Endangered and Threatened Wildlife and Plants; Listing the African Lion Subspecies as Threatened With a Rule Under Section 4(d) of the ESA, 64471-64502 [2014-25731]
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Vol. 79
Wednesday,
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October 29, 2014
Part II
Department of the Interior
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Fish and Wildlife Service
50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Listing the African Lion
Subspecies as Threatened With a Rule Under Section 4(d) of the ESA;
Proposed Rule
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Federal Register / Vol. 79, No. 209 / Wednesday, October 29, 2014 / Proposed Rules
FOR FURTHER INFORMATION CONTACT:
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
Fish and Wildlife Service,
Interior.
ACTION: Proposed rule and 12-month
finding.
AGENCY:
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.
SUMMARY:
You may submit comments
by one of the following methods:
(1) Electronically: Go to the Federal
eRulemaking Portal: https://
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.
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ADDRESSES:
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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 12month 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:
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(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 https://www.regulations.gov. If you
submit information via https://
www.regulations.gov, your entire
submission—including any personal
identifying information—will be posted
on the Web site. If your submission is
made via a hardcopy that includes
personal identifying information, you
may request at the top of your document
that we withhold this information from
public review. However, we cannot
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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 https://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).
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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
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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 https://
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
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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
https://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; https://
www.cites.org/eng/cop/13/prop/E13P06.pdf). An Appendix-I listing
includes species threatened with
extinction whose trade is permitted only
under exceptional circumstances, which
generally precludes commercial trade.
The import of specimens (both live and
dead, as well as parts and products) of
an Appendix-I species generally
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requires the issuance of both an import
and export permit under CITES. Import
permits are issued only if findings are
made that the import would be for
purposes that are not detrimental to the
survival of the species in the wild and
that the specimen will not be used for
primarily commercial purposes. For live
specimens, a finding must also be made
that the recipient must be suitably
equipped to house and care for the
specimens (CITES Article III(3)). Export
permits are issued only if findings are
made that the specimen was legally
acquired and the export is not
detrimental to the survival of the
species in the wild, and that a living
specimen will be so prepared and
shipped as to minimize the risk of
injury, damage to health, or cruel
treatment. (CITES Article III(2)).
Although Kenya had submitted its
proposal to CoP13 for consideration, it
withdrew its proposal due to the lack of
regional consensus on the proposal.
Furthermore, plans were under way at
that time for convening a regional
workshop on lion management in 2005,
the results of which would be reported
to the CITES Animals Committee
(Animals Committee) (https://
www.cites.org/eng/cop/13/rep/E13ComIRep13.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)] (https://
www.cites.org/eng/dec/valid15/E15Dec.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 (https://
www.cites.org/eng/com/ac/25/sum/E25SumRec.pdf). At CoP16 in March 2013,
the Parties adopted a revised Decision
[Decision 13.93 (Rev. CoP16); https://
www.cites.org/common/cop/16/sum/ECoP16-Plen-06.pdf; https://
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 ........................................................
Sub-Saharan Africa:
Western Africa ......................................................
Central Africa ........................................................
Eastern Africa .......................................................
Southern Africa .....................................................
1 Lions
2 Lions
3 Lions
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
Cameroon, CAR, Chad, Congo, DRC, Gabon, Sudan/South Sudan.
Burundi 2, Djibouti 1, Eritrea 1, Ethiopia, Kenya, Rwanda, Somalia, Sudan/South Sudan,
Tanzania, Uganda.
Angola, Botswana, Lesotho 1, Malawi, Mozambique, Namibia, South Africa, Swaziland,
Zambia, Zimbabwe.
extirpated.
considered occasional or transient by Chardonnet 2002.
considered absent by Henschel et al. 2014.
Species Description
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Algeria 1, Egypt 1, Libya 1, Morocco 1, Tunisia.1
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
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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:
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Characteristics include sharp, retractile
claws, a short neck, a broad face with
prominent whiskers, rounded ears and a
muscular body. Lions are typically a tawny
color with black on the backs of the ears and
white on the abdomen and inner legs. Males
usually have a mane around the head, neck
and chest. Lions are sexually dimorphic,
with males weighing about 20–27 percent
more than females. Adult males, on average,
weigh about 188 kg with the heaviest male
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on record weighing 272 kg. Females are
smaller, weighing, on average, 126 kg. The
male body length, not including the tail,
ranges from 1.7 m to 2.5 m with a tail from
0.9 m to 1 m (Nowell & Jackson, 1996).
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.
Taxonomy
In recent years, several genetic studies
The lion (Panthera leo) was first
have provided evidence of an
described by Linnaeus (1758, in Haas et
evolutionary division within lions in
al. 2005, p. 1), who gave it the name
Africa (see Barnett et al. 2014, p. 6;
Felis leo. It was later placed in the genus
Dubach et al. 2013, p. 746; Bertola et al.
Panthera (Pocock 1930, in Haas et al.
2011 (entire); Antunes et al. 2008
2005, p. 1). Although the classification
(entire); Barnett et al. 2006a, pp. 511–
of the modern lion as Panthera leo is
512). These studies include analysis of
accepted within the scientific
DNA samples from all major regions of
community, there is a lack of consensus the species’ range, though some regions
regarding lion intraspecific taxonomy
are represented by few samples. Results
(Mazak 2010, p. 194; Barnett et. al.
of analysis indicate that a major genetic
2006b, p. 2,120).
subdivision among lions occurs in
Based on morphology, traditional
Africa, with lions in southern and
classifications recognize anywhere from
eastern Africa being genetically distinct
zero subspecies (classifying lions as one
from and more genetically diverse than
monotypic species) up to nine
lions elsewhere (western and central
subspecies (Mazak 2010, p. 194, citing
western and central Africa and Asia).
several sources). The most widely
Evidence indicates that lions in western
referenced of the morphology-based
and central Africa (as well as nowtaxonomies is an eight-subspecies (six
extinct north African lions) are more
extant) classification provided by
closely related to lions in India than to
Hemmer (1974, in Nowell and Jackson
lions in southern and eastern Africa
1996, p. 312; Barnett et al. 2006a, p. 507; (Barnett et al. 2014, pp. 4–8; Dubach et
Barnett et al. 2006b, p. 2,120), which is
al. 2013, pp. 741, 746–747, 750–751;
recognized by the Integrated Taxonomic Bertola et al. 2011, entire). According to
Information System (ITIS) (ITIS 2013,
Dubach et al. (2013, p. 753)
www.itis.gov, accessed June 6, 2013). It
contemporary range collapse and
divides the lion species into: Panthera
fragmentation is too recent a
leo persica (India); P. l. leo, commonly
phenomenon to explain the lower
referred to as the Barbary lion (Morocco genetic variability in these regions.
through Tunisia, extinct); P. l.
Rather, the low genetic diversity in and
senegalensis (West Africa east to the
between western and central African
Central African Republic); P. l. azandica lion populations suggests they have a
(northern Zaire); P. l. bleyenberghi
shorter evolutionary history than the
(southern Zaire and presumably
more genetically diverse lions in
neighboring areas of Zambia and
southern and eastern Africa (Bertola et
Angola); P. l. nubica (East Africa); P. l.
al. 2011, p. 1362). Several authors argue
krugeri (Kalahari region east to the
that the origin of these genetically
Transvaal and Natal regions of South
distinct groups may be the result of
Africa), and P. l. melanochaita, also
regional extinctions and recolonizations
called the Cape lion (Cape region of
during major climate (and consequently
South Africa, extinct) (Nowell and
biome) fluctuations during the
Jackson 1996, p. 312).
