitats had fewer than 25 people 

per km2, and in 2000 this number 

decreased to 9.7 million km2. Based on 

analysis of land-use conversion using 

satellite imagery and human population 

densities, Riggio et al. (2013, p. 29) 

found current savannah habitat that is 

suitable for lions to be fragmented and 

to total about 3.4 million km2 (or 25 

percent of African savannah habitat). 

These data suggest a substantial 

decrease in lion habitat over the past 50 

years. 

Projections of future human 

population growth, area of conversion to 

agriculture, and livestock numbers in 

Africa suggest suitable lion habitat will 

continue to decrease into the foreseeable 

future. Africa has the fastest population 

growth rate in the world (UNEP 2012a, 

p. 2). Future population growth in subSaharan Africa is projected to be large 

and rapid (UN 2013, p. 9). Although 

urbanization is increasing in subSaharan Africa (UN 2014, p. 20), the 

majority of the population is rural, and 

about 60–70 percent of the 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 subSaharan 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), Alexandratos and 

Bruinsma (2012, p. 107) project the area 

of cultivated land to increase by 51 

million ha (approximately 21 percent). 

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, 

agriculture, and/or livestock grazing are 

reported as occurring in or on the 

periphery of several of the 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 

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some parts of southern Africa. There are 

only two potential strongholds in 

western and central Africa (one in each 

region). Expansion of agriculture and 

livestock grazing are reported in or 

around both (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), and management of 

protected areas in portions of both is 

reported as weak (Heschel et al. 2014, 

pp. 5–6; IUCN 2008, p. 8). Eastern 

Africa contains over half of all the lions 

in Africa (Table 3). Seven of the 

seventeen African lion strongholds and 

potential strongholds identified by 

Riggio et al. occur in eastern Africa, and 

six of those seven (all four strongholds 

and two of three potential strongholds) 

are located in Tanzania and Kenya 

(Table 6). 

Between 1990 and 2010, Kenya’s 

human population grew from 23 million 

(40/km2) to 41 million (70/km2), 

whereas Tanzania’s grew from 25 

million (27/km2) to 45 million (48/km2) 

(UN 2013, pp. 421, 798). Not 

unexpectedly, sources indicate that 

expansion of agriculture and livestock 

grazing is occurring in these countries 

(Brink et al. 2014, entire; UNEP 2009, p. 

91; Mesochina et al. 2010, p. 74), 

including in or around lion strongholds 

and potential strongholds (Ogutu et al. 

2011, entire; Mesochina et al. 2010, 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. (2010, 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. In Kenya, 

the Kenya Wildlife Service (2009, p. 21) 

indicates that habitat loss due to landuse changes and human encroachment 

into previously wild areas is having a 

major impact on lion range size. By 

2050 the UN projects the human 

population of Tanzania to almost triple 

its 2010 population, reaching a density 

of 137 people per km2, whereas Kenya’s 

population is projected to more than 

double, reaching a density of 167 people 

per km2 (Table 7). 

The human populations of most other 

current and recent lion range countries 

are also expected to have very high 

growth rates (Table 7). It is important to 

note that the country-wide human 

population densities provided here (and 

in Table 7) 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 is 

particularly the case given that much of 

sub-Saharan Africa is rural and locals 

depend on agriculture for their 

livelihood. 

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. Human population 

growth, and resulting pressures exerted 

on habitat, are also expected to vary 

widely in the 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 348 people 

per km 2 (Malawi) (Table 7). 

Summary of Habitat Loss 

In the past several decades the human 

population has been expanding with 

concomitant large decreases in lion 

habitat and lion populations, resulting 

in an extremely large reduction in the 

species’ range. Habitat for African lion 

continues to be threatened with 

destruction, modification, and 

curtailment. Human populations are 

projected to increase dramatically in 

sub-Saharan Africa in coming decades. 

As human populations continue to rise 

in sub-Saharan Africa, the amount of 

land required to meet the expanding 

human population’s needs is constantly 

increasing. In addition, as indicated 

above, lions are increasingly limited to 

protected areas, and human population 

growth rates around protected areas in 

Africa tend to be higher than the average 

rural growth rate (Wittemyer et al. 2008, 

entire). Considering the majority of the 

human population in sub-Saharan 

Africa is rural, and land supports the 

livelihood of most of the population, 

loss and degradation of lion habitat can 

be expected to accompany the rapid 

growth in sub-Saharan Africa’s human 

population. Therefore, overall, because 

(1) lion prides have vast ranges and the 

subspecies requires large areas of 

suitable habitat to survive, (2) the 

subspecies’ range has already declined 

dramatically and is increasingly limited 

to protected areas, and (3) habitat loss 

and degradation is occurring in or 

around several of the remaining lion 

strongholds (viable populations) and 

potential strongholds, we conclude 

based on the best available scientific 

and commercial information that the 

continued destruction, modification, 

and curtailment of lion habitat is likely 

to become a significant threat to the 

African lion throughout its range. 

Human-Lion Conflict 

Human-lion conflict and associated 

retaliatory killing of lions has played a 

major role in the reduction of lion 

populations (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 the greatest threat to remaining lion 

populations (Hazzah et al. 2009, p. 

2,428; 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; Begg 

and Begg 2010, p. 2; Hazzah et al. 2009, 

p. 2,428; Moghari 2009, p. 36; Kissui 

2008, p. 422; Hazzah 2006, pp. 15, 23, 

25). 

Human-wildlife conflict stems from 

human population growth and the 

resulting overlap of humans and 

wildlife habitat (Chardonnet et al. 2010, 

p. 6; 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. However, despite being 

within protected areas, lions continue to 

be impacted by people living on 

adjacent land. Villages are established 

on the borders of protected areas, 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 (Hazzah et al. 2013, p. 1; 

Republic of Namibia 2013, p. 13; 

Chardonnet et al. 2010, pp. 11–12; 

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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). The most significant cause 

of human-lion conflict is livestock 

depredation. 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). 

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 appears to be more 

frequent in southern and eastern Africa 

(Chardonnet et al. 2010, pp. 12, 13; 

Mesochina et al. 2010a, pp. 29–30; 

Frank et al. 2006, pp. 1, 10). Lion attacks 

can have various impacts on those 

communities that coexist with conflictcausing animals, generating resentment 

towards them. When lions 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). 

Loss of Prey Base 

The lion’s prey base has decreased in 

many parts of its range for various 

reasons, but a large factor is due to 

competition for meat by humans. 

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. Historically, subsistence hunting 

with spears was traditionally used to 

hunt wildlife, which had minimal 

impact to wildlife populations. Spears 

have since been replaced by automatic 

weaponry (Chardonnet et al. 2010, 

p. 27), allowing for poaching of large 

numbers of animals for the bushmeat 

trade. 

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 comprises between 

6 percent (southern Africa) and 55 

percent (Central African Republic) of a 

human’s diet within the African lion’s 

range (Chardonnet et al. 2005, p. 9; 

IUCN 2006b, p. 19). In addition, 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). This 

growing demand and widely available 

modern weapons has led to increased 

poaching of native wildlife (Chardonnet 

et al. 2010, pp. 13–14, 27; Packer et al. 

2010, p. 8). Because many wildlife 

species are being hunted at 

unsustainable levels to meet this 

demand within the range of the lion, its 

prey base is becoming depleted in many 

areas, which has led lions to seek out 

livestock (and in some cases, humans) 

for food (Hoppe-Dominik et al. 2011, p. 

452; Chardonnet et al. 2010, pp. 6, 13– 

14; Frank et al. 2006, p. 12). 

Further, the demand for agriculture to 

meet the increasing needs of a growing 

population has been met by intensified 

agricultural and livestock practices 

(Chardonnet et al. 2010, p. 19). As 

natural habitats are converted to 

agricultural or pastoral land, it removes 

the food and cover needed by wildlife, 

and the lion’s natural prey base is 

reduced, causing them to prey on 

domestic livestock (Chardonnet et al. 

2010, p. 27; Gebresenbet et al. 2009, 

p. 9). 

In Tanzania, which is home to more 

than 40 percent of the African lion 

population, conversion of rangeland to 

agricultural use has blocked several 

migratory routes for wildebeest and 

zebra populations, both lion prey 

species, which likely forces lions to rely 

more on livestock (Packer et al. 2010, p. 

9). Conditions worsen as livestock 

numbers and area under cultivation 

increase, leading to overgrazing, further 

habitat destruction, and greater 

depredation rates by lions (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). Additionally, the use of fences to 

subdivide group ranches 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 

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). 

Studies have shown variation in rates 

of livestock depredation 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 

further for forage and water, they use 

temporary bomas (a livestock enclosure) 

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 

(Chardonnet et al. 2010, p. 24; Frank et 

al. 2006, p. 6). In 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 

(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, 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 (Packer et al. 

2010, p. 9; Kissui 2008, p. 427; Patterson 

et al. 2004, p. 514; Frank et al. 2006, p. 

6). Similarly, 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 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 

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(Mesochina et al. 2010a, p. 39; Moghari 

2009, p. 85; Frank et al. 2006, p. 12). 

Attacks on Livestock 

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, these 

traditional practices are being replaced 

by less diligent husbandry practices, 

which are 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). 

Attacks on Humans 

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). Inebriated 

people may walk in an altered manner 

that resembles sick or injured prey, 

attracting the attention of lions (Moghari 

2009, p. 85). 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 

Competition with humans, habitat 

changes, and regional climate variations 

can decrease availability of prey and 

increase human-lion conflict. When 

native prey are unavailable or difficult 

to find and kill, lions will target 

domestic livestock or humans 

(Chardonnet et al. 2010, p. 27; Moghari 

2009, pp. 78, 83; Hazzah 2006, pp. 17– 

18; Patterson et al. 2004, pp. 507, 514). 

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). 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 that do have 

an economic value to people, they are 

subject to retaliatory killing. This greatly 

impacts already-dwindling lion 

populations (Chardonnet et al. 2010, pp. 

12–14; Mesochina et al. 2010a, p. 38; 

Mesochina et al. 2010b, p. 32; 

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). 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. 2,429; 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 

(Government 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. 2,429; 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). Studies have shown 

that lion populations are declining in 

areas where pastoralism persists 

(Hazzah et al. 2009, p. 2,428). 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 km2, yet it 

may be that just under half of the lions 

that are killed are reported (Republic of 

Namibia 2013, p. 14). Although most of 

the information on human-lion conflict 

comes from just a few areas of the lion’s 

range (e.g., Kenya, Tanzania, and 

Uganda), it is reasonable to conclude 

that lions are being killed due to conflict 

in all major range countries, due to their 

depredation on livestock (Frank et al. 

2006, p. 4). 

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). 

Factors That Drive Retaliation 

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 

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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. 2,428; 

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). 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. 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. 2,436; Maclennan 

et al. 2009 in Hazzah et al. 2009, p. 

2,429; 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). 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. The prohibition 

of ritual lion hunts provides a greater 

incentive for participating in retaliatory 

hunts (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). 

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). 

Summary of Human-Lion Conflict 

Human-lion conflict and associated 

retaliatory killing of lions has played a 

major role in the reduction of lion 

populations and is the greatest threat to 

remaining lion populations. The most 

significant cause of human-lion conflict 

is livestock depredation and, to a lesser 

extent, attacks on humans. Expansion of 

human settlements and agricultural and 

pastoral activities into lion habitat, and 

even into protected areas, decreases 

prey availability and increases exposure 

of livestock and humans to lions. 

The most common solution to lion 

attacks is retaliatory killing. Spearing, 

shooting, trapping, and poisoning of 

lions occur regularly. Although a 

majority of information on human-lion 

conflict comes from a few areas of the 

lion’s range, we can reasonably 

conclude that lions are being killed due 

to conflict in all major range countries, 

because of their depredation on 

livestock (Frank et al. 2006, p. 4). 

Impacts on victims of lion attacks 

create resentment towards lions and 

lion conservation, and a greater 

likelihood of retaliation. Even when 

lions are not the predators responsible 

for the majority of attacks, lions incite 

a greater response and are killed more 

often than other predators of livestock. 

In areas of high human density and 

low lion density, mainly in smaller 

reserves and outside large protected 

areas, lion populations may not be 

sustainable. Attacks on humans can 

impact long-term viability for lions as 

people who fear for their lives or safety 

are unlikely to support conservation 

actions and are more likely to retaliate 

by killing any lions found near 

settlements (Frank et al. 2006, p. 12). 

Every year, human-lion conflicts 

intensify due to habitat loss, poor 

livestock management, and decreased 

availability of wild prey, further 

increasing the likelihood that the 

subspecies will be at risk of extinction 

within the foreseeable future (Lion 

Guardians 2013, p. 1). 

Human population growth within the 

lion’s range is projected to be 2.1 billion 

by 2050 (UN 2012, p. 2). The number of 

livestock within the lion’s range is 

projected to increase by about 73 

percent by 2050 (Food and Agriculture 

Organization of the United Nations 

2012, p. 133). Given this expected 

increase in humans and livestock by 

2050, we conclude the conditions 

described above will continue to worsen 

to the point that African lions will likely 

be at risk of extinction within the 

foreseeable future. As livestock numbers 

increase, expansion of agricultural and 

pastoral practices continue, and the 

lion’s prey base is hunted at 

unsustainable levels to meet a growing 

demand for food, livestock depredation 

and retributive killing of lions will 

likely increase (Dickman 2013, p. 379; 

Hoppe-Dominik et al. 2011, p. 452; 

Chardonnet et al. 2010, p. 19; 

Gebresenbet et al. 2009, p. 9; Hazzah 

and Dolrenry 2007, p. 3). Furthermore, 

as the need for grazing land becomes 

more critical, expansion of livestock 

numbers may be partially supported by 

the network of protected areas, seen by 

herders as unused pastures (Chardonnet 

et al. 2010, p. 25). 

Retaliatory killing of lions continue in 

many areas and this practice impacts 

the viability of lion populations 

throughout its range. The killing of lions 

due to human-lion conflict is enough to 

result in the local extirpation of lion 

populations, though at present does not 

place the subspecies in danger of 

extinction. Human-lion conflict is 

exacerbated by an increasing human 

population, the expansion of human 

settlements, loss of prey base due to the 

bushmeat trade and expanding 

agriculture, as well as increasing 

pressures on natural resources to meet 

the needs of the growing human 

population. We expect retaliatory 

killings due to human-lion conflict to 

continue to increase into the foreseeable 

future. We conclude based on the best 

available scientific and commercial 

information that the continuation of this 

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activity is a significant threat to the 

African lion throughout its range. 

Disease 

Wild lions are known to be infected 

with various pathogens (Hunter et al. 

2012, p. 2; Craft 2008, p. 6; Michel et al. 

2006, p. 92; Hofmann-Lehmann et al. 

1996, pp. 559–561). The human 

population within the range of the lion 

is expanding into lion habitat, 

increasing the exposure of lions to 

diseases from domestic animals (IUCN 

2006b, p. 26). Because lions are a top 

predator, they are at a particularly high 

risk of exposure to pathogens (Keet et al. 

2009, p. 11). Some pathogens are 

endemic, meaning they are constantly 

present, but often do not cause disease. 

Others are epidemic and cause a sudden 

severe outbreak with the potential to 

cause high mortality (Craft 2008, pp. 5, 

6). Although lions are known to be 

infected with certain pathogens, 

information on the extent of the 

subspecies’ infections and impacts of 

these diseases on lion populations is 

limited, because few long-term studies 

have been conducted; for example, 

those lion populations found in 

Serengeti National Park, Ngorongoro 

Crater, and Kruger National Park. 

