# Endangered and Threatened Wildlife and Plants; Listing the African Lion Subspecies as Threatened With a Rule Under Section 4(d) of the ESA

> Briefs, arguments, decisions, and more.

URL: https://www.frixlaw.com/law-library/documents/fr%3A2014-25731

## Record

- **Collection:** Federal Register
- **Document type:** Proposed Rule
- **Published:** October 29, 2014
- **Citation:** 79 FR 64472

## Text

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

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

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/E-CoP16-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

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 now-extinct north African lions) are more closely related to lions in India than to lions in southern and eastern Africa (Barnett
et al.
2014, pp. 4-8; Dubach
et al.
2013, pp. 741, 746-747, 750-751; Bertola
et al.
2011, entire). According to Dubach
et al.
(2013, p. 753) 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 (km
2
), while IUCN (2006a, p. 12; 2006b, p. 13) estimates lion historical range in sub-Saharan Africa to be 19.3 million km
2
(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 km
2
(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 km
2
(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.

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 km
2
, 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

(km
2
)

Current
range

(km
2
)

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 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).

1
Current range includes occasional and probable range.

2
Bauer
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.

3
Riggio
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.

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 region-wide estimates, tend to rely to a considerable extent on expert opinion or inference (Riggio
et al.
2013, p. 21; Chardonnet 2002, p. 19). Consequently, there is a large degree of uncertainty in these estimates. In addition, to date all efforts to estimate the size of the African

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.

4
Estimates were made for individual Lion Conservation Units (defined management units), and were given as population size classes rather than specific figures. As calculated by Riggio
et al.
(2013, p. 27).

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 2006
a
and IUCN 2006
b

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).

<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 Björklund (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. Björklund (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).

Table 6—Lion Strongholds and Potential Strongholds Identified by Riggio et al. 2013

Lion area
Country

Area

(km
2
)

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.

5
Two lion areas in central Africa make up one potential stronghold.

6
Riggio
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.

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

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 “fission-fusion” social units with a stable membership that sometimes divide into small groups throughout the range. Lions have no fixed breeding season. Females give birth every 20 months if they raise their cubs to maturity, but the interval can be as short as 4-6 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 km
2
in Selous Game Reserve up to 18 per km
2
in protected areas of eastern Africa and South Africa (Creel and Creel 1997, Nowell and Jackson 1996, in Haas
et al.
2005, p. 4). 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 km
2
, but can be considerably larger (Stander 1992b; Van Orsdol
et al.
1985; Viljoen 1993, in Haas
et al.
2005, p. 4). In areas of low or variable prey biomass, annual range requirements for a single lion pride can exceed 1,000 km
2
(Packer
et al.
2013, p. 636). Funston (2011, p. 5) found the home ranges of lion prides in the dune-savannah habitat of Kgalagadi Transfrontier Park to range from 1,762 to 4,532 km
2
.

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

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 km
2
served as a proxy for the extent of land-use conversion that would render habitat unsuitable for lions. Woodroffe (2000, p. 167) analyzed the impact of people on predators by relating local carnivore extinctions to past and projected human population densities and estimated 26 people per km
2
as the mean human density at which lions went locally or regionally extinct. Riggio
et al.
(2013, p. 29) estimate that there were originally approximately 13.5 million km
2
of savannah habitat in Africa. In 1960, 11.9 million km
2
of these habitats had fewer than 25 people per km
2
, and in 2000 this number decreased to 9.7 million km
2
. 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 km
2
(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 sub-Saharan Africa is projected to be large and rapid (UN 2013, p. 9). Although urbanization is increasing in sub-Saharan 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 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), 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

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/km
2
) to 41 million (70/km
2
), whereas Tanzania's grew from 25 million (27/km
2
) to 45 million (48/km
2
) (UN 2013, pp. 421, 798). Not unexpectedly, 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 land-use 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 km
2
, whereas Kenya's population is projected to more than double, reaching a density of 167 people per km
2
(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;

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 conflict-causing 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

(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 km
2
, yet it may be that just under half of the lions that are killed are reported (Republic of Namibia 2013, p. 14). Although 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

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

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 Hluhluwe-iMfolozi 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

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; Pecon-Slattery
et al.
2008, p. 2; Hofmann-Lehmann
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; Pecon-Slattery
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; Pecon-Slattery
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; Hofmann-Lehmann
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).

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).

7
We 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.

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

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 problem-animal control (to reduce human-wildlife conflict), translocation (to expand conservation), culling (reducing population pressures), and local hunting (for protein/meat or employment) (WWF 1997, pp. 8-10).

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 human-lion 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. T

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Source: Frix Law Library, https://www.frixlaw.com/law-library/documents/fr%3A2014-25731. Public record. Not legal advice.