Pleistocene Epoch (Barnett et al. 2014,
In 1987, O’Brien (1987a, entire;
pp. 5–8; Bertola et al. 2011, pp. 1,362–
1987b, entire) reported the first results
1,364).
of genetic studies conducted on lion
These genetic studies on lion are
samples from some, but not all, regions
based primarily on analysis of
of the species’ range using early genetic
mitochondrial DNA (mtDNA), which is
techniques. Results indicated that lions
inherited only from the mother. Because
in India differed from lions in Africa,
lions display sex-biased dispersal, in
supporting a two-subspecies
which males leave their natal range and
classification for extant lions: P. leo leo
females tend to remain in their natal
and P. leo persica, the African and
range, one would expect gene flow in
Asian lion, respectively (Ellerman et al. females to be lower than in males,
1953, Meester and Setzer 1971, O’Brien
resulting in greater geographic
et al. 1987, in Dubach 2005, p. 16).
differentiation in females (Mazak 2010,
According to Dubach (2005, p. 16), most p. 204). Consequently, some authors
taxonomic authorities recognize this
state that results of mtDNA analyses
two-subspecies taxonomy. This
should be backed up by studies on
taxonomy is also recognized by the
nuclear DNA (nDNA, inherited from
IUCN (Bauer et al. 2012, unpaginated)
both parents) and morphological traits
and, consequently, by several
before assigning taxonomic importance
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to them (Barnett et al. 2014, pp. 1, 8).
Recently, Mazak (2010, entire)
examined morphological characteristics
of 255 skulls of wild lions and found
considerable variation throughout the
species’ range, with variation being
greater within populations than between
them. However, according to Dubach et
al. (2013, p. 742), the genetic distinction
of lions in southern and eastern Africa
from those elsewhere in the species’
range is confirmed by results of studies
by Antunes et al. (2008, entire) which,
in addition to analysis of mtDNA, also
included analysis of nDNA sequence
and microsatellite variation.
The recent results of genetic research
have renewed debate on lion taxonomy
among the experts. For this reason, the
IUCN Species Survival Commission Cat
Specialist Group has commissioned a
Cat Classification Task Force from
among its expert members to determine
a consensus taxonomy for the group.
Until then, we conclude that the
taxonomy of the species is currently
unresolved. However, as required by the
Act, we base this status review on the
best available scientific and commercial
information, which is the most recent
taxonomy that is the most widely
recognized by taxonomic experts: P. leo
leo (Africa) and P. leo persica (India).
Consequently, in this document we
review the status of the petitioned
entity, the African lion, P. leo leo.
Range
Historically, lions occupied most of
the African continent except the West
African coastal rainforest zone, the
Congo Basin rainforest zone, and the
inner Sahara Desert (Bauer 2003, in Ray
et al. 2005, p. 67; IUCN 2006a, p. 10;
IUCN 2006b, p. 10). Ray et al. (2005, p.
52) estimate lion historical range in
Africa (at about 150 years prior to their
study) to be roughly 22.2 million square
kilometers (km2), while IUCN (2006a, p.
12; 2006b, p. 13) estimates lion
historical range in sub-Saharan Africa to
be 19.3 million km2 (Table 2).
Depending on the study and methods
used, the species’ range is reported to
currently cover between 3.0 million and
5.0 million km2 (Table 2). The most
recent range-wide study was based on a
review of all of the most current
available estimates of lion populations
(up through 2012) (Riggio et al, p. 21),
combined with satellite imagery of
savannah habitat, and provided
estimates of current lion range to be 3.4
million km2 (Riggio et al. 2013, p. 26),
or about 25 percent of the subspecies’
historic range in savannah habitat.
According to Chardonnet (2002, pp. 24–
25), about half the range of the African
lion falls within protected areas.
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The African lion is now believed to be
extirpated from between 75 and 83
percent of its former range (Table 2).
The subspecies has been extirpated from
all of its former range in northern Africa
(Black et al. 2013, p. 1). In addition,
according to IUCN (2006a,b; see Table
2), the species’ range has declined by an
estimated 91 percent in western Africa,
79 percent in central Africa, and 68
percent in eastern/southern Africa
(Table 2), with lion occurrence
unknown in an additional 38 percent of
the historical range (Bauer et al 2008, p.
16). More recently, Henschel et al.
(2014, p. 5) estimate the confirmed lion
range in western Africa, based on data
collected between 2006 and 2012, to be
49,000 km2, or an estimated 1.1 percent
of the species’ former range in the
region.
TABLE 2—ESTIMATES OF THE AFRICAN LION RANGE
Source
Region of Africa
Historic
range
(km2)
Ray et al. 2005: ........................................
Chardonnet 2002: .....................................
Continent-wide ..........................................
Western ....................................................
Central ......................................................
Eastern .....................................................
Southern ...................................................
22,200,000
........................
........................
........................
........................
3,800,000
121,980
651,970
1,137,205
1,039,212
Total ..........................................................
Western ....................................................
Central ......................................................
Western + Central ....................................
Southern + Eastern ..................................
........................
3,814,576
3,392,241
7,206,817
12,080,000
2,950,367
331,749
715,482
1,047,231
3,915,000
Total ..........................................................
Western + Central ....................................
19,286,817
7,206,817
4,962,231
1,047,231
Southern + Eastern ..................................
13,010,000
3,564,000
Total ..........................................................
20,216,817
4,611,231
Western ....................................................
Central ......................................................
Eastern .....................................................
Southern ...................................................
Total ..........................................................
........................
........................
........................
13,500,000
133,784
936,465
780,401
1,540,171
3,390,821
Western ....................................................
........................
49,000
IUCN 2006a, b: 1 .......................................
Bauer et al. 2008: 1 thnsp;2 ....................
Riggio 2013 3 (based on estimates of savannah habitat):.
Henschel et al. 2014: ................................
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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
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.
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recently, during surveys of 21 large
protected areas in western Africa,
Henschel et al. (2014, p. 4) considered
lions to be absent from protected areas
in 5 of these 10 countries (Table 1).
Distribution and Abundance
The general distribution of lions in
Africa is summarized by Ray et al.
(2005, p. 67) as follows:
Lions formerly occupied most of the
African continent except for equatorial forest
and the inner-Sahara. Today, they are extinct
in North Africa and have undergone dramatic
range retraction at the limits of their
historical distribution. Currently, lions are
restricted mainly to protected areas and
surrounding conservancies or ‘game
management areas,’ with the largest
populations in East and southern Africa.
Where protection is poor, particularly
outside protected areas, range loss or
population decreases can be significant.
Declines have been most severe in West and
Central Africa, with only small, isolated
populations scattered chiefly through the
Sahel. Lions in the region are declining in
some protected areas and, with the exception
PO 00000
Frm 00006
Fmt 4701
Sfmt 4702
Current
range
(km2)
Current range as
percent of historic
range
(percent of historic
range w/unknown
lion presence)
17 percent.
9 percent.
21 percent.
15 percent.
32 percent.
26 percent.
15 percent.
(0 percent).
23 percent.
(58 percent).
22 percent.
(38 percent).
25 percent.
1 percent.
of southern Chad and northern Central
African Republic, are virtually absent from
unprotected areas (Bauer 2003).
Estimates of lion abundance on a large
geographical scale are few in number.
For a variety of reasons—including low
densities, large ranges, cryptic
coloration, nocturnal and wary habits—
lions are difficult to count (Bauer et al.
2005, p. 6; Riggio et al. 2013, p. 31).
There are large areas of the species’
range in which no data are available on
lion occurrence or abundance (IUCN
2006b, pp. 12–13). Species experts
recognize that estimating the size of the
African lion population is an ambitious
task, involving many uncertainties
(IUCN 2012, p. 2). Estimates,
particularly range-wide or broad regionwide estimates, tend to rely to a
considerable extent on expert opinion or
inference (Riggio et al. 2013, p. 21;
Chardonnet 2002, p. 19). Consequently,
there is a large degree of uncertainty in
these estimates. In addition, to date all
efforts to estimate the size of the African
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lion population have used different
methods; the results of earlier estimates
cannot be directly compared to those of
later estimates to determine population
trend. The earliest estimates of lion
abundance in Africa were educated
guesses made during the latter half of
the 20th Century. Bauer et al. (2008,
unpaginated) summarize the
information as follows:
There have been few efforts in the past to
estimate the number of lions in Africa. Myers
(1975) wrote, ‘‘Since 1950, their [lion]
numbers may well have been cut in half,
perhaps to as low as 200,000 in all or even
less.’’ Later, Myers (1986) wrote, ‘‘In light of
evidence from all the main countries of its
range, the lion has been undergoing decline
in both range and numbers, often an
accelerating decline, during the past two
decades’’. In the early 1990s, IUCN SSC Cat
Specialist Group members made educated
‘‘guesstimates’’ of 30,000 to 100,000 for the
African Lion population (Nowell and Jackson
1996).