Feline calicivirus, feline herpesvirus, 

feline parvovirus, feline coronavirus, 

and feline leukemia virus are endemic 

viruses known to occur in lions of 

Serengeti National Park, Ngorongoro 

Crater, Lake Manyara National Park, 

Kruger National Park, and Etosha 

National Park (but not all viruses are 

known in all parks). However, these 

diseases are not known to affect lion 

survival (Hunter et al. 2012, p. 2; Craft 

2008, p. 6; Hofmann-Lehmann 1996, pp. 

559, 561). 

Lions within Kruger National Park 

and Hluhluwe-iMfolozi Park, South 

Africa, and Serengeti National Park, 

Tanzania, are known to be infected with 

Mycobacterium bovis, a pathogen that 

causes bovine tuberculosis (bTB). This 

pathogen is not endemic to African 

wildlife and was likely introduced from 

cattle imported from Europe. M. bovis is 

transmitted to ungulates, such as 

African buffalo (Syncerus caffer) and 

wildebeest (Connochaetes taurinus) 

from domestic cattle located on the 

periphery of the parks (Maas et al. 2012, 

p. 4,206; Keet et al. 2009, pp. 4, 11; 

Renwick et al. 2007, p. 532; Michel et 

al. 2006, pp. 92, 93; Cleaveland et al. 

2005, pp. 446, 449, 450). Spillover of the 

disease from buffalo to other lion prey 

species, such as kudu (Tragelaphus 

strepsiceros) and warthog 

(Phacochoerus africanus), have also 

been documented (Keet et al. 2009, pp. 

4, 11; Renwick et al. 2007, p. 535; 

Cleaveland et al. 2005, p. 450). Because 

the lion’s primary prey are infected with 

bTB, they are frequently exposed to 

large amounts of infected tissue and are 

at risk of infection (Keet et al. 2009, pp. 

4, 6; Renwick et al. 2007, pp. 532, 536; 

Michel et al. 2006, p. 93; Cleaveland et 

al. 2005, pp. 450, 451). Furthermore, 

predators prey on weak animals and 

scavenge on carcasses, increasing their 

likelihood of being exposed to M. bovis 

(Renwick et al. 2007, p. 536; Michel et 

al. 2006, p. 93). Transmission may also 

occur among lions via scratching and 

biting (Keet et al. 2009, p. 7; Renwick 

et al. 2007, pp. 532–533). M. bovis is a 

pathogen that causes the infected 

animal to remain infectious and, 

therefore, a source of infection, until it 

dies (Renwick et al. 2007, p. 531). 

The social behavior of buffalo and 

lions allows M. bovis to spread to larger 

areas and facilitates the transmission 

within and between prides. Drought 

conditions may also encourage the 

spread of this pathogen as herds must 

move into new areas in search of forage, 

potentially putting them in contact with 

new, uninfected herds (Keet et al. 2009, 

pp. 4, 6; Renwick et al. 2007, p. 533; 

Michel et al. 2006, p. 93). In Kruger 

National Park, bTB was introduced in 

the southeastern corner of the park 

between 1950 and 1960. It gradually 

made a northern progress and reached 

the park’s northern boundary in 2006. In 

2009, the disease was found in buffalo 

across the river boundary in Zimbabwe 

(Keet et al. 2009, pp. 6, 11; Renwick et 

al. 2007, pp. 532, 533; Michel et al. 

2006, pp. 92, 96, 98). In time it will 

likely spread to Mozambique (Keet et al. 

2009, p. 6). In Serengeti National Park, 

infection may be widespread due to the 

large, migratory wildebeest population 

that ranges throughout the Serengeti 

ecosystem, including Maasai Mara 

National Reserve (Cleaveland et al. 

2005, p. 450). Although an eradication 

program has been implemented for 

cattle in South Africa, once an infection 

is established in a free-ranging 

maintenance host, like buffalo, it is 

unlikely to be eradicated (Keet et al. 

2009, p. 11; Renwick et al. 2007, pp. 

537, 538; Michel et al. 2006, p. 96). In 

fact, modeling has predicted that 

prevalence could reach as high as 90 

percent over the next 25 years, with 

similar consequences for predators 

(Renwick et al. 2007, p. 535). 

Clinical signs of bTB in lions include: 

emaciation, respiratory complications, 

swollen lymph nodes, draining sinuses, 

ataxia, and lameness (Keet et al. 2009, 

p. 13; Renwick et al. 2007, pp. 533, 534; 

Cleaveland et al. 2005, p. 450), although 

some lions may be subclinically 

infected but remain asymptomatic until 

they experience another bTB infection, 

suffer from poor nutrition or advancing 

age, or become super-infected with 

other diseases that may exacerbate the 

infection (Renwick et al. 2007, p. 533). 

The impact of bTB on lions is largely 

unknown. Researchers suggest that bTB 

may lower breeding success, reduce 

resiliency, and may be a mortality factor 

based on data that indicate survival is 

shortened in infected lions, with death 

ranging between 2 and 5 years after 

infection (Maas et al. 2012, p. 4,212; 

Renwick et al. 2007, p. 536; Michel et 

al. 2006, p. 93; Cleaveland et al. 2005, 

pp. 450, 451). Thirty percent of the 

inbred populations in HluhluweiMfolozi Park died due to a combination 

of bTB and malnutrition (Hunter et al. 

2012, p. 3). A study from Kruger 

National Park indicated that bTB 

spreads quickly through lion 

populations; in an area with high herd 

prevalence of M. bovis, 90 percent of 

lions became infected (Cleaveland et al. 

2005, p. 451). However, despite bTB 

infection and a high prevalence in prey 

species, the lion population in Kruger 

National Park has remained stable 

(Ferreira and Funston 2010, p. 201). 

Epidemics of canine distemper virus 

(CDV) are known to have occurred in 

the Serengeti-Mara Ecosystem, an area 

that encompasses the Serengeti National 

Park, Ngorongoro Conservation Area, 

and Maasai Mara National Reserve 

(Craft 2008, pp. 13–14; Cleaveland et al. 

2007, pp. 613, 616, 618). CDV is a 

common pathogen in the large 

population of domestic dogs around the 

Serengeti-Mara Ecosystem, which are 

believed to be the source of CDV 

(Cleaveland et al. 2007, pp. 613, 617). 

CDV is assumed to be transferred to 

lions by the sharing of food sources with 

spotted hyenas (Crocuta crocuta) or 

jackals (Canis spp.) that become 

infected by consuming the infected 

carcasses of domestic dogs (Canis lupus 

familiaris). Lions may also transmit 

CDV among themselves via sharing 

food, fights, and mating (Craft et al. 

2009, pp. 1,778, 1,783; Craft 2008, pp. 

13, 18, 71). 

CDV generally lacks clinical signs or 

measurable mortality in lions, and most 

CDV events have been harmless. 

However, in 1994 and 2001, CDV 

epidemics in the Serengeti National 

Park/Maasai Mara National Reserve and 

Ngorongoro Crater, respectively, 

resulted in unusually high mortality 

rates (Hunter et al. 2012, p. 2; Craft 

2008, p. 14; Munson et al. 2008, pp. 1, 

2; Cleaveland et al. 2007, pp. 613, 618; 

Roelke-Parker et al. 1996, pp. 441, 443). 

These outbreaks coincided with climate 

extremes that resulted in a higher 

number of Babesia, a tick-borne 

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parasite, infections (Munson et al. 2008, 

pp. 2, 5). Babesia is common in lions, 

but typically at low levels with no 

measurable impacts on their health 

(Craft 2008, p. 14; Munson et al. 2008, 

p. 3). However, droughts in 1993 and 

2000 in Serengeti National Park/Maasai 

Mara National Reserve and Ngorongoro 

Crater, respectively, led to large-scale 

starvation and widespread die-offs of 

buffalo. This situation combined with 

resumption of rains and fire suppression 

in Ngorongoro Crater favored 

propagation of ticks, vectors of Babesia, 

leading to unusually high tick burdens. 

The compromised health of buffalo 

allowed lions to feed on an inordinate 

number of tick-infested prey (Craft 2008, 

p. 14; Munson et al. 2008, pp. 2, 4, 5). 

Exposure to either CDV or Babesia 

singly is not typically associated with a 

compromise in health or an increase in 

mortality (Craft 2008, p. 14; Munson et 

al. 2008, pp. 1, 2, 3). However, the 

Babesia infections were exacerbated by 

the immunosuppressive effects of CDV 

and led to the unusually high mortality 

rates (Craft 2008, p. 14; Munson et al. 

2008, p. 5). The Serengeti National Park/ 

Maasai Mara National Reserve lion 

population lost 30 percent of its 

population (approximately 1,000 lions), 

but has recovered to its pre-epidemic 

population levels (Craft 2008, pp. v, 14, 

41; Munson et al. 2008, p. 1; Cleaveland 

et al. 2007, pp. 613, 617; Roelke-Parker 

et al. 1996, p. 444). Thirty-four percent 

of the Ngorongoro Crater lion 

population was killed, but frequent 

outbreaks of disease have prevented this 

population from recovering back to its 

carrying capacity (Craft 2008, p. 14; 

Munson et al. 2008, pp. 1, 2; Cleaveland 

et al. 2007, p. 617). The difference in 

recovery is likely due to the highly 

inbred nature of the Ngorongoro Crater 

lion population, compared to the 

Serengeti population, and its greater 

susceptibility to parasitic and viral 

infections (Hunter et al. 2012, p. 2; 

Munson et al. 2008, p. 5; Brown et al. 

1994, pp. 5,953–5,954). 

Feline immunodeficiency virus (FIV) 

is an endemic pathogen in many lion 

populations of southern and eastern 

Africa (Maas et al. 2012, p. 4,206; 

Adams et al. 2011, p. 173; PeconSlattery et al. 2008, p. 2; HofmannLehmann et al. 1996, pp. 555, 558; 

Brown et al. 1994, p. 5,966). FIV is 

believed to have been present in lions 

since the late Pliocene (O’Brien et al. 

2012, p. 243; Troyer et al. 2011, p. 2; 

Roelke et al. 2009, p. 3; Pecon-Slattery 

et al. 2008, p. 8). There are 6 subtypes 

of FIV, A through F, each with a distinct 

geographic area of endemnicity (Adams 

et al. 2011, p. 174; Troyer et al. 2011, 

p. 2; Roelke et al. 2009, p. 3; PeconSlattery et al. 2008, p. 4; O’Brien et al. 

2006, p. 262). The social nature of lions 

allows for viral transmission within and 

between prides through saliva when 

biting (Maas et al. 2012, p. 4210; PeconSlattery et al. 2008, p. 5; Brown et al. 

1994, p. 5,953). Prevalence of FIV in 

infected lion populations is high, often 

approaching 100 percent of adults 

(O’Brien et al. 2012, p. 243; Troyer et al. 

2011, p. 2; Roelke et al. 2009, p. 3; 

O’Brien et al. 2006, p. 262; HofmannLehmann et al. 1996, p. 559). 

FIV causes immune deficiencies that 

allow for opportunistic infections in the 

host (Brown et al. 1994, p. 5,953). 

Chronic effects of FIV are important to 

long-term survival and differ according 

to subtype (Troyer et al. 2011, p. 6). 

Studies have indicated that lions may 

exhibit signs of opportunistic infection 

associated with AIDS, such as swollen 

lymph nodes, gingivitis, tongue 

papillomas, dehydration, poor coat 

condition, and abnormal red blood cell 

parameters, and in some cases death 

(Troyer et al. 2011, p. 2; Roelke et al. 

2009, pp. 2, 3–6). Lions in Botswana 

and Tanzania have demonstrated 

multiple clinical features of chronic 

immune depletion similar to HIV and 

domestic cat AIDS (Troyer et al. 2011, 

pp. 2–3). However, there is no evidence 

that it poses a threat to wild populations 

(Frank et al. 2006, p. 1); FIV does not 

appear to be impacting lions in Kruger 

National Park (Maas et al. 2012, p. 

4,212), and no evidence of AIDS-like 

illnesses or decreased lifespan has been 

found in FIV lion populations in the 

Serengeti (O’Brien et al. 2006, p. 263). 

Infection with a single disease does 

not appear to have detrimental impacts 

on lions, although general body 

condition, health, and lifespan may be 

compromised. Co-infections, however, 

could have synergistic effects that lead 

to greater impacts on lions than a single 

infection. Lions impacted by the 1994 

CDV outbreak in Serengeti National 

Park/Maasai Mara National Reserve may 

have been more susceptible to CDV due 

to depleted immunity caused by FIV 

(O’Brien et al. 2006, p. 263). Troyer et 

al. (2011, pp. 5–6) found that survival 

during the CDV/Babesia outbreak in 

Serengeti National Park/Maasai Mara 

National Reserve was significantly less 

for lions infected with FIV A and/or C 

than FIV B. This finding suggests that 

FIV A and C may predispose carriers to 

CDV pathogenesis and may increase the 

risk of mortality (O’Brien et al. 2012, p. 

243). Additionally, certain 

environmental conditions may 

exacerbate the effects of an otherwise 

innocuous infection. For example, as 

discussed above, CDV and Babesia 

infections generally have no measurable 

impacts on lion health, but climatic 

conditions increased exposure of lions 

to Babesia infections, which were 

exacerbated by the immunosuppressive 

effects of CDV and led to unusually high 

mortality rates. Furthermore, species 

with reduced genetic variation may be 

less able to mount an effective immune 

response against an emerging pathogen 

(O’Brien et al. 2006, p. 255). Some lions 

infected with bTB may remain 

asymptomatic until conditions change 

and they suffer from poor nutrition due 

to low prey density, advancing age, or 

become super-infected with other 

diseases that may exacerbate the 

infection (Renwick et al. 2007, p. 533). 

Impacts of coinfections of FIV with 

FCV, FPV, FHV, and FCoV on 

individual lions are negligible and do 

not endanger the lion population, at 

least in the absence of other aggravating 

cofactors (Hofmann-Lehmann et al. 

1996, p. 561). Pathogen–pathogen 

interactions may become more 

important when lions are under 

additional stress (e.g., increased parasite 

load or low prey density) (Maas et al. 

2012, p. 4,212). 

Although disease is known in several 

populations, the impacts are known in 

only a couple of populations where 

disease has been frequently studied. 

Disease can be a factor in the decline of 

lions when combined with other factors, 

including environmental changes, 

reduced prey density, and inbreeding 

depression. However, this type of 

impact has been observed in some small 

populations that are at a higher risk, but 

has not been observed at the species 

population level. Therefore, we 

conclude, based on the best scientific 

and commercial information available, 

that disease is not a significant threat to 

the species. 

Deleterious Effects Due to Small 

Population Sizes 

The risk of extinction is related to the 

moment when a declining population 

becomes a small population and is often 

estimated using minimum viable 

population (MVP) sizes (Traill et al. 

2010, p. 28). The viability of a lion 

population is complex, but it partly 

depends on the number of prides and 

ability of males to disperse and interact 

with other prides, which affects 

exchange of genetic material (Bjorklund 

2003, p. 518). Without genetic exchange, 

or variation, individual fitness is 

reduced and species are less able to 

adapt to environmental changes and 

stress, increasing the risk of extinction 

(Bijlsma and Loeschcke 2012, pp. 117, 

119; Segelbacher et al. 2010, p. 2; Traill 

et al. 2010, p. 31; Bjorklund 2003, p. 

515). 

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7We found conflicting data on Cameroon, which 

was reported to prohibit trophy hunting (CITES 

2014, p. 14), although other information provided 

by Lindsey (2013, pers. comm.) and Jackson (2013, 

p. 8) state that trophy hunting is legal in Cameroon. 

Some scientists believe that the 

minimum viable population size (MVP) 

to maintain genetic viability is between 

500 and 5,000 individuals, although this 

estimate is not specific to lion (Bijlsma 

and Loeschcke 2012, p. 122; Traill et al. 