Ferreras and Cousins (1996, entire)
provided the first quantitatively derived
estimate using a GIS-based model
calibrated with information obtained
from lion experts. Ferreras and Cousins
predicted African lion abundance in
1980 to be 75,800. Later, four additional
efforts—Chardonnet (2002), Bauer and
Van Der Merwe (2004), IUCN (2006a,
2006b), and Riggio et al. 2013—
estimated lion population sizes ranging
from 23,000 to 40,000 (Table 3).
Currently, about 90 percent of all
African lions occur in southern and
eastern Africa (Table 3). According to
most studies, most African lions are in
eastern Africa (Table 3). According to
Riggio et al. (2013, p. 27), only nine
countries contain resident populations
of at least 1,000 free-ranging lions
(Central African Republic, Kenya,
Tanzania, Mozambique, Zambia,
Zimbabwe, South Africa, Botswana, and
possibly Angola). Approximately 40
percent of all lions are found in
Tanzania (Riggio et al. 2013, p. 27).
Only about 10 percent of all lions occur
in western and central Africa (Table 3).
According to the most recent survey
effort, numbers in western Africa are
extremely low. Henschel et al. (2014, p.
5) estimate that only 400 lions in the
entire region, with most (about 350, or
88 percent) concentrated in a single
population.
TABLE 3—ESTIMATES OF AFRICAN LION ABUNDANCE
[Rows may not tally due to rounding]
Western
Africa
(percent of total)
Source
Ferreras & Cousins 1996 (estimate for
lion abundance in 1980).
Central
Africa
(percent of total)
.........................
Eastern Africa
(percent of total)
.........................
Chardonnet 2002 ................................
1,163 (3 percent)
2,815 (7 percent)
Bauer & Van Der Merwe 2004 ...........
850 (4 percent) ....
950 (4 percent) ....
IUCN 2006 4 (as calculated by Riggio
et al. 2013).
Riggio 2013 (based on estimates of
savannah habitat).
Henschel et al. 2014 ...........................
1,640 (5 percent)
2,410 (7 percent)
480 (1 percent) ....
2,419 (7 percent)
Southern Africa
(percent of total)
.........................
15,744 (40
cent).
11,000 (48
cent).
17,290 (52
cent).
19,972 (57
cent).
per-
.........................
19,651 (50
cent).
10,000 (44
cent).
11,820 (37
cent).
12,036 (34
cent).
perperper-
Total
per-
75,800 (18,600 in
protected
areas).
39,373.
per-
23,000.
per-
33,160.
per-
34,907.
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).
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TABLE 4—LION CONSERVATION UNITS (LCUS) AS IDENTIFIED AND CHARACTERIZED IN IUCN 2006a AND IUCN 2006b
Western &
Central Africa
Number of LCUs
Total .................................................................................................................................
Estimated to contain:
>500 lions .................................................................................................................
50–500 lions .............................................................................................................
4 Estimates were made for individual Lion
Conservation Units (defined management units),
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and were given as population size classes rather
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Eastern &
Southern
Africa
All regions
(percent)
20
66
86.
2
5
12
28
14 (16 percent).
33 (38 percent).
than specific figures. As calculated by Riggio et al.
(2013, p. 27).
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TABLE 4—LION CONSERVATION UNITS (LCUS) AS IDENTIFIED AND CHARACTERIZED IN IUCN 2006a AND IUCN 2006b—
Continued
Western &
Central Africa
Number of LCUs
<50 lions ...................................................................................................................
Considered:
Viable ........................................................................................................................
Potentially Viable ......................................................................................................
Doubtful Viability .......................................................................................................
With Populations Considered to be:
Increasing .................................................................................................................
Stable ........................................................................................................................
Decreasing ................................................................................................................
Unknown ...................................................................................................................
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
Eastern &
Southern
Africa
All regions
(percent)
13
39 (45 percent).
4
12
4
19
34
13
23 (27 percent).
46 (53 percent).
17 (20 percent).
3
5
12
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 Bjorklund (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. Bjorklund (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
26
5
21
24
16
8 (9 percent).
26 (30 percent).
36 (42 percent).
16 (19 percent).
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
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Number of lion areas
Western
Total .........................................................................
# Estimated to contain:
≥500 lions .........................................................
250–499 lions ...................................................
50–249 lions .....................................................
<50 ....................................................................
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
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Central
Eastern
8
8
28
23
0
1
0
7
1
2
2
3
7
1
12
8
7
2
6
8
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
PO 00000
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(percent)
Southern
Frm 00008
Fmt 4701
Sfmt 4702
67.
15 (22 percent).
6 (9 percent).
20 (30 percent).
26 (39 percent).
4,000 lions. More than 6,000 lions are
located in areas not considered
strongholds or potential strongholds and
have a very high risk of being extirpated
(Riggio et al. 2013, p. 33).
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64479
TABLE 6—LION STRONGHOLDS AND POTENTIAL STRONGHOLDS IDENTIFIED BY RIGGIO ET AL. 2013
Lion area
Area
(km2)
Country
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.
Potential 5 .....
Potential 6 .....
400
1,244
140
148
Stable.
Stable.
500
271
731
3,779
7,644
3,673
880
∼ 500
46
208
2,235
4,953
3,516
820
Central Africa
SE Chad ...............................
E CAR ..................................
Chad .....................................
Central African Republic ......
133,408
328,721
Eastern Africa
Boma-Gambella ...................
Laikipia-Samburu .................
Tarangire ..............................
Ruaha-Rungwa ....................
Selous ..................................
Serengeti-Mara .....................
Tsavo-Mkomazi ....................
Ethiopia, South Sudan .........
Kenya ...................................
Tanzania ..............................
Tanzania ..............................
Tanzania ..............................
Kenya, Tanzania ..................
Kenya, Tanzania ..................
106,941
35,511
28,771
195,993
138,035
35,852
39,216
Potential .......
Potential .......
Potential .......
Stronghold ....
Stronghold ....
Stronghold ....
Stronghold ....
Unknown.
Stable.
Decreasing.
Stable.
Stable.
Increasing.
Decreasing.
Southern Africa
Etosha-Kunene .....................
Kafue ....................................
Great Limpopo .....................
Kgalagadi .............................
Luangwa ...............................
Mid-Zambezi .........................
Niassa ..................................
Okavango-Hwange ...............
Angola, Namibia ...................
Zambia .................................
Mozambique, South Africa,
Zimbabwe.
Botswana, South Africa .......
Malawi, Zambia ....................
Mozambique, Zambia,
Zimbabwe.
Mozambique, Tanzania ........
Botswana, Zimbabwe ...........
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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
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.
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123,800
58,898
150,347
Potential .......
Potential .......
Stronghold ....
455
386
2,311
∼ 315–595
386
2,179
Increasing.
Stable.
Increasing.
163,329
72,992
64,672
Stronghold ....
Stronghold ....
Stronghold ....
800
574
755
∼ 800
574
∼ 350–650
Stable.
Stable.
Stable.
177,559
99,552
Stronghold ....
Stronghold ....
1,573
2,300
1,080
∼ 2,300
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
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Fmt 4701
Sfmt 4702
Increasing.
Stable.
increased or remained stable (see
Distribution and Abundance), and some
have been restored to areas from which
they were previously extirpated (Packer
et al. 2013, p. 636). Reports from the
IUCN Species Survival Commission Cat
Specialist Group (IUN 2006a, b)
characterize the population as
increasing in 3 of the lion strongholds
identified by Riggio et al. (Table 6), as
stable in 6 of the strongholds, and as
decreasing in 1 stronghold. While four
of the lion strongholds or potential
strongholds identified by Riggio et al.
(Table 6) are considered to be
increasing, several African lion
populations, containing a total of more
than 6,000 individuals, have a very high
risk of local extinction (Riggio et al.
2013, p. 33). During the 2005–2006
African lion workshops, lion experts
characterized lion populations in 36 (42
percent) of the 86 LCUs as decreasing.
In extensive surveys recently conducted
within 15 of the 20 LCUs in western and
central Africa, Henschel et al. (2010,
entire) were able to confirm lion
presence in only four. The work of
Packer et al. (2013) suggests future
declines within a number of protected
areas. Craigie et al. (2010, entire)
provide evidence of declining large
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mammal populations in Africa’s
protected areas, indicating that
protected areas in Africa have generally
failed to mitigate threats to large
mammal populations, including African
lion. Although Craigie et al. (2010, p.
2,225) found large regional differences
(from large declines in western Africa to
positive rates of change in southern
Africa), they found overall populations
decreased steadily from 1970 to 2005.
Biology/Ecology
Habitat
Historically, the species occurred in
all habitats in Africa, except rainforest
and the hyper-arid interior of the Sahara
(Ray et al. 2006, p. 66). Today they are
found primarily in savannah, although
there are some remnant populations in
other habitat types (Riggio et al. 2013, p.
19). According to Nowell and Jackson
(1996, p. 19), optimal habitat appears to
be open woodlands and thick bush,
scrub, and grass complexes, where
sufficient cover is provided for hunting
and denning. The highest lion densities
are reached in savannah woodlands
plains mosaics of eastern and southern
Africa (Ray et al. 2005, p. 66). The
species is intolerant of anthropogenic
(human-caused) habitat conversion,
such as farming or overgrazing by
livestock (Ray et al. 2005, p. 66).
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General Biology
Lions are well studied. Much
information exists on African lion
habits, behavior, and ecology. CITES
(2014a, p. 3) provides a general
overview as follows:
Lions are generalist, cooperative hunters,
with foraging preferences changing with
season and with lion group size. Lions live
in groups called ‘‘prides’’, which are ‘‘fissionfusion’’ social units with a stable
membership that sometimes divide into
small groups throughout the range. Lions
have no fixed breeding season. Females give
birth every 20 months if they raise their cubs
to maturity, but the interval can be as short
as 4–6 weeks if their litter is lost. Gestation
lasts 110 days, litter size ranges 1–4 cubs,
and sex ratio at birth is 1:1. At about four
years of age, females will have their first litter
and males will become resident in a pride.
Pride takeovers by male lions and subsequent
infanticide of cubs sired by the ousted male
lions greatly influences reproductive success.
Lionesses defending their cubs from the
victorious males are sometimes killed during
the takeover. Infanticide accounts for 27
percent of cub mortality. Adult mortality is
typically caused by humans, starvation,
disease or attacks from other lions. Injury and
death can also occur during hunting attempts
on some of their larger prey.
Haas et al. (2005, entire) provide a
summary of information on lion,
including the following:
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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,
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Namibia, where Stander (1992) showed that
lions hunting in pairs met their minimum
requirements hunting springboks which, at
<50 kg, are the smallest preferred prey
species recorded.
Prey availability affects the
reproduction, recruitment, and foraging
behavior of lions and, as a result,
strongly influences lion movements,
abundance, and population viability
(Winterbach et al. 2012, p. 7, citing
several sources). Lion densities are
directly dependent on prey biomass
(Van Orsdol et al. 1985, in Packer et al.
2013a, p. 636; Hayward et al. 2007,
entire), and range from 0.08–0.13 adults
and subadults per 100 km2 in Selous
Game Reserve up to 18 per km2 in
protected areas of eastern Africa and
South Africa (Creel and Creel 1997,
Nowell and Jackson 1996, in Haas et al.
2005, p. 4). Aside from human-related
mortality, prey availability is likely the
primary determinant of lion density
(Fuller & Sievert 2001, in Winterbach et
al. 2012, p. 7). In areas of low natural
prey density, or high human contact,
lions may prey on livestock (see
Human-Lion Conflict).
Movements/Home Range
Availability of prey is perhaps the
primary factor that determines the
ranging behavior of large carnivores
(Gittleman & Harvey 1982, Van Orsdol
et al. 1985, Grant et al. 2005, Hayward
et al. 2009, in Winterbach et al. 2012,
p. 4). Home-range sizes of lion prides
correlate with lean-season prey biomass
(Van Orsdol et al. 1985, in Haas et al.
2005, p. 4) and, therefore, vary widely
among habitats. Average range sizes of
African lion prides are 26–226 km2, but
can be considerably larger (Stander
1992b; Van Orsdol et al. 1985; Viljoen
1993, in Haas et al. 2005, p. 4). In areas
of low or variable prey biomass, annual
range requirements for a single lion
pride can exceed 1,000 km2 (Packer et
al. 2013, p. 636). Funston (2011, p. 5)
found the home ranges of lion prides in
the dune-savannah habitat of Kgalagadi
Transfrontier Park to range from 1,762
to 4,532 km2.
Because lion home ranges can be very
large, many protected areas are not large
enough to sustain them (Winterbach et
al. 2014, p. 1; Funston 2011, p. 1, citing
several sources). Where lion ranges
approximate protected area size, lions
roam near or beyond the protected area
boundary, increasing human-lion
contact and human-caused lion
mortality. In these situations, local or
regional extirpation probability is high
due to the population sink created
around the boundary of the protected
area (Davidson et al. 2011, in
Winterbach et al. 2012, p. 5; Funston
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2011, p. 1, citing several sources;
Brashares et al. 2001, entire). This ‘‘edge
effect’’ is a major threat to carnivore
populations inside protected areas
throughout the world (Woodroffe 2001,
in Winterbach et al. 2012, p. 5) (also see
Human-Lion Conflict).
Habitat Loss
Habitat loss and degradation is
reported to be among the main threats
to African lions (IUCN 2006a, p. 18; Ray
et al. 2005, pp. 68–69). The main cause
of lion habitat loss and degradation is
expansion of human settlements and
activities, particularly agriculture and
intensive livestock grazing in lion
habitat (IUCN 2006a, p. 18; IUCN 2006b,
p. 23; Ray et al. 2005, pp. 68–69;
Chardonnet 2002, pp. 103–106).
Expansion of human settlements and
activities into lion habitat renders the
habitat unsuitable for lions primarily
because it results in reduced availability
of the wild prey that lions depend on for
survival (see Loss of Prey Base) and
increased human-lion conflict resulting
in lion mortality (see Human-Lion
Conflict)—two of the main factors that
influence the distribution and
population viability of large carnivores
such as lions (Winterbach et al. 2014, p.
1). Ray et al. (2005, p. 69) note that,
although lions have a wide tolerance for
habitats, they are generally incompatible
with humans and human-caused habitat
alteration and loss. Lions are sensitive
to loss of cover or prey. Riggio et al.
(2013, p. 18) state that dense human
populations and widespread conversion
of land to human use preclude use by
lions.
Habitat destruction and degradation
has been extensive throughout the range
of the African lion, resulting in local
and regional lion population
extirpations, reduced lion densities, a
dramatically reduced subspecies range
(see Range), and small, fragmented, and
isolated lion populations that are
increasingly limited to protected areas
(see Distribution and Abundance) (Ray
et al. 2005, p. 69; Bauer and Van der
Merwe 2004, pp. 29–30; Nowell and
Jackson 1996, pp. 20–21). Lions appear
to have one of the lowest levels of
ecological resilience to human-caused
habitat fragmentation; they are the least
successful large African carnivore
outside conservation areas (Woodroffe
2001, in Winterbach et al. 2012, p. 6).