2010, p. 30; Willi et al. 2006, p. 449). 

The MVP for the African lion has not 

been formally established and agreed 

upon by species experts (Riggio et al. 

2011, p. 5; CITES 2004, p. 2; Bjorkland 

2003, p. 521); however, it has been 

suggested that, to conserve genetic 

diversity populations of 50 to 100 prides 

(250 to 500 individuals), with no limits 

to dispersal, are necessary because 

inbreeding increases significantly when 

populations fall below 10 prides. If 

there are less than 10 prides, inbreeding 

will increase from an F-value of 0.0 in 

the initial state to an F-value 0.26–0.45 

after 30 generations, while if the number 

of prides is 100 this F-value is only 

around 0.05 assuming no migration into 

the population (Bjorkland 2003, p. 515). 

F is the probability that the two alleles 

of a gene in an individual are identical 

by descent. Therefore, the Service 

considers the MVP to be 50 prides. 

Because the number of prides and male 

dispersal are the most important factors 

for maintaining viability, sufficient 

areas are needed to support 50 or more 

prides and allow unrestricted male 

dispersal. Unfortunately, few lion 

populations meet these criteria, and few 

protected areas are large enough to 

support viable populations (Bauer et al. 

2008, unpaginated; Riggio 2011, p. 5; 

Hazzah 2006, p. 2; Bauer and Van Der 

Merwe 2004, pp. 28–30; Bjorklund 

2003, p. 521). Even within large areas, 

inbreeding will increase if dispersal is 

limited, (Bjorklund 2003, pp. 521–522). 

More than 6,000 lions are in 

populations where their probability of 

survival is likely to be at risk of 

extinction within the foreseeable future 

(Riggio et al. 2013, p. 33). Furthermore, 

research indicates that there is a general 

lack of gene flow in most lion 

conservation units (Dubach et al. 2013, 

pp. 749, 750; Bertola et al. 2011, p. 

1364; Chardonnet et al. 2009, p. 54). 

Small populations (e.g. fewer than 50 

lions) can persist in the wild for some 

time; however, the lack of dispersal and 

genetic variation can negatively impact 

the reproductive fitness of lions in these 

populations and local extirpation is 

likely (Traill et al. 2010, p. 30; O’Brien 

1994, p. 5,748). 

Increasing human population growth 

between now and 2050 will continue to 

decrease and fragment large areas of 

habitat needed to support viable lion 

populations and disrupt dispersal routes 

for genetic exchange. Additionally, as 

the human population grows and lion 

populations decline, as discussed above, 

more lion populations could reach 

levels below the suggested minimum of 

10 prides to maintain genetic diversity, 

putting more populations at risk of 

inbreeding and extirpation. Therefore, 

we conclude, based on the best 

scientific and commercial information 

available, that small population sizes 

currently pose a threat to the species. 

Trophy Hunting 

Trophy hunting (also known as sport 

hunting) has been identified by the 

petitioners as one of the factors 

contributing to the decline of African 

lions (Petition 2011, p. 24). Lions are a 

key species in sport hunting as they are 

considered one of the ‘‘big five’’ (lion, 

leopard, elephant, rhino, and cape 

buffalo), touted to be the most 

challenging species 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, the sport hunting of 

lions for trophies has become a highly 

complex issue that has raised 

considerable controversy among 

stakeholders. 

Range Countries 

As of May 2014, approximately 18 

countries in Africa permit lions to be 

hunted for trophies: Benin, Burkina 

Faso, Central African Republic (CAR), 

Democratic Republic of Congo (DRC), 

Ethiopia, Ivory Coast, Mali, 

Mozambique, Namibia, Senegal, 

Somalia, South Africa (RSA), Sudan, 

Tanzania, Togo, Uganda, Zambia, and 

Zimbabwe. However, in 2013 lion 

trophy hunting was only documented to 

occur in nine countries, specifically 

Benin, Burkina Faso, CAR, 

Mozambique, Namibia, RSA, Tanzania, 

Zambia, and Zimbabwe (Lindsey 2013, 

personal communication). Four 

countries, Burundi, Guinea Bissau, 

Lesotho, and Swaziland, provide no 

legal protection for lions (CITES 2014a, 

p. 14). 

Hunting Moratoriums 

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. 

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. 

Therefore, you will see us refer to 

countries like Zambia and Botswana, 

each of which has hunting quotas and 

bans in place. Trophy hunting is 

currently banned in 12 countries: 

Angola, Botswana, Cameroon,7 Congo, 

Gabon, Ghana, Kenya, Malawi, 

Mauritania, Niger, Nigeria, and Rwanda 

(CITES 2014a, p.14; Lindsey et al. 

2013a, entire; Lindsey 2013, pers. 

comm.; Jackson 2013, pp. 7–8). 

Botswana banned lion hunting between 

2001 and 2004, and then again from 

2008 to the present (Davison et al. 2011, 

p. 114). Kenya banned all sport hunting 

in 1977 (African Wildlife Foundation 

1998, p. 3). Trophy hunting is restricted 

to problem or dangerous animals in 

Ethiopia and Uganda (Lindsey 2008, p. 

42). Zambia banned all sport hunting in 

January of 2013; while restrictions were 

lifted from other trophy species in 

August 2014, the ban on lions and 

leopards remains in place (ABC News 

2014, unpaginated; Flocken 2013, 

unpaginated). In 2011, researchers in 

Cameroon suggested that there should 

be an immediate moratorium of at least 

5 years on the hunting of lions in 

Cameroon, during which lions are 

allowed to recover and a management 

plan for lion hunting is established 

(Croes et al. 2011). 

Quotas 

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). For a quota to be 

scientifically based, it must be based 

upon available monitoring data of the 

species. Although varying by country 

and by economic resources, monitoring 

data used to determine quotas have 

included, but are not limited to, past 

hunting off-take records, trophy quality 

data, ground transect surveys, wildlife 

ranger and safari operator input, the 

species’ reproductive biology, and aerial 

population census data, although 

usually aerial data is limited to species 

that can be easily observed from the air, 

such as elephants and buffalo (Barnett & 

Patterson 2005, p. 102). Generally, the 

conservation principle behind 

scientifically based quotas is to limit 

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offtake of the species to either equal or 

slightly lower than the growth rate of 

the target specimens (e.g., males vs. 

female), provided the offtake does not 

damage the integrity and sustainability 

of that population. 

In order for scientifically based quotas 

to result in offtake less than the growth 

rate of target specimens, many factors 

are evaluated including the species’ 

biological factors (reproductive rate, 

gender, age, and behavior), as well as 

community and client objectives (WWF 

1997, pp. 14–19). Each quota should be 

then assigned to a geographical area 

and/or population based on this 

information. Thus, for lions, a 

scientifically based quota defines the 

specific number of lions that can be 

removed from a specific geographical 

area and population, for any purpose, 

within a particular year. Scientifically 

based quotas do not apply solely to 

sport hunting, but set the limits for all 

offtake for a particular year; other 

potential offtake includes problemanimal control (to reduce humanwildlife conflict), translocation (to 

expand conservation), culling (reducing 

population pressures), and local hunting 

(for protein/meat or employment) (WWF 

1997, pp. 8–10). 

While each of these uses offers 

advantages and disadvantages, quotas 

are typically utilized only for sport 

hunting, as it may provide the highest 

all-around benefits to local 

communities. For example, a portion of 

a quota could be used to kill a problem 

animal; the benefits to the community 

would then include the use of the 

animal parts for meat or trade and it 

would theoretically reduce the conflict. 

However, this provides a more limited 

economic benefit to the community than 

would selling the same quota for trophy 

hunting, which could potentially 

eliminate the problem animal, provide 

meat and parts for trade, and provide 

revenue for the community (WWF 1997, 

pp. 31–33). 

There are two primary types of 

quotas, ‘‘fixed’’ and ‘‘optional.’’ Trophy 

fees for ‘‘optional’’ quotas are paid only 

when the lion is shot, whereas, ‘‘fixed’’ 

quotas require the payment of a portion 

(40–100 percent) of the lion trophy fee, 

regardless of whether the hunt is 

successful. Until 1999, male lions were 

typically on ‘‘fixed’’ quotas, whereas 

female lions were under ‘‘optional’’ 

quotas. Due to this approach, trophies 

collected in the 1990’s were often of 

lower quality, younger, less desirable 

male lions, as operators and hunters had 

no incentive to be selective (e.g. the 

hunter had already paid for it). 

Therefore, current recommendation for 

all quotas is to be the ‘‘optional’’ type 

(Lindsey et al. 2013a, p. 9; Packer et al. 

2006, pp. 5, 9). 

Two primary concerns have been 

raised by the scientific and international 

community with regards to current lion 

quotas. Specifically, that existing quotas 

are set above sustainable levels and the 

data used for setting quotas is 

inconsistent and not scientifically based 

(Hunter et al. 2013, unpaginated; 

Lindsey et al. 2006, p. 284). For 

example, recent quotas appear rarely to 

address safeguards for sustainability or 

establish a systematic approach to 

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 humanlion conflict levels (Lindsey et al. 

2013b, p. 4). Apparently, in recognition 

of these inconsistencies, 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). We 

have no information on whether these 

best practices have been implemented 

by the lion range states. However, most 

countries that allow trophy hunting of 

lions appear to be reviewing their 

trophy hunting practices (Jackson 2013, 

pp. 2–3; White 2013, pp. 12–13). Benin 

halved their quotas in 2002 after the 

first population census of lions was 

conducted and resulted in the current 

quota of six lions every 2 years in 

Pendjari and four lions every 2 years in 

western Benin or one lion annually in 

each of the five hunting zones. This was 

largely due to impacts to lions from 

habitat degradation and fragmentation 

(particularly exacerbated by the increase 

of human population), loss of prey by 

poaching, trade (both legal and illegal), 

and human-lion conflict. (CITES 2014a, 

p. 5; Sogbohossou 2014, p. 1). 

Throughout the countries in Africa, 

most appear to have reduced their 

offtake considerably since the 1990’s. 

According to Packer et al. (2006, pp. 2– 

3), regardless of population estimates, 

countries are allowing for only a small 

proportion of their lion populations to 

be hunted, with most countries ranging 

from 2–4 percent annually (excluding 

offtake from South Africa, where offtake 

has been increasing from the trophy 

hunting of primarily captive-born lions, 

and Zimbabwe, where offtake was 2–3 

percent higher than other countries from 

1998–2004. 

Regardless of these reductions, many 

stakeholders consider the quota system 

to be outdated and ineffective because it 

does not address the biological and 

social impacts of trophy hunting on lion 

prides. Opponents also state that trophy 

hunting affects the social structure of 

the pride and results in increased 

infanticide of lion cubs. This 

supposition is inconclusive and not 

well supported (CITES 2014a, p. 14; 

Dagg 2000, pp. 831–835) (See 

Infanticide and Age-based Hunting 

Strategies). Regardless, since 2006, 

researchers have recommended the 

implementation of age-based hunting 

strategies; these are discussed below 

(Packer et al. 2006, pp. 6–8). 

Five countries maintain quotas to 

allow for approximately 6–15 lion 

trophies to be taken per year: Benin, 

Burkina Faso, Cameroon,7 Mozambique, 

and Namibia. Tanzania allows the take 

of approximately 50 lions annually, and 

Zimbabwe allows approximately 70 

animals annually to be taken (Jackson 

2013 pp. 7–8, CITES WCMC–UNEP 

trade database, accessed December 

2013). In Ethiopia and Uganda, trophy 

hunting is restricted to problem or 

dangerous animals only (Lindsey 2008, 

p. 42), and Botswana and Zambia 

currently ban all trophy hunting (CITES 

2014a, p.14). 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). 

Below is a summary of estimated 

annual hunting quotas for the African 

lion: 

TABLE 7—ANNUAL TROPHY QUOTAS

(APPROXIMATE) AS OF 2013 

Country 

Annual lion 

trophy quotas 

(Jackson 2013, 

pp. 7–8) 

Benin ................................... 6 

Botswana (moratorium) ...... 30 

Burkina Faso ...................... 6 

Cameroon 7 ......................... 6 

Mozambique ....................... 15 

Namibia ............................... 10 

Tanzania (as of 2012) ........ 50 

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TABLE 7—ANNUAL TROPHY QUOTAS

(APPROXIMATE) AS OF 2013—Continued 

Country 

Annual lion 

trophy quotas 

(Jackson 2013, 

pp. 7–8) 

Zambia (moratorium) .......... 50 

Zimbabwe ........................... 70 

Import/Export of Lion Trophies 

Although each country has its own 

method of regulating trophy hunting, 

international trade of lion trophies must 

adhere to CITES (see Conservation 

Status). International trade of lion parts 

and products (including trophies) are 

reported by both the exporting and 

importing countries and tracked by the 

United Nations Environment 

Programme World Conservation 

Monitoring Centre (UNEP–WCMC). The 

international trade data on the African 

lion that has been compiled in the 

CITES UNEP–WCMC Trade Database is 

extensive. Therefore, it is likely that the 

actual numbers of African lion parts and 

products in international trade is 

slightly smaller than what we have 

reported using the UNEP–WCMC ‘‘gross 

exports’’ report (CITES lion gross 

exports, http://trade.cites.org, accessed 

April 23, 2014). 

In 2012, the most recent year for 

which CITES trade data are available, 

U.S. CITES Annual Report trade data 

indicated that the United States allowed 

the direct import of African lion 

trophies from eight African countries, as 

follows: 

Central African Republic = 1 trophy 

Ethiopia = 1 trophy 

Mozambique = 5 trophies 

Namibia = 5 trophies 

South Africa = 413 trophies (the majority of 

which are reported to be of captive-born 

origin) 

Tanzania = 42 trophies 

Zambia = 32 trophies 

Zimbabwe = 49 trophies 

According to the CITES UNEP– 

WCMC database, between 2005 and 

2012, exports of lion trophies have 

demonstrated a decreasing trend when 

exports of captive-born lions from South 

Africa are excluded (CITES lion gross 

exports, http://trade.cites.org, accessed 

April 23, 2014). For example, in 2005 

there were 874 lion trophy exports 

reported in UNEP–WCMC, 521 if South 

Africa were excluded; whereas in 2012, 

there were 1,237 lion trophy exports 

reported in UNEP–WCMC, 336 if South 

Africa is excluded. 

Here it should be noted that there are 

limitations to interpreting the above 

reported information. The 2004 guide to 

using the CITES Trade Database 

indicates that the outputs produced by 

the CITES Trade Database can be easily 

misinterpreted if one is not familiar 

with it (CITES 2004b, p. 5). The number 

of ‘‘trophies’’ reported does not 

necessarily equate to the number of 

lions hunted. Additionally, the number 

of trophies reported for a given year in 

the trade report does not equate directly 

to the number of animals hunted in that 

given year (CITES export permits may 

be valid for 6 months, and a trophy 

could in theory be exported the year 

after it was hunted). The second 

limitation to interpreting this 

information is, although many permits 

may indicate that an animal is of wild 

origin (source code ‘‘W’’), these permits 

may be incorrectly coded. This is true 

for South Africa, where during the 

period of 2000 to 2009, animals that 

were captive-born and released into 

private reserve systems were assigned 

an incorrect source code of ‘‘wild.’’ 

South Africa has since requested their 

provincial authorities to use the correct 

source code for ‘‘captive bred’’ in order 

to correctly reflect the source of sporthunted lion trophies; however, some 

provinces are still not complying (RSA 

2013, pp. 8–9). However, based on 

South African trade data, the bulk of the 

exports of lions and their parts and 

products (including trophies) from 

South Africa were from captive-born 

lions (RSA 2013, p. 7). 