Large carnivores with low ecological
resilience have a high risk of local
extinction. In order to survive, they
require larger contiguous habitats with
lower negative human impacts than do
more resilient species (Winterbach et al.
2012, p. 5). As human populations
continue to rise in sub-Saharan Africa,
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the amount of land required to meet the
needs of those populations is constantly
increasing (Brink et al. 2014, entire;
Brink and Eva 2009, entire; Eva et al.
2006, p. 4), a problem accentuated by
slow rates of technological progress in
food production and land degradation
from both overuse and natural causes
(United Nations Environment
Programme (UNEP) 2012a, p. 3;
Chardonnet et al. 2010a p. 19;
International Assessment of Agricultural
Knowledge, Science and Technology for
Development (IAASTD) 2009, pp. 3–4,
8; United Nations Economic
Commission for Africa 2008, pp. 3–5).
The result of this process is accelerated
transformation of natural landscapes at
the expense of wilderness that sustains
species such as lions and their prey
(Chardonnet et al. 2010a p. 19). From
1970 to 2000, the human population in
sub-Saharan Africa increased by 126
percent (from 282 million to 639
million) (United Nations (UN) 2013, p.
9), while at about the same time (1975
to 2000), there was a 57 percent increase
in agriculture area (from just over 200
million ha to almost 340 million ha) and
21 percent decrease in natural
vegetation in the region (Brink and Eva
2009, p. 507). In 2009, approximately
1.2 billion ha, or 40 percent, of Africa’s
land area was in permanent pasture or
crops, with the vast majority (31
percent) in pasture (UNEP 2012b, p. 68).
Growing human populations have
been associated with declines in large
carnivore populations all over the
world, and high human density is
strongly associated with local
extirpation of large carnivores (Linnell
et al. 2001, Woodroffe 2001, in
Woodroffe and Frank 2005, p. 91;
Woodroffe 2000, entire). Chardonnet et
al. (2002, p. 103) indicate that the
distribution maps of lion
subpopulations tend to confirm a direct
inverse correlation of lion density and
numbers with human activity and
presence. Further, Packer et al. (2013,
entire) found that lions in unfenced
reserves are highly sensitive to human
population densities in surrounding
communities.
Based on a comparison of land-use
and human population data, Riggio et
al. (2013, p. 23) determined that a
density of 25 or more people per km2
served as a proxy for the extent of landuse conversion that would render
habitat unsuitable for lions. Woodroffe
(2000, p. 167) analyzed the impact of
people on predators by relating local
carnivore extinctions to past and
projected human population densities
and estimated 26 people per km2 as the
mean human density at which lions
went locally or regionally extinct. Riggio
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et al. (2013, p. 29) estimate that there
were originally approximately 13.5
million km2 of savannah habitat in
Africa. In 1960, 11.9 million km2 of
these habitats had fewer than 25 people
per km2, and in 2000 this number
decreased to 9.7 million km2. Based on
analysis of land-use conversion using
satellite imagery and human population
densities, Riggio et al. (2013, p. 29)
found current savannah habitat that is
suitable for lions to be fragmented and
to total about 3.4 million km2 (or 25
percent of African savannah habitat).
These data suggest a substantial
decrease in lion habitat over the past 50
years.
Projections of future human
population growth, area of conversion to
agriculture, and livestock numbers in
Africa suggest suitable lion habitat will
continue to decrease into the foreseeable
future. Africa has the fastest population
growth rate in the world (UNEP 2012a,
p. 2). Future population growth in subSaharan Africa is projected to be large
and rapid (UN 2013, p. 9). Although
urbanization is increasing in subSaharan Africa (UN 2014, p. 20), the
majority of the population is rural, and
about 60–70 percent of the population
relies on agriculture and livestock for
their livelihood (UNEP 2006, pp. 82,
100, 106; IAASTD 2009, p. 2). Much of
the agriculture and livestock-raising is
at subsistence level (IAASTD 2009, pp.
8, 28). As a result, a large portion of the
growing population will depend
directly on expansion of agriculture and
livestock grazing to survive. Between
2010 and 2050 the population of subSaharan Africa is projected to more than
double to more than 2 billion (from 831
million to 2.1 billion) (UN 2013, p. 9).
During about this same time period
(2005 to 2050), Alexandratos and
Bruinsma (2012, p. 107) project the area
of cultivated land to increase by 51
million ha (approximately 21 percent).
However, this figure does not include
range land, and the majority of
agricultural land in Africa is devoted to
grazing (UNEP 2012b, p. 68). The
number of livestock (cattle, sheep, and
goats) in sub-Saharan Africa is projected
to increase about 73 percent, from 688
million to 1.2 billion, by 2050
(Alexandratos and Bruinsma 2012, p.
133).
Expansion of human settlements,
agriculture, and/or livestock grazing are
reported as occurring in or on the
periphery of several of the areas
identified by Riggio et al. (2013, suppl.
1) as lion strongholds (viable
populations) and potential strongholds
(IUCN 2006a, p. 16; IUCN 2006b, pp.
20–22), and are particularly a threat in
western, central, and eastern Africa and
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some parts of southern Africa. There are
only two potential strongholds in
western and central Africa (one in each
region). Expansion of agriculture and
livestock grazing are reported in or
around both (Heschel et al. 2014, pp. 5–
6; Houessou et al. 2013, entire;
Chardonnet et al. 2010, pp. 24–26; IUCN
2008, pp. 8, 28–29), and management of
protected areas in portions of both is
reported as weak (Heschel et al. 2014,
pp. 5–6; IUCN 2008, p. 8). Eastern
Africa contains over half of all the lions
in Africa (Table 3). Seven of the
seventeen African lion strongholds and
potential strongholds identified by
Riggio et al. occur in eastern Africa, and
six of those seven (all four strongholds
and two of three potential strongholds)
are located in Tanzania and Kenya
(Table 6).
Between 1990 and 2010, Kenya’s
human population grew from 23 million
(40/km2) to 41 million (70/km2),
whereas Tanzania’s grew from 25
million (27/km2) to 45 million (48/km2)
(UN 2013, pp. 421, 798). Not
unexpectedly, sources indicate that
expansion of agriculture and livestock
grazing is occurring in these countries
(Brink et al. 2014, entire; UNEP 2009, p.
91; Mesochina et al. 2010, p. 74),
including in or around lion strongholds
and potential strongholds (Ogutu et al.
2011, entire; Mesochina et al. 2010, pp.
71–74, 76; Packer et al. 2010, pp. 8–9;
UNEP 2009, pp. 98–99; Newmark 2008,
pp. 322–324; IUCN 2006b, pp. 20–22;
Ogutu et al. 2005, entire). Mesochina et
al. (2010, p. 74) state that widespread
destruction of wildlife habitat and
human encroachment in wildlife
corridors are major threats to lion
conservation in Tanzania and consider
loss of suitable habitat as a top threat to
lion survival in the country. In Kenya,
the Kenya Wildlife Service (2009, p. 21)
indicates that habitat loss due to landuse changes and human encroachment
into previously wild areas is having a
major impact on lion range size. By
2050 the UN projects the human
population of Tanzania to almost triple
its 2010 population, reaching a density
of 137 people per km2, whereas Kenya’s
population is projected to more than
double, reaching a density of 167 people
per km2 (Table 7).
The human populations of most other
current and recent lion range countries
are also expected to have very high
growth rates (Table 7). It is important to
note that the country-wide human
population densities provided here (and
in Table 7) are not directly comparable
to the density thresholds determined by
Riggio et al. (discussed above) due to the
differences in scale at which they were
made. However, country-wide
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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
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human population in sub-Saharan
Africa is rural, and land supports the
livelihood of most of the population,
loss and degradation of lion habitat can
be expected to accompany the rapid
growth in sub-Saharan Africa’s human
population. Therefore, overall, because
(1) lion prides have vast ranges and the
subspecies requires large areas of
suitable habitat to survive, (2) the
subspecies’ range has already declined
dramatically and is increasingly limited
to protected areas, and (3) habitat loss
and degradation is occurring in or
around several of the remaining lion
strongholds (viable populations) and
potential strongholds, we conclude
based on the best available scientific
and commercial information that the
continued destruction, modification,
and curtailment of lion habitat is likely
to become a significant threat to the
African lion throughout its range.
Human-Lion Conflict
Human-lion conflict and associated
retaliatory killing of lions has played a
major role in the reduction of lion
populations (Lion Guardians 2013, p. 1;
Lion Guardians 2011, p. 2; Hazzah and
Dolrenry 2007, p. 21; Frank et al. 2006,
p. 1; Patterson et al. 2004, p. 508) and
is the greatest threat to remaining lion
populations (Hazzah et al. 2009, p.
2,428; Moghari 2009, p. 31; Kissui 2008,
p. 422; Frank et al. 2006, pp. 1, 3, 10;
Ray et al. 2005 in Hazzah 2006, p. 2;
IUCN 2006b, p. 18). Conflict between
humans and wildlife has been linked to
population declines, reduction in range,
impacts to small population
demographics, and even species
extinctions (Dickman 2013, p. 377; Begg
and Begg 2010, p. 2; Hazzah et al. 2009,
p. 2,428; Moghari 2009, p. 36; Kissui
2008, p. 422; Hazzah 2006, pp. 15, 23,
25).
Human-wildlife conflict stems from
human population growth and the
resulting overlap of humans and
wildlife habitat (Chardonnet et al. 2010,
p. 6; Hazzah 2006, pp. 14, 15). Lion
populations are increasingly restricted
to protected areas, due to human
expansion and associated expansion of
livestock husbandry and agricultural
activities. However, despite being
within protected areas, lions continue to
be impacted by people living on
adjacent land. Villages are established
on the borders of protected areas, cattle
herders enter the protected areas, and
lions move beyond the borders of
protected areas in search of food,
increasing interactions between humans
and lions and the risk of human-lion
conflict (Hazzah et al. 2013, p. 1;
Republic of Namibia 2013, p. 13;
Chardonnet et al. 2010, pp. 11–12;
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Mesochina et al. 2010a, p. 39;
Mesochina et al. 2010b, p. 33; Packer
et al. 2010, pp. 2, 6; Gebresenbet et al.
2009, p. 9; Moghari 2009, pp. 1, 14, 25,
26, 78; Kissui 2008, p. 422; Hazzah
2006, p. 2). The most significant cause
of human-lion conflict is livestock
depredation. Poor husbandry practices
and grazing of livestock within or
adjacent to protected areas increase
exposure of livestock to lions and
increase livestock loss (Uganda Wildlife
Authority 2010, p. 27; Woodroffe and
Frank 2005 in Moghari 2009, p. 35;
Hazzah and Dolrenry 2007, pp. 22–23).
Although lions generally avoid people,
they will occasionally prey on humans,
causing serious injury or death
(Dickman 2013, pp. 380, 384;
Chardonnet et al. 2010, pp. 11, 12, 13;
Moghari 2009, pp. 14, 49, 26, 88; Bauer
et al. 2001 in Moghari 2009, pp. 31, 78,
84; Frank et al. 2006, p. 1; Hazzah 2006,
pp. 14, 17; Patterson et al. 2004, p. 507).
Attacks on humans appears to be more
frequent in southern and eastern Africa
(Chardonnet et al. 2010, pp. 12, 13;
Mesochina et al. 2010a, pp. 29–30;
Frank et al. 2006, pp. 1, 10). Lion attacks
can have various impacts on those
communities that coexist with conflictcausing animals, generating resentment
towards them. When lions cause or are
perceived to cause damage to livestock,
property, or people, the response is
generally to kill them (Dickman 2013,
pp. 378–379; Moghari 2009, p. 25; Frank
et al. 2006, p. 1).
Loss of Prey Base
The lion’s prey base has decreased in
many parts of its range for various
reasons, but a large factor is due to
competition for meat by humans.
Humans in Africa rely on protein
obtained from bushmeat, resulting in
direct competition for prey between
humans and lions, and commercial
poaching of wildlife is becoming a
significant threat to many species,
including those that lions rely upon for
food. Historically, subsistence hunting
with spears was traditionally used to
hunt wildlife, which had minimal
impact to wildlife populations. Spears
have since been replaced by automatic
weaponry (Chardonnet et al. 2010,
p. 27), allowing for poaching of large
numbers of animals for the bushmeat
trade.
The human population in a majority
of African countries within the range of
the lion has quadrupled since the 1960s
(Riggio et al. 2013, p. 29; IUCN 2009,
p. 15), increasing the demand for
bushmeat. Bushmeat comprises between
6 percent (southern Africa) and 55
percent (Central African Republic) of a
human’s diet within the African lion’s
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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
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can be expected, and some lions can
become habitual livestock killers (Frank
et al. 2006, p. 9).
Studies have shown variation in rates
of livestock depredation with regional
rainfall that correlate with prey
availability, including changes in
herding strategies, movement of prey,
and movement of lions (Lion Guardians
2011, p. 6; Moghari 2009, p. 32; Hazzah
2006, pp. 17, 18; Patterson et al. 2004,
p. 514). For example, in some parts of
Zimbabwe, Kenya, and Tanzania,
livestock losses occur during the dry
season. During this time, herders travel
further for forage and water, they use
temporary bomas (a livestock enclosure)
that are typically weak, they are
unfamiliar with carnivore movements in
these new areas, and livestock are weak
due to disease, which makes them more
vulnerable to predator attacks by lions
(Hazzah 2006, p. 17). Additionally,
herders are dependent on resources
within protected areas, and livestock
may be left to wander for days or weeks
during a prolonged drought to find
forage, increasing opportunities for
attacks on livestock by lions
(Chardonnet et al. 2010, p. 24; Frank et
al. 2006, p. 6). In other parts of Kenya,
the Maasai Steppe region of Tanzania,
and Queen Elizabeth National Park,
Uganda, livestock losses were greater
during or following the rainy season
(Moghari 2009, p. 88; Kissui 2008, pp.
427, 428; Frank et al. 2006, p. 6;
Patterson et al. 2004, pp. 510, 514).
Weakened prey and readily available
carcasses provide easy meals during
times of drought, leading to fewer
livestock attacks. However, when rains
return, the abundant grass makes wild
prey harder to catch and lions may turn
to livestock. Migratory prey species,
such as zebra and wildebeest, will move
to other areas for forage and replenished
water sources, leaving lions to turn to
livestock as an alternate food source.
Migratory prey may also move outside
of protected areas. Opportunities for
livestock predation on communal land
increase when lions follow (Packer et al.
2010, p. 9; Kissui 2008, p. 427; Patterson
et al. 2004, p. 514; Frank et al. 2006, p.
6). Similarly, environmental factors
such as vegetative cover, habitat,
climate, seasonality, and prey
availability may affect the rate of attacks
on humans. A certain amount of
vegetative cover is crucial for hunting
success; however, in some cases, the
vegetative cover may make it more
difficult to catch prey, leading to more
attacks on humans. Additionally, dense
cover near settlements allows lions to
hide or stalk humans at a close distance
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protecting crops (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).
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(Mesochina et al. 2010a, p. 39; Moghari
2009, p. 85; 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
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
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rates of lions killed per 100 km2, yet it
may be that just under half of the lions
that are killed are reported (Republic of
Namibia 2013, p. 14). Although most of
the information on human-lion conflict
comes from just a few areas of the lion’s
range (e.g., Kenya, Tanzania, and
Uganda), it is reasonable to conclude
that lions are being killed due to conflict
in all major range countries, due to their
depredation on livestock (Frank et al.
2006, p. 4).
In areas of high conflict, identifying
the responsible animal is often difficult,
and a token animal may be killed
instead (Hazzah 2006, p. 25), leaving the
problem lion to continue to attack and
the potential for additional retaliatory
killings. In Tanzania, game officers kill
numerous lions each year in retaliation
for attacks (Frank et al. 2006, p. 12).