Tanzania, with the highest lion 

populations (Hamunyela et al. 2013, pp. 

29, 283; Riggio et al. 2013, p. 32; Ikanda 

2008, p. 4; Baldus 2004, pp. 5, 6), was 

the largest exporter of wild-origin lion 

trophies, but their exports have 

decreased significantly since 2006. In 

2008, approximately 138 lions had been 

estimated to be killed in Tanzania as 

trophies. In 2010, Tanzania’s numbers 

declined to 128 exports, 55 in 2011, and 

42 in 2012 (CITES lion gross exports, 

http://trade.cites.org/, accessed April 

25, 2014). In 2012, Tanzania established 

an annual quota to limit trophy hunting 

to no more than 50 animals (Jackson 

2013, p. 7). Again, it should be noted 

that there may be discrepancies between 

the annual quota and the actual number 

of trophies exported in a given year (see 

http://www.cites.org/common/

resources/TradeDatabaseGuide.pdf for 

additional information). Regardless, the 

numbers of lion trophies exported by 

Tanzania according to the UNEP– 

WCMC database suggest a decreasing 

trend. 

In other areas within the range of the 

African lion, the number of lions hunted 

or authorized to be hunted annually has 

remained fairly consistent. In Burkina 

Faso, approximately 12 lions per year 

have been hunted over the past two 

decades (IUCN 2009, pp. 36–37; Bauer 

and Nowell 2004, p. 36), although their 

current annual quota is 6 animals. In 

Botswana, a quota of 30 lions per year 

was authorized for nearly two decades; 

however, Botswana has recently 

implemented a hunting moratorium 

(Jackson 2013, p. 8). (CITES lion gross 

exports, http://trade.cites.org, accessed 

April 23, 2014; CITES UNEP–WCMC 

database, accessed January 8, 2014, and 

August 16, 2013). 

Potential Impacts of Trophy Hunting 

Infanticide and Age-Based Hunting 

Strategies 

Tourist safari hunting of males has 

been suggested by the petitioners to 

increase infanticide rates (when males 

kill young lion cubs sired by other 

males) (Petition 2011, p. 24; Whitman et 

al. 2004, p. 175), due in part to trophy 

hunters taking males under a certain 

age. Removing a younger male lion is 

purported to allow another male to take 

over the pride, and kill the former 

patriarch’s cubs. This supposition is 

inconclusive and not well supported 

(CITES 2014a, p. 14; Dagg 2000, pp. 

831–835). Infanticide is a common 

practice among many species, including 

lions (Hausfater et al. 1984, pp. 31, 145, 

173, 487). When an adult male lion in 

a pride is killed, surviving males who 

form the pride’s coalition become 

vulnerable to takeover by other male 

coalitions, and this often results in 

injury or death of the defeated males 

(Davidson et al. 2011, p. 115). In some 

cases, replacement males who take over 

the pride will kill all cubs less than 9 

months of age in the pride (Whitman et 

al. 2004, p. 175). One range country 

specifically addressed this issue; the 

Republic of Namibia indicates that lion 

populations reproduce at similar rates 

in both harvested and non-harvested 

populations, but it is unclear whether 

cub survival is consistent in harvested 

vs. non-harvested lion populations. 

While utilizing individual-based 

simulation models, Whitman et al. 

(2004, pp. 175–177) found that if offtake 

is restricted to males older than 6 years 

of age, then trophy hunting will likely 

have minimal impact on the pride’s 

social structure and young (Packer et al. 

2006, p. 6). This 6-year age restriction 

approach for lion trophies is in the 

process of being self-implemented, 

along with other best practices, by 

professional hunting guides, and is 

being adopted by certain range states 

(White 2013, p. 14; Davidson et al. 2011, 

p. 114; Whitman et al. 2004, p. 176). It 

involves conducting an age assessment 

of male lions using identification 

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techniques, such as mane development, 

facial markings, nose pigmentation and 

tooth-aging, to establish the relative age 

of male lions. 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). 

Although these characteristics may be 

subjective, as regional differences may 

occur between lion populations, there 

are clear attempts by the trophy hunting 

community to establish and implement 

best practices. Promoting the removal of 

males 6 years of age or older, 

theoretically allows younger males the 

opportunity to remain resident long 

enough to rear a cohort of cubs 

(allowing their genes to enter the gene 

pool; increasing the overall genetic 

diversity). By removing 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; Whitman 

et al. 2004, p. 176). The governments of 

Tanzania, western Zimbabwe, 

Mozambique in the Niassa National 

Reserve, Zambia, and most recently 

Benin have instituted or are in the 

process of instituting reforms such as 6- 

year age restrictions on lion trophies to 

increase the likelihood that trophy 

hunting of lion is sustainable in those 

countries (Van der Merwe 2013, p. 2; 

Jackson 2013, p. 3; White 2013, p. 14; 

Dallas Safari Club 2013, pp. 1–2; Hunter 

et al. 2013, p. 2). 

In addition to quota-setting, 

moratoriums, and the 6 year age limit, 

it has been reported that more protective 

standards and guidelines are 

implemented, such as the best practices 

listed below (Jackson 2013, pp. 3, 8–10, 

Dallas Safari Club 2013, pp. 1–2). 

• Minimum trophy quality, sizes, and 

standards; 

• Wildlife hunting regulations 

enacted and enforced; 

• Professional hunting associations 

formed; 

• Professional hunting training 

courses; 

• Professional hunter standards 

established; 

• Quota-setting procedures; 

• Compliance with CITES 

demonstrated; 

• Monitoring; and 

• Information and data collection and 

analysis. 

While the supposition of increased 

infanticide due to the remove of 

established males from a pride is 

inconclusive and not well supported, it 

is clear that improved management 

practices are beneficial to maintaining 

viable lion populations. Developing and 

implementing best management 

practices, while not categorically 

establishing a direct correlation with 

increased population numbers and 

health, do appear to have practical 

impacts on lion populations. Based on 

the best available scientific and 

commercial information, infanticide, as 

a result of the removal of lions through 

hunting, is not a threat to African lions. 

Further, it is not likely to become a 

threat in the foreseeable future since the 

science is not well supported as to 

whether infanticide resulting from 

offtake due to trophy hunting is a 

significant threat to the subspecies 

(Whitman et al. 2004, pp. 175–176; 

CITES 2014a, p. 14). 

Corruption 

Corruption is common in some areas 

within the range of the African lion, 

particularly in areas with extreme 

poverty (Michler 2013, pp. 1–3; Kimati 

2012, p. 1; Garnett et al. 2011, p. 1; 

IUCN 2009, p. 89; Leader-Williams et al. 

2009, p. 296–298; Kideghesho 2008, pp. 

16–17; http://www.transparency.org). 

Several of the range countries of African 

lion have experienced political 

instability for many years, which 

appears to be a contributing factor in 

intensifying levels of corruption. 

Political instability results in war and 

famine, which essentially halt 

conservation efforts and the 

enforcement of existing wildlife 

protection laws (Barnett & Patterson 

2005, p. 82). Corruption manifests itself 

in several ways, including 

embezzlement of funds and acceptance 

of bribes to overlook illegal activities or 

for political influence (Garnett et al. 

2011, p. 1). Given the financial aspects 

of sport hunting, it is reasonable to 

assume that corruption and the inability 

to control it could have a negative 

impact on decisions made in lion 

management by overriding biological 

rationales with financial concerns. 

Corruption has complex roots and 

will not end immediately, but from all 

appearances, it is being addressed in 

many of the African lion range countries 

where it has occurred in the past. 

Countries throughout the range of the 

African lion are putting tools in place to 

combat corruption and create awareness 

(http://www.transparency.org/cpi2012/

results, accessed June 20, 2013). In 

recent years, in several African lion 

range countries, leadership 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 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 currently 

prohibited throughout the country 

(Michler 2013, pp. 1–3). According to 

some authors (Martin 2012, pp. 4, 104; 

Kimati 2012, p. 1; Kideghesho 2008, pp. 

16–17), corruption in the wildlife sector 

has often been one of the most 

discussed topics in Tanzania’s National 

Assembly, which presumably would 

indicate the awareness of and 

willingness to address the corrupting 

factors in the wildlife sector. 

Provided that countries continue to 

address corruption within the wildlife 

sector, we conclude, based on the best 

scientific and commercial information 

available, that corruption, in and of 

itself, does not currently pose a threat to 

the species. However, if efforts to 

address corruption do not continue, it 

could become a threat to African lions 

in the future. 

Revenue From Trophy Hunting 

The high value of lions makes them 

one of the most expensive large game 

species to hunt. The revenue derived 

from lion hunting is substantial. Lions 

are reported to generate the highest 

daily rate of any mammal hunted (USD 

$2,650 per day), the longest number of 

days that must be booked, and the 

highest trophy fee ($24,500) (Jackson 

2013, p. 6; Lindsey et al. 2012a, p. 5). 

According to Groom (2013, p. 4), a 21- 

day lion hunt in Zimbabwe may be sold 

for approximately $2,500 per day, with 

an additional trophy fee of $10,000. 

Depending on the country in which a 

hunter visits, there may be several 

different fees required, 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 & 

Patterson 2005, p. 71). In the late 1990’s, 

Tanzania reported annual revenue of 

$29.9 million from all trophy hunting, 

South Africa reported $28.4 million, 

Zimbabwe reported $23.9 million from 

all trophy hunting, Botswana reported 

$12.6 million, and Namibia reported 

$11.5 million; the revenue generated 

solely from lion hunting was not broken 

out (Barnett & Patterson 2005, p. iv). In 

the past, government and private land 

owners were the primary beneficiaries 

of the revenue gained; however, a 

portion of the revenue derived from 

hunting, in some countries, is now 

being distributed to local communities 

as well, which benefits the livelihoods 

of local people as well as contributes to 

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national economies of African range 

states (Barnett & Patterson 2005, p. vi). 

Trophy Hunting as a Wildlife 

Management Tool 

The concept of using trophy hunting 

to support lion conservation is complex 

and counterintuitive to many. Many 

range countries rely heavily on tourism 

(predominantly ecotourism and safari 

hunting) to provide funding for wildlife 

management (IUCN 2006a, p. 24). The 

countries that rely most on lion hunting 

are proportionally the highest in 

Mozambique, Tanzania, and Zambia 

(Lindsey et al. 2012a, pp. 7–8). The 

revenue generated from these industries 

provides jobs for locals, such as game 

guards, cooks, drivers, and security 

personnel, and often brings in revenue 

for local microbusinesses that sell art, 

jewelry, and other native crafts. 

Revenue generated from scientifically 

based management program is used to 

build and maintain fences, provide 

security personnel with weapons and 

vehicles, provide resources for antipoaching activities, and provides 

resources for habitat acquisition and 

management (Chardonnet et al. 2010, 

pp. 33–34; Newmark 2008, p. 321). 

Revenue from trophy hunting increases 

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). 

Proponents and most species experts 

support trophy hunting as a 

conservation tool for the African lion 

(Hunter 2011, entire; van der Merwe 

2013, entire; Hunter et al. 2013, entire) 

because it provides: (1) Incentives for 

the conservation of large tracts of prime 

habitat, and (2) funding for park and 

reserve management, anti-poaching, 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 the concerns about 

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). In 

Tanzania, 25–33 percent of the total 

area, encompassing 190 hunting units 

and over 247,000 km2, 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 & Patterson 2005, p. 61). 

In Botswana, despite the current ban 

on lion hunting, the country currently 

has over 128,000 km2 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 

is in addition to 111,000 km2 (or 19.1 

percent) that has been set aside as 

habitat in the form of National Parks, 

Game Reserves, and Forest Reserves 

(Barnett & Patterson 2005, p. 7). 

Tanzania 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 & Patterson 

2005, pp. 76–77). In 2000, five countries 

in southern Africa (Botswana, Namibia, 

South Africa, Tanzania, and Zimbabwe) 

had set aside a combined 420,000 km2

of communal land, 188,000 km2 of 

commercial land, and 420,089 km2 of 

state land totaling over 1,028,000 km2

for sport hunting purposes (Barnett & 

Patterson 2005, p. iii). As a species with 

a considerable range (up to 1,000 km2) 

(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 for use in sport hunting has 

helped to reduce, but not eliminate, the 

impact of habitat loss for the African 

lion. 

Cost estimates for maintaining lion 

populations range, from an annual 

budget of $500 per km2 in smaller 

fenced reserves to $2,000 per km2

annually for unfenced populations 

(Packer et al. 2013, p. 640; Lindsey et al. 

2012a, p. 9). This includes but is not 

limited to costs associated with 

permanent and temporary staff, fencing 

installation and maintenance (fences 

can cost $3,000 per km to install), 

infrastructure maintenance, antipoaching activities such as surveillance 

and snare/trap removal, wildlife 

restocking fees (both for lions killed by 

illegal poaching/snares as well as other 

trophy species killed by lions on the 

reserves), community outreach, and 

compensation for loss of livestock in 

surrounding communities (Packer et al. 

2013, p. 640; Groom 2013, pp. 4–5; 

Lindsey et al. 2012a, p. 9; Barnett & 

Patterson 2005, p. 82). For example, in 

the past, the Save ́ Valley Conservancy in 

Zimbabwe invested $546,000 annually 

on anti-poaching activities and 

employed 186 permanent scouts, while 

operators in Coutada 16, Mozambique, 

spent $60,000 annually on antipoaching (such as the removal of 5,000 

gin traps) (Groom 2013, p. 5; Lindsey et 

al. 2012a, p. 9). According to Barnett 

and Patterson (2005, p. 82), in 

Zimbabwe: 

Land invasions, resettlement and political 

instability has had dire consequences for 

wildlife occurring in the commercial sector. 

Land invasions have affected all wildlife 

management activities, and resulted in severe 

habitat destruction, increased poaching and 

infrastructure damage with thousands of 

kilometers of fences being destroyed to make 

wire snares . . . A typical questionnaire 

response from an invaded 50,000 acre farm 

in Masvingo Province . . . indicates 

substantial poaching losses of up to 

$1,819,040, with over 3,400 snares recovered 

and 134 poachers arrested in just two 

months. 

Niassa National Reserve, 

Mozambique, incurs annual costs of 

approximately $1.9–2 million to 

maintain a 42,000-km2 area (Lindsey et 

al. 2012a, p. 9). As a single source of 

revenue, the trophy hunting of lions 

provides a substantial source of funds to 

pay for the management of lion habitat. 

According to Lindsey et al. (2012a, p. 5), 

with the exception of rhinoceros and 

exceptional elephant trophies, ‘‘lions 

generate the highest revenue per hunt of 

any species in Africa.’’ In Niassa 

National Reserve, lion trophy hunting 

has generated $380,000–400,000 

annually (Lindsey et al. 2012a, p. 9). In 

the Save ́ Valley Conservancy, between 

2005 and 2011, lion hunting in 

Zimbabwe provided an estimated net 

income (based on 26 lions) of 

approximately $1,365,000 in per-night 

charges and roughly $260,000 in trophy 

fees (Groom 2013, p. 4). 

Trophy hunting of lions, if part of a 

scientifically based management 

program, can provide direct benefits to 

the species and its habitat, both at the 

national and local level (See: Role of 

Local Communities in Lion 

Conservation). 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 

hunting alone will not address all of the 

issues that are contributing to the 

declined status of the species, it can 

provide benefits to the species. 

Role of Local Communities in Lion 

Conservation 

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, 

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they have incentive to protect it. 

Therefore, utilizing the wildlife sector 

as a land-use option and source of 

income for rural populations has 

increasingly been employed throughout 

the range countries of the African lion. 