Whereas shooting or spearing target
specific problem animals, poisoning is
indiscriminate and is known to remove
entire prides at once (Frank et al. 2006,
pp. 2, 10, Living with Lions no date,
unpaginated). In the absence of reliable
methods for protecting livestock, rural
people often turn to indiscriminant
methods, like poisoning, to control
livestock depredation. Poisoning is an
easy method for lethal control since it
is readily available, and reinforcing
bomas or more carefully tending
livestock requires time and effort. The
use of Furadan, a widely available and
cheap agricultural pesticide, is
particularly lethal to wildlife and is
increasingly being used to kill predators
in small pastoralist areas of Kenya and
Tanzania. Livestock carcasses are
doused with the poison, killing
predators and scavengers that feed on
them (Frank et al. 2006, pp. 2, 10, Living
with Lions no date, unpaginated).
Poisoning of bush pig carcasses to kill
lions is not uncommon after attacks on
humans. These practices have serious
negative impacts on lion populations
(Frank et al. 2006, p. 9).
Factors That Drive Retaliation
Several anthropogenic factors drive
the level of resentment towards lions
and the extent of retaliatory killing
(Dickman 2013, pp. 379, 385), including
the extent of the loss caused by the
lions, and the wealth and security of the
people affected (Dickman 2013, p. 381;
Mesochina et al. 2010b, p. 54; Moghari
2009, pp. 14, 25; Hazzah 2006, p. 81).
Depending on alternative assets or
incomes, the economic impact of lions
killing livestock can be significant.
Domestic livestock can provide manure,
milk, and meat, and are the basis of
many family incomes, savings, and
social standing; losses can amount to a
large proportion of a subsistence
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herder’s annual income. These losses
are generally uncompensated,
reinforcing negative community
attitudes toward lions and causing
retaliation (Dickman 2013, pp. 380, 381;
Chardonnet et al. 2010, pp. 11, 12, 18,
29; Hazzah et al. 2009, p. 2,428;
Moghari 2009, pp. 14, 25, 27, 36; Kissui
2008, pp. 422–423). Furthermore, a
common perception among local
communities is that lions are conserved
at the cost of community safety and
uncompensated financial losses. When
the people who suffer significant costs
from wildlife feel that the wildlife’s
needs are being put before their own
needs, their frustration can lead to
retaliatory killings (Dickman 2013, p.
382). This situation further contributes
to negative attitudes toward lion
conservation programs (Moghari 2009,
p. 37).
Lions are particularly vulnerable to
retributive killing because they are often
driven by a perceived level of lion
predation on livestock rather than actual
levels of conflict. In some locations,
other predators (e.g., baboons (Papio
ursinus), spotted hyenas (Crocuta
crocuta), and leopards (Panthera
pardus)) as well as disease are
responsible for the majority of livestock
losses and human casualties, yet it is
lions that are sought and killed more
often. Negative perceptions of lions may
be based on an over-estimated number
of lions in a community or protected
area and an over-estimated number of
human-lion conflicts (Dickman 2013, p.
380; Begg and Begg 2010, p. 20;
Chardonnet et al. 2010, pp. 12, 21–22;
Hazzah et al. 2009, p. 2,436; Maclennan
et al. 2009 in Hazzah et al. 2009, p.
2,429; Moghari 2009, pp. 77–78, 107,
150; Holmern et al. 2007 in Moghari
2009, p. 34; Butler 2001 in Moghari
2009, p. 34; Kissui 2008, pp. 426, 428,
429; Hazzah 2006, pp. 18–19, 83–85, 96,
98, 107, 111; Patterson et al. 2004, pp.
514, 515). One cause for the
disproportionate blame put on lions is
that the lion is a highly visible species.
It is a large-bodied species that lives in
groups and has cultural significance.
Because of its physical presence, there
is often a ‘‘hyper-awareness’’ of the
potential risk for lion attacks and lions
may be blamed simply because they
have been seen in an area (Dickman
2013, pp. 380–381).
Cultural beliefs and traditions can
have a negative impact on lions.
Because cattle are of great cultural
significance to Maasai, their loss can
impose social or cultural costs and
incite greater resentment and higher
levels of retributive killing (Dickman
2013, p. 384; Kissui 2008, p. 429;
Hazzah 2006, p. 99). In some areas of
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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
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sustainable. Attacks on humans can
impact long-term viability for lions as
people who fear for their lives or safety
are unlikely to support conservation
actions and are more likely to retaliate
by killing any lions found near
settlements (Frank et al. 2006, p. 12).
Every year, human-lion conflicts
intensify due to habitat loss, poor
livestock management, and decreased
availability of wild prey, further
increasing the likelihood that the
subspecies will be at risk of extinction
within the foreseeable future (Lion
Guardians 2013, p. 1).
Human population growth within the
lion’s range is projected to be 2.1 billion
by 2050 (UN 2012, p. 2). The number of
livestock within the lion’s range is
projected to increase by about 73
percent by 2050 (Food and Agriculture
Organization of the United Nations
2012, p. 133). Given this expected
increase in humans and livestock by
2050, we conclude the conditions
described above will continue to worsen
to the point that African lions will likely
be at risk of extinction within the
foreseeable future. As livestock numbers
increase, expansion of agricultural and
pastoral practices continue, and the
lion’s prey base is hunted at
unsustainable levels to meet a growing
demand for food, livestock depredation
and retributive killing of lions will
likely increase (Dickman 2013, p. 379;
Hoppe-Dominik et al. 2011, p. 452;
Chardonnet et al. 2010, p. 19;
Gebresenbet et al. 2009, p. 9; Hazzah
and Dolrenry 2007, p. 3). Furthermore,
as the need for grazing land becomes
more critical, expansion of livestock
numbers may be partially supported by
the network of protected areas, seen by
herders as unused pastures (Chardonnet
et al. 2010, p. 25).
Retaliatory killing of lions continue in
many areas and this practice impacts
the viability of lion populations
throughout its range. The killing of lions
due to human-lion conflict is enough to
result in the local extirpation of lion
populations, though at present does not
place the subspecies in danger of
extinction. Human-lion conflict is
exacerbated by an increasing human
population, the expansion of human
settlements, loss of prey base due to the
bushmeat trade and expanding
agriculture, as well as increasing
pressures on natural resources to meet
the needs of the growing human
population. We expect retaliatory
killings due to human-lion conflict to
continue to increase into the foreseeable
future. We conclude based on the best
available scientific and commercial
information that the continuation of this
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activity is a significant threat to the
African lion throughout its range.
Disease
Wild lions are known to be infected
with various pathogens (Hunter et al.
2012, p. 2; Craft 2008, p. 6; Michel et al.
2006, p. 92; Hofmann-Lehmann et al.
1996, pp. 559–561). The human
population within the range of the lion
is expanding into lion habitat,
increasing the exposure of lions to
diseases from domestic animals (IUCN
2006b, p. 26). Because lions are a top
predator, they are at a particularly high
risk of exposure to pathogens (Keet et al.
2009, p. 11). Some pathogens are
endemic, meaning they are constantly
present, but often do not cause disease.
Others are epidemic and cause a sudden
severe outbreak with the potential to
cause high mortality (Craft 2008, pp. 5,
6). Although lions are known to be
infected with certain pathogens,
information on the extent of the
subspecies’ infections and impacts of
these diseases on lion populations is
limited, because few long-term studies
have been conducted; for example,
those lion populations found in
Serengeti National Park, Ngorongoro
Crater, and Kruger National Park.
Feline calicivirus, feline herpesvirus,
feline parvovirus, feline coronavirus,
and feline leukemia virus are endemic
viruses known to occur in lions of
Serengeti National Park, Ngorongoro
Crater, Lake Manyara National Park,
Kruger National Park, and Etosha
National Park (but not all viruses are
known in all parks). However, these
diseases are not known to affect lion
survival (Hunter et al. 2012, p. 2; Craft
2008, p. 6; Hofmann-Lehmann 1996, pp.