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 

populations living in severe poverty 

(United Nations Development 

Programme’s (UNDP) Human 

Development Report, http://

hdr.undp.org/en/data, accessed July 7, 

2014; Barnett & 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 & 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 

high 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 [editor] 1990, pp. 1, 5– 

15). 

Studies have indicated that, in order 

for species such as the African lion to 

persist, the 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 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 beneficial 

value of lions being present in their 

communal areas, sustainable utilization 

of lions as a land-use 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 & 

Patterson 2005, p. iii). When the value 

of lions in areas outside of 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 them 

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 km2 (61,776 mi2) 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. For example, in 2012, the Save ́

Valley Conservancy (Zimbabwe) 

‘‘provided over US$100,000 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 at an estimated value 

of $250,000 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. 

Lastly, local communities benefit from 

the trophy hunting industry by gaining 

employment as cooks, drivers, game 

guards, security, and anti-poaching 

personnel, and they also obtain revenue 

for items purchased by trophy hunters 

such as jewelry, art, and native 

handicrafts. 

Trophy hunting as part of a 

scientifically based management 

program may provide direct economic 

benefits to the local communities and 

can create incentives for local 

communities to conserve lions, reduce 

the pressure on lion habitat, and end 

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 are perceived to kill livestock, 

which do have value to communities 

(see Human-lion Conflict). 

Many range countries have realized 

local communities must benefit from the 

conservation of the species because 

[why?] and have revised their land 

management and ownership policies to 

reflect this. Of the ten countries where 

lion trophy hunting currently occurs 

(including Cameroon), 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 has incorporated sustainable 

natural resource development as a main 

priority, and emphasized benefit 

distribution and management to rural 

communities (Benin 2000, unpaginated; 

Burkina Faso 2000, unpaginated; 

unpaginated; CAR 2000, p. 45; 

Mozambique 2000, unpaginated; 

Tanzania 2000, pp. 13, 21; Zambia 2000, 

unpaginated). As a result, an increase in 

participation by local communities in 

managing natural resources that are 

adjacent to reserves is occurring in 

several areas. 

Captive Lions 

In analyzing threats to a species, the 

Service focuses its analysis on threats 

acting upon wild specimens within the 

native range of the species, because the 

goal of the Act is survival and recovery 

of the species within its native 

ecosystem. We do not separately 

analyze ‘‘threats’’ to captive-held 

specimens because the statutory five 

factors under section 4 (16 U.S.C. 1533) 

are not well-suited to consideration of 

specimens in captivity and captive-held 

specimens are not eligible for separate 

consideration for listing. However, we 

do consider the extent to which 

specimens held in captivity create, 

contribute to, reduce, or remove threats 

to the species. 

Captive-held African lions, including 

those that are managed for trophy 

hunting in South Africa and lions held 

in captivity in zoos, are believed to 

number between a few thousand and 

5,000 worldwide (Republic of South 

Africa 2013, p. 5; Barnett et al. 2006a, 

p. 513). Captive lions in general are not 

suitable for reintroduction due to their 

uncertain origins (Barnett et al. 2006a, 

p. 513; Hunter et al. 2012, p. 3), 

potential maladaptive behaviors, and 

higher failure risk compared to 

translocated individuals (Hunter et al. 

2012, pp. 2–3). There may be cases 

where captive specimens provide a 

benefit to the species under certain 

circumstances. For example, the display 

of Giant pandas in U.S. zoos has 

generated considerable revenue that is 

used for in-situ conservation of the 

species in China. It may be possible that 

captive lions could also serve a purpose 

of generating revenue for in-situ 

conservation. 

Summary of Trophy Hunting 

Although there is some indication 

that trophy hunting could contribute to 

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local declines in lion populations 

through unsustainable quotas, 

corruption, and possible disruption of 

pride structure through infanticide and 

take of males that are too young, we do 

not find that any of these activities rises 

to the level of a threat to the African 

lion subspecies at this time. It appears 

that most range countries that allow 

trophy hunting of African lions restrict 

offtake to approximately 2–4 percent of 

their lion populations for trophy 

hunting annually, excluding South 

Africa, where offtake is from 

predominantly captive-born animals, 

and Zimbabwe, where offtake is 2–3 

percent higher than in other countries 

(Packer et al. (2006, pp. 2–3). Exports of 

lion trophies have demonstrated a 

decreasing trend when exports of likely 

captive-born lions from South Africa are 

excluded (CITES lion gross exports, 

http://trade.cites.org, accessed April 23, 

2014), and lions from South Africa are 

likely captive-born (RSA 2013, p. 5). 

Most of the range countries that allow 

trophy hunting have quotas in place to 

limit take. Tanzania, with a population 

of approximately 16,000 lions, has a 

quota of 50 animals per year. Many 

other range countries have laws in effect 

that address trophy hunting, and several 

have moratoriums in place. The hunting 

community is taking the lead in 

developing best management practices 

to address take of males that are under 

6 years of age, and they are guiding the 

development of scientifically based 

tools for minimizing the impact of 

trophy hunting on the social structure of 

lion populations. This 6-year age 

restriction on lion trophies is in the 

process of being self-implemented by 

professional hunting guides, and is 

being adopted by certain range states, 

such as Tanzania (White 2013, p. 14; 

Whitman et al. 2004, p. 176). 

Currently, most countries that allow 

trophy hunting of lions appear to be 

reviewing their trophy hunting practices 

(Jackson 2013, pp. 2–3; White 2013, pp. 

12–13). Range countries have 

recognized the need to incorporate best 

management practices, and have been 

progressively updating the policies and 

management systems in order to 

implement them (Lindsey et al. 2013a, 

pp. 4–10). 

Finally, we found that, if trophy 

hunting of lions is part of a scientifically 

based management program, it could 

provide considerable benefits to the 

species, by reducing or removing 

incentives by locals to kill lions in 

retaliation for livestock losses, and by 

reducing the conversion of lion habitat 

to agriculture. Trophy hunting, if 

managed well and with local 

communities in mind, can bring in 

needed revenue, jobs, and a muchneeded protein source to local people, 

demonstrating the value of lions to local 

communities (Groom 2013, pp. 1–3; 

Lindsey et al. 2006, pp. 283, 289). In 

addition, the amount of habitat that has 

been set aside by range countries 

specifically for trophy hunting has 

greatly increased the range and habitat 

of lions and their prey base, which is 

imperative given the current ongoing 

rate of habitat destruction occurring in 

Africa. The total amount of land set 

aside for trophy hunting throughout 

Africa exceeds the total area of the 

national parks, providing half the 

amount of viable lion habitat 

(Chardonnet et al. 2010, p. 34; Packer et 

al. 2006, pp. 9–10). However, expanding 

protected areas without taking the 

human population into consideration 

could lead to more resentment and 

retaliatory killing of lions (Nelson et al. 

2009, p. 315). 

Therefore, we conclude, based on the 

best scientific and commercial 

information available, that trophy 

hunting is not a significant threat to the 

species. 

Traditional Use of Lion Parts and 

Products 

CITES (2014, p. 8) reports that many 

African countries, including Somalia, 

Nigeria, Burkina Faso, Kenya, and 

Cameroon, maintain local markets in 

lion products, which include teeth, 

claws, fat, whiskers, bone, bile, testicles, 

meat, and tails for use as talismans, 

decorations, and in traditional African 

medicine. In Ghana, lion parts and 

products are used for ceremonial, 

medicinal, and nutritional purposes 

(Burton et al. 2010, p. 4). Skins and 

claws of lions were observed for sale in 

a market in Tamale, Ghana. Lions in and 

around Mole National Park in Ghana 

have been killed for traditional 

consumptive purposes (Burton et al. 

2010, p. 4). In some cases, lions (either 

alive or dead) have been ‘‘laundered’’ 

through other countries so that their 

country of origin is unknown. As an 

example, lions have been found to be 

shot in Zimbabwe and Mozambique and 

declared as South African trophies (Lion 

Aid 2011, p. 20). In other cases, there 

have been reports of captive-born lions 

being smuggled between Botswana and 

South Africa and described as wild 

(Mouton 2013, pp. 1–2). Lion products, 

such as the trade in lion bone, seem to 

be primarily byproducts of trophy 

hunting; hunters are primarily 

interested in the trophy and skin and, 

therefore, the bones and other parts are 

sold separately (CITES 2014a, p. 10). 

However, since the reports of these 

types of activities are primarily 

anecdotal in nature, based on the best 

available scientific and commercial 

information, we find that the sale of 

these byproducts does not currently 

pose a threat to the species. Further, 

without a significant shift in the market, 

it is not likely to become a threat in the 

foreseeable future. 

Conservation Measures in Place To 

Protect Lions 

There has been awareness for several 

years that conservation strategies need 

to be implemented for the African lion 

due to the apparent decrease in its 

population numbers (Hamunyela et al. 

2013, p. 1; Henschel et al. 2010, p. 34; 

Gebresenbet et al. 2009, p. 5; IUCN 

2006a, b, entire). Prior to 2006, 

institutional inconsistencies throughout 

the African lion’s range resulted in poor 

lion conservation policies and little to 

no enforcement of existing laws (IUCN 

2006b, p. 18). As mentioned, in 2005 

and 2006, nongovernmental 

organizations (NGOs) and several 

governments at various levels organized 

two regional lion conservation 

workshops. Species specialists, wildlife 

managers, and government officials 

attended these regional workshops in 

order to provide range country 

governments with frameworks for 

developing their own national action 

plans for the conservation of lions. Over 

50 lion specialists, representing all lion 

range countries, participated in these 

workshops (Henschel et al. 2010, p. 34). 

During the workshops lion experts 

collectively assessed what they believed 

to be the then-current status of African 

lions based on a variety of information, 

and subsequently identified 86 African 

LCUs. This information was then used 

as a framework to identify lion areas, 

strongholds, and potential strongholds 

by Riggio et al. (2013, p. 32). 

Many countries with very small lion 

populations have developed or updated 

their conservation plans for the African 

lion. Some of these include Benin, 

Cameroon, Uganda, and Malawi. Some 

range countries participate in 

transboundary conservation projects 

and are collaborating on transboundary 

lion conservation initiatives for shared 

lion populations. Most range countries 

have a national lion action plan or 

strategies in place, particularly if there 

are economic incentives for them to 

have viable lion populations (Groom 

2013; Nghidinwa et al. 2013, pp. 11–12; 

Zambia Wildlife Authority 2012; Lion 

Aid 2011, pp. 1–2; Mesochina et al. 

2010; Government of Tanzania 2010; 

Begg and Begg 2010). Range states have 

also implemented a number of 

conservation strategies designed to 

conserve habitat, reduce human-lion 

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conflict, and preserve the lion’s preybase. 

Conservation Measures To Stem Habitat 

Loss 

Habitat loss represents one of the 

main threats facing the African lion 

(Bauer et al. 2008, unpaginated). 

Attempts by range countries to address 

this decline in habitat are manifested in 

a number of ways, such as the creation 

of protected areas and the establishment 

of wildlife corridors to connect 

fragmented habitats. 

Two conservation tools utilized by 

range countries for African lions include 

the establishment of protected areas and 

the enforcement of protections in these 

areas (Mesochina et al. 2010a and b; 

Treves et al. 2009, pp. 60, 64). Over the 

past few decades, the effectiveness of 

protected areas in protecting habitat has 

been studied, particularly in Africa 

(Pfeifer et al. 2012, p. 1; Craigie et al. 

2010, pp. 2,221–2,222). A study 

conducted by the Wildlife Conservation 

Society in 2005 found that most lion 

populations in protected areas of 

southern and eastern Africa have been 

essentially stable over the previous 

three decades (Ray et al. 2005, p. 67). 

However, several problems have 

emerged. For example, certain landtenure systems do not recognize 

community ownership of land and 

wildlife and undermine the extent to 

which benefits are converted into 

incentives for conservation. Protectedarea ‘‘boundaries’’ are not always 

visible. Additionally, law enforcement 

in protected areas can be sporadic, and 

parks are often understaffed (Pfeifer et 

al. 2012, pp. 1, 7). Lastly, despite the 

Wildlife Conservation Society’s 

findings, more recent evidence suggests 

that some protected areas are being 

more commonly encroached upon as 

human populations expand and search 

for resources. 

Despite encroachment, protected 

areas are somewhat effective at 

protecting wildlife and habitat as rates 

of habitat loss tend to be lower in 

protected areas than outside them 

(Green et al. 2013, p. 70; Pfeifer et al. 

2012, p. 2). African countries are 

realizing the benefits of managing their 

wildlife populations and parks for 

tourism; however, conservation of vast 

areas of land for megafauna such as the 

African lion is not only complex, but 

also expensive. As an example, the 28- 

km (17-mi) elephant corridor, 

completed in 2011 in Kenya, cost $1 

million (The Nature Conservancy 2013, 

unpaginated). Additionally, the overall 

costs of anti-poaching and 

compensation is expected to increase in 

range states concurrently with growing 

human populations, declining 

purchasing power of external funds, and 

corruption (Garnett et al. 2011, pp. 1–2; 

Wittemyer et al. 2008, pp. 123, 125). 

Another mechanism for protecting 

habitat is to reconnect fragmented 

habitat across national boundaries. 

Corridors are being restored, fences are 

being removed, and protected areas are 

being connected. Restoration of these 

corridors allows wildlife to travel 

between areas of suitable habitat (Jones 

et al. 2012, pp. 469–470). In some areas, 

fences have been constructed to protect 

grazing resources for domestic livestock 

as well as to provide barriers to disease 

(Gadd 2012, pp. 153, 176). One aspect 

of these fences is that they separate 

lions from their prey. In southern 

Africa, the trend now is to take down 

fences to increase the size of connected 

habitat and link it to reserves and 

national parks (IUCN 2009, p. 101; 

IUCN 2008, various). The Limpopo 

Transfrontier Park is another example of 

where this is being implemented 

(Newmark 2008, p. 327). Boundary 

fences along national borders that 

separate many reserves are being 

removed to form a 35,000-km2 park. 

Limpopo National Park (formerly 

known as Coutada 16) in Mozambique; 

Kruger National Park in South Africa; 

Gonarezhou National Park, Manjinji Pan 

Sanctuary, and Malipati Safari Area in 

Zimbabwe will all be connected, as will 

be the area between Kruger and 

Gonarezhou, and the Sengwe communal 

land in Zimbabwe and the Makuleke 

region in South Africa (Newmark 2008, 

p. 327). However, in some locations, 

areas that have previously been 

designated as corridors have been 

encroached upon by human settlements 

and agriculture (Estes et al. 2012, pp. 

258–261; Jones et al. 2012, p. 469). 

Tanzania is an example of a country 

attempting to reconnect habitat. As of 

2002, the Tanzanian Government, with 

donor and NGO support, was 

reconnecting the nine largest blocks of 

forest in the East Usambara Mountains 

using wildlife corridors (Newmark 2002, 

various). Additionally, the 2009 

Wildlife Act of Tanzania allows the 

Minister, in consultation with relevant 

local authorities, to designate wildlife 

corridors, dispersal areas, buffer zones, 

and migratory routes. The 2010–2015 

National Elephant Management Plan of 

Tanzania indicates that corridors are the 

primary objective of the plan, and 

although primarily designed for 

elephants, these corridors allow for 

continuity of populations of other large 

mammal species such as lions (Jones et 

al. 2012, p. 470). 

In 2011, Kenya (which neighbors 

Tanzania to the North), completed a 28- 

km corridor through an area that had 

been heavily impacted by humanwildlife conflict. The purpose of the 

corridor was primarily to reduce 

human-elephant conflict and appears to 

have been successful (Mount Kenya 

Trust 2011, p. 1). The corridor also 

allows other wildlife such as lions to 

disperse through habitat that otherwise 

would have been unfavorable for 

wildlife to travel through (Mount Kenya 

Trust 2011, p. 1). It was an expensive 

project, but recent reports indicate that 

the effort has served its purpose: 

Elephants are using the corridor on a 

regular basis (particularly an underpass 

under a highway), and humans are 

reporting less human-wildlife conflict 

(Mount Kenya Trust 2011, p. 1). 

However, connectivity alone does not 

ensure the dispersal of animals (Roever 

et al. 2013, pp. 19–21). The Tanzania 

Wildlife Research Institute (TAWIRI) is 

a parastatal organization under 

Tanzania’s Ministry of Natural 

Resources and Tourism, and is 

responsible for conducting and 

coordinating wildlife research activities 

in Tanzania (http://tawiri.or.tz/). In this 

role, TAWIRI has been actively involved 

in promoting the development of and 

monitoring the use of wildlife corridors 

in Tanzania (http://

www.tzwildlifecorridors.org). Surveys 

conducted in 2009 and 2010 suggest 

that the Nyanganje Corridor in Tanzania 

is no longer being used by elephants 

and other wildlife. This corridor is at a 

narrow passage in the Kilombero Valley 

and is the shortest distance for animals 

to cross between the Udzungwa and 

Selous ecosystems. Despite efforts in 

place, much of the corridor is being 

encroached upon by conversion of land 

to rice farming and cattle grazing (Jones 

et al. 2012, p. 469). Because these 

activities often deter wildlife from 

passing through, the corridor is 

ineffective (Jones et al. 2012, p. 469). 

TAWIRI reminds wildlife managers that 

they need to continue to implement 

steps to ensure that corridors are 

functioning properly. 

Conservation Measures in Place To 

Stem the Loss of Prey Base 

Lions, like most large carnivores, prey 

upon a variety of species including 

buffalo, plains zebra, wildebeest, giraffe, 

gemsbok, kob, and warthog (Kenya 

Wildlife Service 2013, p. 13; Niassa 

National Reserve Technical Report 

2011, p. 4; Nowell and Jackson 1996, p. 

18). Depletion of these prey species due 

to competition with humans represents 

a threat to the lion (Chardonnet et al. 

2005, pp. 8–9). As noted, the increase in 

the human population in Africa is a 

major contributor to the increase in the 

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demand for bushmeat, which in turn 

increases human encroachment into 

wildlife lands (Lindsey et al. 2012b, p. 

36). In addition to the increase in the 

human population, lack of an 

alternative livelihood, lack of alternate 

food sources, and lack of clear rights 

over land or wildlife are contributing 

factors toward the increase in demand 

for bushmeat (Lindsey et al. 2012b, pp. 

36–41). The advent of automatic 

weapons in the bushmeat trade impacts 

the lion’s prey base, which is being 

hunted at unsustainable levels. 

Reconnecting fragmented habitat has 

the additive effects of not only 

conserving the biodiversity of the 

African lion’s habitat, but also that of its 

prey base (Lindsey et al. 2012b, p. 43). 

These types of restoration practices 

enhance the health of species by 

allowing genetic interchange to occur 

and, thus, conserve the genetic diversity 

of all wildlife. Wildlife management 

entities are linking many of the major 

protected areas by removing boundary 

fences along national borders that 

separate many reserves in addition to 

creating or improving corridors to link 

good-quality habitat for wildlife (Gadd 

2012, p. 179; Newmark 2008, pp. 323– 

324). To address the increasing 

consumption of bushmeat, host 

countries have employed a variety of 

different strategies, including the 

development of alternative industries 

for communities. Helping local 

communities develop alternate 

industries represents one of the ways 

range countries can reduce their 

dependence on bushmeat. Throughout 

Africa, several ideas have been 

attempted with varying levels of 

success. For example, the Anne Kent 

Taylor Fund (AKTF) helps local Maasai 

women to buy beads and other supplies 

to produce traditional items for the local 

tourist industry (AKTF 2012, p. 7; 

Lindsey et al. 2012b, p. 45; van Villet 

2011, p. 17). In addition, AKTF helps 

organize local men into anti-poaching 

and de-snaring teams (AKTF 2012, p. 5; 

van Villet 2011, p. 17). By creating 

programs targeting both men and 

women, AKTF creates an environment 

that provides communities with 

financial stability as well as direct 

community interest in protecting local 

wildlife. With 13 years assisting local 

communities, the AKTF represents one 

of the more successful attempts to 

encourage locals to shift away from 

relying on bushmeat. 

Studies compiled by Huzzah 2013 

(pp. 1, 8) have shown that local 

communities who lived near protected 

areas with more lenient policies have a 

more positive attitude and relationship 

with both the manager and the protected 

area as a whole. This open approach to 

protected area management reflects a 

trend in recent years to bring in local 

communities to assist in the 

management of protected areas (Lindsey 

et al. 2012b, p. 53). Wildlife 

management programs run by local 

communities are defined by two goals: 

Conserving wildlife and providing 

economic aids to the community 

(Bandyopadhyay et al. 2010, p. 5). With 

regards to discouraging the 

consumption of bushmeat, this new 

approach is seen in the creation of 

community-based wildlife management 

programs (van Villet 2011, p. 26). The 

purpose of these programs is to give the 

local community a direct stake in the 

management of wildlife areas. One use 

for these areas is to turn them into game 

ranches. These areas are used both for 

legal bushmeat production as well as 

trophy hunting and ecotourism. 

One such program is the Chivaraidze 

Game Ranch in Zimbabwe (van Villet 

2011, pp. 28–29). The Chivaraidze Game 

Ranch started in 1996 with the stated 

goal of reducing poaching through 

providing bushmeat at a reduced price. 

However, internal infighting in the 

organization over the devolution of 

power to local communities, between 

those in favor of devolution and a 

powerful local interest group, limited 

the effectiveness of the organization. In 

the span of 8 years (between 2001 and 

2009), the Chivaraidze Game Ranch has 

had six different boards of directors 

(Mombeshora and Le Bel 2010, p. 5). 

Furthermore, a power shake-up in local 

communities along party lines and 

kinship affiliation limited the abilities 

for communities to cooperate with each 

other (van Villet 2011, pp. 28–29; 

Mombeshora and Le Bel 2010, p. 7). The 

result was that the cost of maintaining 

the program exceeded the benefits to the 

local community. The decline in 

economic benefits to the local 

community coincided with a resurgence 

in poaching within areas of the park 

(Mombeshora and Le Bel 2010, p. 3). 

The result of the Chivaraidze Game 

Ranch project reflects the difficulty in 

shifting wildlife management from a 

centralized national government 

approach towards a more decentralized, 

community-based approach. 

Unlike the difficulties encountered in 

Zimbabwe, Namibia has had greater 

success in setting up community-run 

conservancies. After gaining 

independence in 1990, Namibia began 

to turn over ownership of wildlife areas 

to local communities (van Vliet 2011, p. 

29; Bandyopadhyay et al. 2010, p. 6). By 

2011, Namibia had 64 communities that 

covered 17 percent of the country total 

area (van Vliet 2011, p. 29; Connif 2011, 

npn; NASCO 2010, p. 4). The majority 

of the incomes from these conservancies 

come from ecotourism, followed by 

trophy hunting (NASCO 2010, p. 22). 

These incomes are then used to support 

infrastructure improvement in the 

community. In addition, legal bushmeat 

acquired within conservancy lands is 

distributed to local families (NASCO 

2010, p. 25). The success of the program 

in Namibia has been attributed to 

Namibia’s unique characteristics, 

including low population density and 

favorable seasonal rain, which helps 

prey species recover (van Vliet 2011, p. 

30). Despite the successes in Namibia, 

the country’s unique characteristics 

mean that adapting Namibia’s success to 

other, more densely populated countries 

will be difficult. 

Conservation Measures To Stem 

Human-Lion Conflict 

As the human population expands, 

the potential for conflict with wildlife 

increases. In Africa, conflict between 

villagers and lions, who prey upon 

livestock, represent a threat to the 

species (Chardonnet et al. 2010, p. 12; 

Moghari 2009, p. 14; IUCN 2006a, p. 

23). In addition, habitat loss due to 

conversion of land increases the chance 

of villagers coming into direct contact 

with lions (Chardonnet et al. 2010, p. 

24). In an attempt to address these 

problems, range countries have 

employed a variety of different 

strategies to help the lion. Such 

strategies involve education, an effective 

conservation plan, and interacting with 

the local community. 

Historically, range countries seek to 

mitigate human-lion conflict through 

controlling rather than conserving the 

predator population. In countries such 

as Malawi, for example, the Department 

of Game, Fish and Tsetse Control would 

shoot large carnivores that prey upon 

livestock. The result of this policy was 

that, between 1948 and 1961, over 560 

predators (which include lions and 

leopards) were killed in the country 

(Mesochina et al. 2010b, p. 35). While 

this department was disbanded in 1963 

and jurisdiction shifted to the new 

Department of Forestry, crop and 

livestock protection still remains an 

important part of its function. Despite 

the department focusing on protecting 

crops and livestock, the number of lions 

killed in the country has declined. 

Between 1977 and 1982, eight lions 

were killed, whereas six lions were 

killed between 1998 and 2007 

(Mesochina et al. 2010b, p. 35). While 

fewer lions are being killed than in the 

previous decades, problems remain, 

including lack of resources, lack of 

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8ECOLEX is a comprehensive database on 

environmental law, maintained by the International 

Union for Conservation of Nature (IUCN), the 

United Nations Environment Programme (UNEP), 

and the Food and Agriculture Organization of the 

United Nations (FAO). Our search terms used with 

respect to wildlife laws were ‘‘African lion’’ and 

‘‘country’’, e.g., ‘‘Angola’’, ‘‘Benin’’, etc. See 

Appendix A. 

manpower, and corruption within the 

range countries. 

Current governmental management of 

lions in countries such as Malawi, 

Tanzania, and Zambia are managed by 

the Problem Animal Control units 

(Mesochina et al. 2010a, p. 41; 

Mesochina et al. 2010b, p. 36). When 

lion attack incidents occur, Problem 

Animal Control dispatches officials to 

investigate the problems. If the problem 

lion is located, it is either removed or 

eliminated. When properly funded, this 

program has helped in reducing not 

only conflicts between lions and 

humans but also has driven down the 

numbers of lions killed. Between 2005 

and 2009, there were 116 reported cases 

of lions killed, with the number of lions 

killed being less than 50 per year in 

Tanzania (Mesochina et al. 2010a, p. 

41). However, limitations of resources 

(including both manpower and funds) 

have hampered the effectiveness of 

these officials in responding to these 

incidents. In addition, many Problem 

Animal Control interventions resulted 

in the death of the lion (Mesochina et 

al. 2010a, p. 41; Chardonnet et al. 2009, 

p. 36). Even in cases of translocation, 

the lions that were being transported 

often end up injured or continue to pose 

problems to the community (Bauer et al. 

2007, p. 91). 

NGOs are also assisting in protecting 

lions. Intervention by NGOs often takes 

the form of interacting with the local 

community (Winterbach et al. 2010, p. 

98). Lion Guardians, which operate in 

Kenya, recruits and educates local 

young men. These men then monitor 

and track lion movement and warn 

herders of lion presence in the area, 

thereby mitigating or preventing 

possible lion-human conflict (Hazzah et 

al. 2014, p. 853; Lion Guardians 2013, 

p. 7; Lion Guardians 2012, p. 3). In 

addition, Lion Guardians work with 

tribal elders to dissuade young men 

from killing lions for ceremonial 

purposes. Historically, the killing of 

lions through ritualized lion hunts 

called ilmurran is rewarded with gifting 

of cows and other rewards (Lion 

Guardians 2012, p. 5; Goldman et al. 

2010, p. 334). After introducing village 

elders to the Lion Guardians program 

first hand, many return home to their 

village and give their blessings to the 

project. This education led to significant 

results; on August 11, 2013, two Lion 

Guardians stopped a group of hunters 

who were planning to hunt a lion in 

retaliation for the lion preying on their 

livestock. The local village elders fined 

the potential hunters two cattle each for 

going on a lion hunt, marking a gradual 

but significant shift in the cultural 

attitudes regarding the lion (Hazzah et 

al. 2014, p. 858; Lion Guardians 2013, 

p. 20). Since its establishment in 2007, 

only five lions had been killed in 

territories where Lion Guardians 

operates, in contrast to more than 100 

lions killed in adjacent areas (Lion 

Guardians 2013, p. 5). Furthermore, 

reduced lion mortality was sustained 

across multiple years, resulting in the 

reserve having one of the highest lion 

densities in Africa (Hazzah et al. 2014, 

p. 857; Schuette et l. 2013, p. 149). 

Despite the success of this program, 

retaliatory as well as ceremonial killings 

of lions outside the program areas 

remain a threat to the species. 

We found that many of the lion range 

states are trying to address lion 

conservation through the establishment 

of protected areas, wildlife management 

areas, wildlife corridors, and 

reconnecting habitat. In some areas, 

creating incentives for lion conservation 

is occurring through community 

conservation programs in range 

countries. In other cases, participatory 

strategies have been implemented to 

enhance local tolerance for large 

carnivores in Africa. An increasing 

number of programs encourage local 

communities to solve problems that 

arise from human-lion conflict without 

killing lions. However, the effectiveness 

of these measures still ranges from 

successful to unsuccessful, due in part 

to lack of resources, political will, and 

infighting. It is imperative that range 

countries continue to recognize and 

support the role that local communities 

play in lion conservation. Greater 

support by countries to address the 

needs of local communities, and thereby 

address the needs of lions, may be the 

single-most important role these 

countries can play in changing the 

trajectory of lion declines. 

Regulatory Mechanisms 

Regulatory mechanisms in place to 

provide protections to African lions 

vary substantially throughout Africa. As 

mentioned in the Conservation Status of 

African Lions CITES section, lions are 

listed in Appendix II under CITES, and 

with the exception of South Sudan, all 

of the lion range states are parties to 

CITES. According to the draft CITES 

Periodic Review of the Status of African 

Lions (CITES 2014a, pp. 14–15) outside 

of CITES, lions have no legal protections 

in four countries: Burundi, Guinea 

Bissau, Lesotho, and Swaziland. 

However, CITES 2014a (p. 15), states 

that most of the southern and eastern 

lion range states have regulatory 

mechanisms in place to protect lions. 

We found that most of the range states 

have national environmental legislation 

to establish national parks and 

conservation areas, and to conserve and 

regulate the take, hunting, and trade of 

wildlife, including parts and products, 

but could find no legislation specific to 

lions, nor to the main threats affecting 

lions: habitat loss, human-lion conflict, 

and loss of prey base (See: Appendix A, 

Ecolex information was accessed July 7– 

10, 2014, at http://www.ecolex.org.8). 

Our status review did not reveal 

regulatory mechanisms in place that 

specifically address the main threats 

affecting lions. We are requesting 

comments or information from lion 

range states, other concerned 

governmental agencies, the scientific 

community, or any other interested 

parties concerning regulatory 

mechanisms that address the three main 

threats to lions: habitat loss, human-lion 

conflict, and loss of prey base. 

Finding 

Section 4 of the Act (16 U.S.C. 1533) 

and its implementing regulations at 50 

CFR 424 set forth the procedures for 

adding a species to, and/or removing a 

species from, the Federal Lists of 

Endangered and Threatened Wildlife 

and Plants. As noted in the Information 

Requested section, a species may be 

determined to be an endangered or 

threatened species due to one or more 

of the five factors set forth in section 

4(a)(1) of the Act: 

(A) The present or threatened 

destruction, modification, or 

curtailment of its habitat or range; 

(B) Overutilization for commercial, 

recreational, scientific, or educational 

purposes; 

(C) Disease or predation; 

(D) The inadequacy of existing 

regulatory mechanisms; 

(E) Other natural or manmade factors 

affecting its continued existence. 

In assessing whether the African lion 

meets the definition of an endangered or 

threatened species, we considered the 

five factors in section 4(a)(1) of the Act. 

A species is ‘‘endangered’’ for purposes 

of the Act if it is in danger of extinction 

throughout all or a significant portion of 

its range and is ‘‘threatened’’ if it is 

likely to become endangered within the 

foreseeable future throughout all or a 

significant portion of its range. The 

‘‘foreseeable future’’ is the period of 

time over which events or effects 

reasonably can or should be anticipated, 

or trends extrapolated. 

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When considering what factors might 

constitute threats to a species, we must 

look beyond the mere exposure of the 

species to a factor to evaluate whether 

the species may respond to the factor in 

a way that causes actual impacts to the 

species. If there is exposure to a factor 

and the species responds negatively, the 

factor may be a threat and we attempt 

to determine how significant a threat it 

is. The threat is significant if it drives, 

or contributes to, the risk of extinction 

of the species such that the species may 

warrant listing as endangered or 

threatened as those terms are defined in 

the Act. We conducted a review of the 

best scientific and commercial data 

available regarding the status of the 

African lion and assessed whether the 

African lion is endangered or threatened 

throughout all of its range. 

There is consensus within the 

research community as well as lion 

range states that the African lion is 

impacted by a number of factors actively 

contributing to its population decline 

throughout Africa: habitat loss 

(fragmentation and degradation) (Factor 

A); decreased access to food prey 

sources (aka loss of prey base) (Factor 

B); retaliatory killing, snaring, and 

poaching (both intentional and 

unintentional), and deleterious effects 

in its viability due to small populations 

in some areas within its range (Factor E) 

(Nyanganji et al. 2012, p. 12; Seguya et 

al. 2010, p. 26). 

We find three main threats, habitat 

loss, loss of prey base, and human-lion 

conflict, are impacting lions, alone and 

in combination, such that the 

subspecies is likely to become 

endangered within the foreseeable 

future throughout all of its range. In the 

past several decades, the human 

population has been expanding with 

concomitant large decreases in lion 

habitat and likely lion numbers, 

resulting in an extremely large 

reduction in the species’ range. As 

human populations continue to rise in 

sub-Saharan Africa, the amount of land 

required to meet the expanding human 

population’s needs is constantly 

increasing. Lions are increasingly 

limited to protected areas, and human 

population growth rates around 

protected areas in Africa tend to be 

higher than the average rural growth 

rate (Wittemyer et al. 2008, entire). 

Considering the majority of the human 

population in sub-Saharan Africa is 

rural, and land supports the livelihood 

of most of the population, loss and 

degradation of lion habitat, loss of prey 

base, and increased human-lion conflict 

can reasonably be expected to 

accompany the rapid growth in subSaharan Africa’s human population into 

the foreseeable future. 

Africa has the fastest population 

growth rate in the world (UNEP 2012a, 

p. 2). The majority of the population is 

rural, and about 60–70 percent of the 

population relies on agriculture and 

livestock for their livelihood (UNEP 

2006, pp. 82, 100, 106; IAASTD 2009, p. 

2). As a result, a large portion of the 

growing population will depend 

directly on expansion of agriculture and 

livestock grazing to survive in the 

future. 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 rangeland, and the majority of 

agricultural land in Africa is devoted to 

grazing (UNEP 2012b, p. 68), thus that 

figure may be much larger. 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). Therefore, in the case of African 

lion, the best available scientific and 

commercial data that we rely upon in 

projecting future conditions for the 

purpose of this listing determination 

establish the foreseeable future to be 

2050. 

Human settlements and agricultural 

and pastoral activities have expanded 

into lion habitat and protected areas, 

decreasing prey availability and 

increasing exposure of livestock and 

humans to lions. Human-lion conflict 

and associated retaliatory killing of 

lions will continue to play a major role 

in the reduction of lion populations and 

is the greatest current threat to 

remaining lion populations. The lion’s 

prey base has decreased in many parts 

of its range in large part due to the 

bushmeat trade 

Bushmeat is the primary source of 

protein for humans in much of the lion’s 

range (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), 

comprising between 6 percent (southern 

Africa) and 55 percent (Central African 

Republic) of a human’s diet (Chardonnet 

et al. 2005, p. 9; IUCN 2006b, p. 19). 

This reliance by humans on protein 

obtained from bushmeat results in direct 

competition for prey species between 

humans and lions, and commercial 

poaching of wildlife through the use of 

automatic weapons is a significant 

threat to lion prey (Chardonnet et al. 

2010, p. 27). Because many wildlife 

species are being hunted at 

unsustainable levels to meet this 

demand within the range of the lion, its 

prey base is becoming depleted in many 

areas and has led to lion attacks on 

livestock and humans (Hoppe-Dominik 

et al. 2011, p. 452; Chardonnet et al. 

2010, pp. 6, 13–14; Frank et al. 2006, p. 

12). Given the rapid increase in humans 

and livestock by 2050, we can 

reasonably expect the conditions 

described above to worsen. Also, as 

livestock numbers increase and as 

expansion of agricultural and pastoral 

practices continue to deplete and 

degrade the habitat that lion’s prey rely 

on, the lion’s prey base is expected to 

further decline. As the lion’s prey base 

is hunted at unsustainable levels to 

meet a growing demand for food, 

livestock depredation and retributive 

killing of lions through spearing, 

shooting, trapping, and poisoning will 

continue to occur, and will likely 

increase (Dickman 2013, p. 379; HoppeDominik et al. 2011, p. 452; Chardonnet 

et al. 2010, p. 19; Gebresenbet et al. 

2009, p. 9; Hazzah and Dolrenry 2007, 

p. 3). 

Lion range countries are aware of the 

threats affecting lions, and many are 

working to address them. NGOs and 

several governments at various levels 

have organized regional lion 

conservation workshops, which have 

helped them to identify Lion 

Conservation Units. Most range 

countries have a national lion action 

plan or strategy in place (Groom 2013; 

Nghidinwa et al. 2013, pp. 11–12; 

Zambia Wildlife Authority 2012; Lion 

Aid 2011, pp. 1–2; Mesochina et al. 

2010; Government of Tanzania 2010; 

Begg and Begg 2010). Some range 

countries participate in transboundary 

conservation projects to create wildlife 

corridors and reconnect habitat, and are 

collaborating on transboundary lion 

conservation initiatives for shared lion 

populations. Reconnecting fragmented 

habitat has the additive effects of not 

only strengthening the biodiversity of 

the African lion but also that of its prey 

species (Lindsey et al. 2012b, p. 43). 

Wildlife management entities are 

linking many of the major protected 

areas by removing boundary fences 

along national borders that separate 

many reserves, in addition to creating or 

improving corridors to link good-quality 

habitat for wildlife (Gadd 2012, p. 179; 

Newmark 2008, pp. 323–324). 

Range states have also implemented a 

number of conservation strategies 

designed to conserve habitat, reduce 

human-lion conflict, and preserve lion 

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prey-base. In order to address the 

increasing consumption of bushmeat, 

host countries have employed a variety 

of different strategies, including the 

development of alternative industries 

for communities, which can reduce 

their dependence on bushmeat. For 

example, the Anne Kent Taylor Fund 

(AKTF) helps local Maasai women to 

buy beads and other supplies to produce 

traditional items for the local tourist 

industry (AKTF 2012, p. 7; Lindsey et 

al. 2012b, p. 45; van Villet 2011, p. 17) 

and has organized local men to 

participate in anti-poaching and desnaring teams (AKTF 2012, p. 5; van 

Villet 2011, p. 17). By targeting both 

men and women in the community, 

such programs provide communities 

with financial stability as well as direct 

community interest in protecting local 

wildlife. African countries are realizing 

the benefits of managing their wildlife 

populations and parks for tourism; 

however, conservation of vast areas of 

land for megafauna such as the African 

lion is expensive. The costs of antipoaching and compensation is expected 

to increase in range states concurrently 

with growing human populations, 

declining purchasing power of external 

funds, and corruption (Garnett et al. 

2011, pp. 1–2; Wittemyer et al. 2008, 

pp. 123, 125). 

Studies have shown that local 

communities who live near protected 

areas (PAs) with community-based 

conservation policies have more 

positive attitudes and relationships with 

both the park manager and the PA as a 

whole (Huzzah 2013, pp. 1, 8). This 

open approach to PA management 

reflects a trend in recent years to bring 

in local communities to assist in the 

management of PAs (Lindsey et al. 

2012b, p. 53). Wildlife management 

programs run by local communities are 

defined by two goals: conserving 

wildlife and providing economic aids to 

the community (Bandyopadhyay et al. 

2010, p. 5). NGOs are also assisting in 

protecting lions. Intervention by NGOs 

often takes the form of interacting with 

the local community (Winterbach et al. 

2010, p. 98). For example, Lion 

Guardians, which operates in Kenya, 

has shown great success with its Lion 

Guard program. Lion Guardians 

educates local young men who monitor 

and track lion movement and warn 

herders of lion presence in the area, 

thereby mitigating or preventing 

possible lion-yhuman conflict (Hazzah 

et al. 2014, p. 853; Lion Guardians 2013, 

p. 7; Lion Guardians 2012, p. 3). 

Outreach to tribal elders has 

successfully helped elders to dissuade 

young men from killing lions for 

ceremonial purposes. The result of such 

programs has been a gradual change in 

cultural attitudes towards lions (Hazzah 

et al. 2014, p. 858; Lion Guardians 2013, 

p. 20). 

Finally, many range countries rely 

heavily on tourism (predominantly 

ecotourism and safari hunting) to 

provide funding for wildlife 

management (IUCN 2006a, p. 24). The 

revenue generated from these industries 

can be critical to fund wildlife 

management programs in range states. 

Tourism, through ecotourism and 

trophy hunting, can provide jobs to 

locals (such as game guards, cooks, 

drivers, security personnel) and often 

brings in revenue for local 

microbusinesses that sell art, jewelry, 

and other native crafts. Lions can 

generate the highest daily rate of any 

mammal hunted (USD $2,650 per day), 

the longest number of days that must be 

booked, and the highest trophy fee 

($24,500) (Jackson 2013, p. 6; Lindsey et 

al. 2012a, p. 5), thus generating 

significant revenue for range countries. 

Creating community-based incentives to 

conserve lions from revenue derived 

from trophy hunting may ameliorate the 

human-lion conflict that arises from 

lions and humans coexisting in the 

same area. 

Revenue from scientifically based 

management programs that include 

trophy hunting can 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). In the past, 

government and private land owners 

were the primary beneficiaries of the 

revenue gained; however, a portion of 

the revenue derived from hunting is 

reportedly now being distributed to 

local communities, creating a value for 

lions that encourages their conservation 

(Barnett & Patterson 2005, p. iv). 

Revenue from trophy hunting is 

purported to create: (1) Incentives for 

countries to conserve large tracts of 

prime habitat; and (2) funding for park 

and reserve management, anti-poaching, 

and security activities. Because habitat 

loss has been identified as one of the 

primary threats to lion populations, it is 

notable that trophy hunting has 

provided lion range states incentives to 

set land aside for hunting throughout 

Africa, and the land set aside exceeds 

the total area of the national parks, 

accounting for approximately half of the 

amount of viable lion habitat 

(Chardonnet et al. 2010, p. 34; Packer et 

al. 2006, pp. 9–10). 

In Tanzania, which is home to 40 

percent of all lions, land set aside for 

sport hunting purposes has resulted in 

an area 5.1 times greater than Tanzania’s 

fully protected and gazetted parks 

(Jackson 2013, p. 6; Barnett & Patterson 

2005, p. 61). In Botswana, despite the 

current ban on lion hunting, the country 

currently has more than 128,000 km2 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 

is in addition to 111,000 km2 (or 19.1 

percent) that has been set aside as 

habitat in the form of National Parks, 

Game Reserves, and Forest Reserves 

(Barnett & Patterson 2005, p. 7). In 2000, 

five countries in southern Africa 

(Botswana, Namibia, South Africa, 

Tanzania, and Zimbabwe) had set aside 

a combined 420,000 km2 of communal 

land, 188,000 km2 of commercial land, 

and 420,089 km2 of state land totaling 

more than 1,028,000 km2 for sport 

hunting purposes (Barnett & Patterson 

2005, p. iii). As a species with a 

considerable range (up to 1,000 km2) 

(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. The 

habitat currently preserved for use in 

sport hunting has helped to reduce the 

impact of habitat loss for the African 

lion, but as discussed previously, 

habitat loss remains a significant threat 

to the species. 

Within its current range, the African 

lion exists in 10 stronghold populations 

containing approximately 24,000 lions 

(70 percent of the current African lion 

population), 19,000 of which are in 

protected areas, and in 7 potential 

stronghold populations containing 

another 4,000 lions. Reports from the 

IUCN Species Survival Commission Cat 

Specialist Group (IUN 2006a, b) 

characterize the population as 

increasing in 3 of those strongholds, as 

stable in 6 of the strongholds, and as 

decreasing in 1 stronghold. Most lion 

populations in protected areas of 

southern and eastern Africa have been 

essentially stable over the last three 

decades (Ray et al. 2005, p. 67). In 

contrast to the stronghold or potential 

stronghold populations, other African 

lion populations, containing a total of 

more than 6,000 individuals, have a 

very high risk of local extinction (Reggio 

et al. 2013, p. 33. During the 2005–2006 

African lion workshops, lion experts 

characterized lion populations in 36 (42 

percent) of the 86 LCUs as decreasing. 

In extensive surveys recently conducted 

within 15 of the 20 LCUs in western and 

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64500 Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules 

central Africa, Henschel et al. (2010, 

entire) were able to confirm lion 

presence in only four. The work of 

Packer et al. (2013) suggests future 

declines within a number of protected 

areas. Craigie et al. (2010, entire) 

provide evidence of declining large 

mammal populations in Africa’s 

protected areas, indicating that 

protected areas in Africa have generally 

failed to mitigate threats to large 

mammal populations, including African 

lion. Although Craigie et al. (2010, p. 

2,225) found large regional differences 

(from large declines in western Africa to 

positive rates of change in southern 

Africa), they found overall populations 

decreased steadily from 1970 to 2005. 

The best available scientific and 

commercial information leads us to 

conclude that the African lion is in 

danger of extinction within the 

foreseeable future throughout all of its 

range. Accordingly, we find that listing 

is warranted and we propose to list it as 

a threatened species throughout its 

range, wherever found. 

Significant Portion of Its Range 

Under the Act and our implementing 

regulations, a species may warrant 

listing if it is endangered or threatened 

throughout all or a significant portion of 

its range. The term ‘‘species’’ includes 

‘‘any subspecies of fish or wildlife or 

plants, and any distinct population 

segment [DPS] of any species of 

vertebrate fish or wildlife which 

interbreeds when mature.’’ We 

published a final policy interpreting the 

phrase ‘‘Significant Portion of its 

Range’’ (SPR) (79 FR 37578, July 1, 

2014). The final policy states that (1) if 

a species is found to be endangered or 

threatened throughout a significant 

portion of its range, the entire species is 

listed as endangered or threatened, 

respectively, and the Act’s protections 

apply to all individuals of the species 

wherever found; (2) a portion of the 

range of a species is ‘‘significant’’ if the 

species is not currently endangered or 

threatened throughout all of its range, 

but the portion’s contribution to the 

viability of the species is so important 

that, without the members in that 

portion, the species would be in danger 

of extinction, or likely to become so in 

the foreseeable future, throughout all of 

its range; (3) the range of a species is 

considered to be the general 

geographical area within which that 

species can be found at the time FWS 

or NMFS makes any particular status 

determination; and (4) if a vertebrate 

species is endangered or threatened 

throughout an SPR, and the population 

in that significant portion is a valid 

DPS, we will list the DPS rather than the 

entire taxonomic species or subspecies. 

We found the African lion to be in 

danger of extinction within the 

foreseeable future throughout all of its 

range. Therefore, no portions of the 

species’ range are ‘‘significant’’ as 

defined in our SPR policy and no 

additional SPR analysis is required. 

Proposed 4(d) Rule 

The purposes of the ESA are to 

provide a means whereby the 

ecosystems upon which endangered 

species and threatened species depend 

may be conserved, to provide a program 

for the conservation of such endangered 

species and threatened species, and to 

take such steps as may be appropriate to 

achieve the purposes of the treaties and 

conventions set forth in the ESA. When 

a species is listed as endangered, certain 

actions are prohibited under section 9 of 

the ESA, as specified in 50 CFR 17.21. 

These include, among others, 

prohibitions on take within the United 

States, within the territorial seas of the 

United States, or upon the high seas; 

import; export; and shipment in 

interstate or foreign commerce in the 

course of a commercial activity. 

The ESA does not specify particular 

prohibitions and exceptions to those 

prohibitions for threatened species. 

Instead, under section 4(d) of the ESA, 

the Secretary, as well as the Secretary of 

Commerce depending on the species, 

was given the discretion to issue such 

regulations as deemed necessary and 

advisable to provide for the 

conservation of such species. The 

Secretary also has the discretion to 

prohibit by regulation with respect to 

any threatened species any act 

prohibited under section 9(a)(1) of the 

ESA. Exercising this discretion, the 

Service has developed general 

prohibitions (50 CFR 17.31) and 

exceptions to those prohibitions (50 

CFR 17.32) under the ESA that apply to 

most threatened species. Under 50 CFR 

17.32, permits may be issued to allow 

persons to engage in otherwise 

prohibited acts for certain purposes. 

Under section 4(d) of the ESA, the 

Secretary, who has delegated this 

authority to the Service, may also 

develop specific prohibitions and 

exceptions tailored to the particular 

conservation needs of a threatened 

species. In such cases, the Service issues 

a 4 (d) rule that may include some or all 

of the prohibitions and authorizations 

set out in 50 CFR 17.31 and 17.32 but 

which also may be more or less 

restrictive than the general provisions at 

50 CFR 17.31 and 17.32. For the African 

lion, the Service has determined that a 

4(d) rule is appropriate. 

We propose to add a 4(d) (special) 

rule for the African lion (Panthera leo 

leo) at 50 CFR 17.40(n). This 4(d) rule 

would maintain all of the prohibitions 

and exceptions codified in 50 CFR 17.31 

and 17.32 and would supersede with 

regard to African lion the import 

exemption found in 50 CFR 17.8 for 

threatened wildlife listed in Appendix II 

of CITES, such that a threatened species 

import permit under 50 CFR 17.32 

would be required for the importation of 

all African lion specimens. Through the 

promulgation of the proposed 4(d) rule, 

the presumption of legality provided 

under Section 9(c)(2) of the Act for the 

otherwise lawful importation of wildlife 

listed in Appendix II of CITES that is 

not an endangered species listed 

pursuant to section 4 of the Act would 

not apply to this subspecies. Thus, 

under the proposed 4(d) rule, all 

otherwise prohibited activities, 

including all imports of African lion 

specimens, would require prior 

authorization or permits under the Act. 

Under our regulations, permits or 

authorization to carry out an otherwise 

prohibited activity could be issued for 

scientific purposes, the enhancement of 

propagation or survival of the species, 

economic hardship, zoological 

exhibitions, educational purposes, or 

special purposes consistent with the 

purposes of the Act. Applications for 

these activities are available from 

http://www.fws.gov/forms/3-200-37.pdf. 

The intent of this proposed 4(d) rule 

is to provide for the conservation of the 

African lion consistent with the 

purposes of the Act. Under the proposed 

4(d) rule, the prohibitions would, in 

part, make it illegal for any person 

subject to the jurisdiction of the United 

States to ‘‘take’’ (includes harass, harm, 

pursue, hunt, shoot, wound, kill, trap, 

capture, or to attempt any of these) 

within the United States or upon the 

high seas; import or export; deliver, 

receive, carry, transport, or ship in 

interstate or foreign commerce, by any 

means whatsoever, in the course of 

commercial activity; or sell or offer for 

sale in interstate or foreign commerce 

any lion specimens. It would also be 

illegal to possess, sell, deliver, carry, 

transport, or ship any such wildlife that 

has been taken in violation of the Act. 

We believe that these protections, 

including the requirement for an import 

permit for all African lion specimens, 

will support and encourage 

conservation actions for the African lion 

and require that permitted activities 

involving lions are carried out in a 

manner that is consistent with the 

purposes of the Act and our 

implementing regulations. 

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Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules 64501 

In connection with this proposed 4(d) 

rule, the Service notes that the African 

lion is listed in Appendix II of CITES, 

and thus can be imported into the U.S. 

pursuant to Section 9(c)(2) of the Act 

and upon presentation of a proper 

CITES export permit from the country of 

origin. Section 9(c)(2) of the Act 

provides that the otherwise lawful 

importation of wildlife that is not an 

endangered species listed pursuant to 

section 4 of the Act, but that is listed in 

Appendix II of CITES, shall be 

presumed to be in compliance with 

provisions of the Act and implementing 

regulations. While there has been 

question as to whether this provision of 

the Act might automatically require 

allowing the importation of a species 

that is both listed as threatened and in 

Appendix II, and preclude the issuance 

of more restrictive 4 (d) rules covering 

importation, the Service has concluded 

that such 4 (d) rules may be issued to 

provide for the conservation of the 

involved species. Section 9(c)(2) does 

not expressly refer to threatened species 

or prevent the issuance of appropriate 4 

(d) rules and could not logically have 

been intended to allow the addition of 

a species to an appendix of an 

international convention to override the 

needs of U.S. law, where there is 

reliable evidence to affect the 

presumption of validity. Finally, the 

term ‘‘presumed’’ implies that the 

established presumption is rebuttable 

under certain circumstances, including 

through the promulgation of a protective 

regulation pursuant to section 4(d) of 

the Act. 

In the case of the African lion, there 

are substantive grounds on which to 

challenge the presumption. For the 

import of sport-hunted trophies, while 

there is evidence that many of the range 

countries are implementing lion 

management plans, we want to 

encourage and support efforts by these 

countries to develop plans that are 

based on sound scientific information. 

As noted, the proposed 4(d) rule for 

African lion would provide for the 

importation into the United States of 

trophies taken legally in range countries 

upon the issuance of a threatened 

species import permit. While the 

Service cannot control hunting of 

foreign species such as African lion, we 

can regulate their importation and 

thereby require that U.S. imports of 

sport-hunted African lion trophy 

specimens are obtained in a manner that 

is consistent with the purposes of the 

Act and the conservation of the 

subspecies in the wild, by allowing 

importation from range countries that 

have management plans that are based 

on scientifically sound data and are 

being implemented to address the 

threats that are facing lions within that 

country. 

Such management plans would be 

expected to address, but are not limited 

to, evaluating population levels and 

trends; the biological needs of the 

species; quotas; management practices; 

legal protection; local community 

involvement; and use of hunting fees for 

conservation. In evaluating these 

factors, we will work closely with the 

range countries and interested parties to 

obtain the best available scientific and 

commercial data. By allowing entry into 

the United States of African lion 

trophies from range countries that have 

scientifically based management plans, 

the range countries would be 

encouraged to adopt and financially 

support the sustainable management of 

lions that benefits both the species and 

local communities. In addition to 

addressing the biological needs of the 

subspecies, a scientifically based 

management plan would provide 

economic incentives for local 

communities to protect and expand 

African lion habitat. 

As stated, anyone wishing to conduct 

any otherwise prohibited activity, such 

as interstate commerce or imports, must 

first obtain a permit under the current 

permitting regulations found at 50 CFR 

13 and 50 CFR 17. As will all permits, 

the individual requesting authorization 

to carry out an otherwise prohibited 

activity under the Act must submit a 

permit application to the Service with 

specific information concerning the 

proposed activity and the benefits/

impacts of the activity on the species. In 

some cases, such as imports of sporthunted trophies, it is not always 

possible for the applicant to provide all 

of the necessary information needed by 

the Service to make a positive 

determination under the Act to 

authorize the activity. For the import of 

sport-hunted trophies, it is typical for 

the Service to consult with the range 

country and other interested parties to 

obtain the necessary information. To 

date, the Service typically has made the 

required findings on sport-hunted 

trophy imports on a country-wide basis, 

although individual import permits are 

issued for each applicant. While the 

Service encourages the submission of 

information from individual applicants, 

we would primarily rely on information 

from other sources when making a 

permitting decision. 

Effects of This Rule 

This rule, if made final, would revise 

50 CFR 17.11(h) to add the African lion 

to the List of Endangered and 

Threatened Wildlife. This rule, if 

adopted, would also establish a 4(d) rule 

for the African lion, which implements 

all of the prohibitions and exceptions 

under 50 CFR 17.31 and 17.32 and 

requires a threatened species import 

permit under 50 CFR 17.32 for the 

importation of all African lion 

specimens. Under the proposed 4(d) 

rule, the import exemption found in 50 

CFR 17.8 for threatened wildlife listed 

in Appendix II of CITES would not 

apply to this subspecies. Through the 

promulgation of the proposed 4(d) rule, 

the presumption of legality provided 

under Section 9(c)(2) of the Act for the 

otherwise lawful importation of wildlife 

listed in Appendix II of CITES that is 

not an endangered species listed 

pursuant to section 4 of the Act would 

not apply to this subspecies. (See: 

Proposed Special Rule section). 

Available Conservation Measures 

Conservation measures provided to 

species listed as endangered or 

threatened under the Act include 

recognition of conservation status, 

requirements for Federal protection, and 

prohibitions against certain practices. 

Recognition through listing encourages 

and results in public awareness and 

conservation actions by Federal and 

State governments in the United States, 

foreign governments, private agencies 

and groups, and individuals. 

Section 7(a) of the Act, as amended, 

and as implemented by regulations at 50 

CFR part 402, requires Federal agencies 

to evaluate their actions that are to be 

conducted within the United States or 

upon the high seas, with respect to any 

species that is proposed to be listed or 

is listed as endangered or threatened. 

Because the African lion is not native to 

the United States, no critical habitat is 

being proposed for designation with this 

rule. Regulations implementing the 

interagency cooperation provision of the 

Act are codified at 50 CFR part 402. 

Section 7(a)(2) of the Act requires 

Federal agencies to ensure that activities 

they authorize, fund, or carry out are not 

likely to jeopardize the continued 

existence of a listed species or to 

destroy or adversely modify its critical 

habitat. If a proposed Federal action 

may adversely affect a listed species, the 

responsible Federal agency must enter 

into formal consultation with the 

Service. Currently, with respect to the 

African lion, no Federal activities are 

known that would require consultation. 

Section 8(a) of the Act authorizes the 

provision of limited financial assistance 

for the development and management of 

programs that the Secretary of the 

Interior determines to be necessary or 

useful for the conservation of 

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64502 Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules 

endangered or threatened species in 

foreign countries. Sections 8(b) and 8(c) 

of the Act authorize the Secretary to 

encourage conservation programs for 

foreign listed species, and to provide 

assistance for such programs, in the 

form of personnel and the training of 

personnel. 

Section 9 of the Act and its 

implementing regulations at 50 CFR 

17.31 set forth a series of general 

prohibitions that apply to all threatened 

wildlife, except where a 4(d) rule 

applies, in which case the 4(d) rule will 

contain all the applicable prohibitions 

and exceptions. If the 4(d) rule is 

adopted as proposed, these prohibitions 

would apply to the African lion. These 

prohibitions, in part, make it illegal for 

any person subject to the jurisdiction of 

the United States to ‘‘take’’ (includes 

harass, harm, pursue, hunt, shoot, 

wound, kill, trap, capture, or to attempt 

any of these) within the United States or 

upon the high seas; import or export; 

deliver, receive, carry, transport, or ship 

in interstate or foreign commerce, by 

any means whatsoever, in the course of 

commercial activity; or sell or offer for 

sale in interstate or foreign commerce 

any lion specimens. It also is illegal to 

possess, sell, deliver, carry, transport, or 

ship any such wildlife that has been 

taken in violation of the Act. Permits 

may be issued to carry out otherwise 

prohibited activities involving 

threatened wildlife species under 

certain circumstances. Certain 

exceptions apply to agents of the 

Service and State conservation agencies. 

Required Determinations 

National Environmental Policy Act (42 

U.S.C. 4321 et seq.) 

We have determined that we do not 

need to prepare an environmental 

assessment, as defined under the 

authority of the National Environmental 

Policy Act of 1969, in connection with 

regulations adopted under section 4(a) 

of the Act. We published a notice 

outlining our reasons for this 

determination in the Federal Register 

on October 25, 1983 (48 FR 49244). 

References Cited 

A list of all references cited in this 

document is available at http://

www.regulations.gov at Docket No. 

FWS–R9–ES–2012–0025, or upon 

request from the U.S. Fish and Wildlife 

Service, Endangered Species Program, 

Branch of Foreign Species (see FOR

FURTHER INFORMATION CONTACT). 

Authors 

The primary authors of this proposed 

rule are staff of the Branch of Foreign 

Species, Ecological Services, U.S. Fish 

and Wildlife Service. 

Proposed Regulation Promulgation 

For the reasons described in the 

preamble, we propose to amend part 17, 

subchapter B of chapter I, title 50 of the 

Code of Federal Regulations, as follows: 

PART 17—[AMENDED] 

■ 1. The authority citation for part 17 

continues to read as follows: 

Authority: 16 U.S.C. 1361–1407; 1531– 

1544; and 4201–4245; unless otherwise 

noted. 

■ 2. In § 17.11(h), the List of Endangered 

and Threatened Wildlife, add an entry 

for ‘‘Lion, African’’ under Mammals to 

read as follows: 

§ 17.11 Endangered and threatened 

wildlife. 

* * * * * 

(h) * * * 

Species 

Historic range 

Vertebrate 

population where 

endangered or 

threatened 

Status When 

listed 

Critical 

habitat 

Special 

rules Common name Scientific name 

MAMMALS

* * * * * * * 

Lion, African ............. Panthera leo leo ..... Africa ...................... Entire ...................... T .................... NA 17.40(n) 

* * * * * * * 

* * * * * 

■ 3. Amend § 17.40 by adding paragraph 

(n) to read as follows: 

§ 17.40 Special rules—mammals. 

* * * * * 

(n) African lion (Panthera leo leo). 

(1) General requirements. All 

prohibitions and provisions of §§ 17.31 

and 17.32 of this part apply to this 

subspecies. 

(2) The import exemption found in 

§ 17.8 of this part for threatened wildlife 

listed in Appendix II of the Convention 

on International Trade in Endangered 

Species of Wild Fauna and Flora 

(CITES) does not apply to this 

subspecies. A threatened species import 

permit under § 17.32 of this part is 

required for the importation of all 

African lion specimens. 

(3) All applicable provisions of 50 

CFR parts 13, 14, 17, and 23 must be 

met. 

* * * * * 

Dated: October 20, 2014. 

Stephen Guertin, 

Director, Fish and Wildlife Service. 

[FR Doc. 2014–25731 Filed 10–28–14; 8:45 am] 

BILLING CODE 4310–55–P 

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