559, 561).
Lions within Kruger National Park
and Hluhluwe-iMfolozi Park, South
Africa, and Serengeti National Park,
Tanzania, are known to be infected with
Mycobacterium bovis, a pathogen that
causes bovine tuberculosis (bTB). This
pathogen is not endemic to African
wildlife and was likely introduced from
cattle imported from Europe. M. bovis is
transmitted to ungulates, such as
African buffalo (Syncerus caffer) and
wildebeest (Connochaetes taurinus)
from domestic cattle located on the
periphery of the parks (Maas et al. 2012,
p. 4,206; Keet et al. 2009, pp. 4, 11;
Renwick et al. 2007, p. 532; Michel et
al. 2006, pp. 92, 93; Cleaveland et al.
2005, pp. 446, 449, 450). Spillover of the
disease from buffalo to other lion prey
species, such as kudu (Tragelaphus
strepsiceros) and warthog
(Phacochoerus africanus), have also
been documented (Keet et al. 2009, pp.
4, 11; Renwick et al. 2007, p. 535;
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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
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they experience another bTB infection,
suffer from poor nutrition or advancing
age, or become super-infected with
other diseases that may exacerbate the
infection (Renwick et al. 2007, p. 533).
The impact of bTB on lions is largely
unknown. Researchers suggest that bTB
may lower breeding success, reduce
resiliency, and may be a mortality factor
based on data that indicate survival is
shortened in infected lions, with death
ranging between 2 and 5 years after
infection (Maas et al. 2012, p. 4,212;
Renwick et al. 2007, p. 536; Michel et
al. 2006, p. 93; Cleaveland et al. 2005,
pp. 450, 451). Thirty percent of the
inbred populations in HluhluweiMfolozi Park died due to a combination
of bTB and malnutrition (Hunter et al.
2012, p. 3). A study from Kruger
National Park indicated that bTB
spreads quickly through lion
populations; in an area with high herd
prevalence of M. bovis, 90 percent of
lions became infected (Cleaveland et al.
2005, p. 451). However, despite bTB
infection and a high prevalence in prey
species, the lion population in Kruger
National Park has remained stable
(Ferreira and Funston 2010, p. 201).
Epidemics of canine distemper virus
(CDV) are known to have occurred in
the Serengeti-Mara Ecosystem, an area
that encompasses the Serengeti National
Park, Ngorongoro Conservation Area,
and Maasai Mara National Reserve
(Craft 2008, pp. 13–14; Cleaveland et al.
2007, pp. 613, 616, 618). CDV is a
common pathogen in the large
population of domestic dogs around the
Serengeti-Mara Ecosystem, which are
believed to be the source of CDV
(Cleaveland et al. 2007, pp. 613, 617).
CDV is assumed to be transferred to
lions by the sharing of food sources with
spotted hyenas (Crocuta crocuta) or
jackals (Canis spp.) that become
infected by consuming the infected
carcasses of domestic dogs (Canis lupus
familiaris). Lions may also transmit
CDV among themselves via sharing
food, fights, and mating (Craft et al.
2009, pp. 1,778, 1,783; Craft 2008, pp.
13, 18, 71).
CDV generally lacks clinical signs or
measurable mortality in lions, and most
CDV events have been harmless.
However, in 1994 and 2001, CDV
epidemics in the Serengeti National
Park/Maasai Mara National Reserve and
Ngorongoro Crater, respectively,
resulted in unusually high mortality
rates (Hunter et al. 2012, p. 2; Craft
2008, p. 14; Munson et al. 2008, pp. 1,
2; Cleaveland et al. 2007, pp. 613, 618;
Roelke-Parker et al. 1996, pp. 441, 443).
These outbreaks coincided with climate
extremes that resulted in a higher
number of Babesia, a tick-borne
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parasite, infections (Munson et al. 2008,
pp. 2, 5). Babesia is common in lions,
but typically at low levels with no
measurable impacts on their health
(Craft 2008, p. 14; Munson et al. 2008,
p. 3). However, droughts in 1993 and
2000 in Serengeti National Park/Maasai
Mara National Reserve and Ngorongoro
Crater, respectively, led to large-scale
starvation and widespread die-offs of
buffalo. This situation combined with
resumption of rains and fire suppression
in Ngorongoro Crater favored
propagation of ticks, vectors of Babesia,
leading to unusually high tick burdens.
The compromised health of buffalo
allowed lions to feed on an inordinate
number of tick-infested prey (Craft 2008,
p. 14; Munson et al. 2008, pp. 2, 4, 5).
Exposure to either CDV or Babesia
singly is not typically associated with a
compromise in health or an increase in
mortality (Craft 2008, p. 14; Munson et
al. 2008, pp. 1, 2, 3). However, the
Babesia infections were exacerbated by
the immunosuppressive effects of CDV
and led to the unusually high mortality
rates (Craft 2008, p. 14; Munson et al.
2008, p. 5). The Serengeti National Park/
Maasai Mara National Reserve lion
population lost 30 percent of its
population (approximately 1,000 lions),
but has recovered to its pre-epidemic
population levels (Craft 2008, pp. v, 14,
41; Munson et al. 2008, p. 1; Cleaveland
et al. 2007, pp. 613, 617; Roelke-Parker
et al. 1996, p. 444). Thirty-four percent
of the Ngorongoro Crater lion
population was killed, but frequent
outbreaks of disease have prevented this
population from recovering back to its
carrying capacity (Craft 2008, p. 14;
Munson et al. 2008, pp. 1, 2; Cleaveland
et al. 2007, p. 617). The difference in
recovery is likely due to the highly
inbred nature of the Ngorongoro Crater
lion population, compared to the
Serengeti population, and its greater
susceptibility to parasitic and viral
infections (Hunter et al. 2012, p. 2;
Munson et al. 2008, p. 5; Brown et al.
1994, pp. 5,953–5,954).
Feline immunodeficiency virus (FIV)
is an endemic pathogen in many lion
populations of southern and eastern
Africa (Maas et al. 2012, p. 4,206;
Adams et al. 2011, p. 173; PeconSlattery et al. 2008, p. 2; HofmannLehmann et al. 1996, pp. 555, 558;
Brown et al. 1994, p. 5,966). FIV is
believed to have been present in lions
since the late Pliocene (O’Brien et al.
2012, p. 243; Troyer et al. 2011, p. 2;
Roelke et al. 2009, p. 3; Pecon-Slattery
et al. 2008, p. 8). There are 6 subtypes
of FIV, A through F, each with a distinct
geographic area of endemnicity (Adams
et al. 2011, p. 174; Troyer et al. 2011,
p. 2; Roelke et al. 2009, p. 3; Pecon-
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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; PeconSlattery et al. 2008, p. 5; Brown et al.
1994, p. 5,953). Prevalence of FIV in
infected lion populations is high, often
approaching 100 percent of adults
(O’Brien et al. 2012, p. 243; Troyer et al.
2011, p. 2; Roelke et al. 2009, p. 3;
O’Brien et al. 2006, p. 262; HofmannLehmann et al. 1996, p. 559).
FIV causes immune deficiencies that
allow for opportunistic infections in the
host (Brown et al. 1994, p. 5,953).
Chronic effects of FIV are important to
long-term survival and differ according
to subtype (Troyer et al. 2011, p. 6).
Studies have indicated that lions may
exhibit signs of opportunistic infection
associated with AIDS, such as swollen
lymph nodes, gingivitis, tongue
papillomas, dehydration, poor coat
condition, and abnormal red blood cell
parameters, and in some cases death
(Troyer et al. 2011, p. 2; Roelke et al.
2009, pp. 2, 3–6). Lions in Botswana
and Tanzania have demonstrated
multiple clinical features of chronic
immune depletion similar to HIV and
domestic cat AIDS (Troyer et al. 2011,
pp. 2–3). However, there is no evidence
that it poses a threat to wild populations
(Frank et al. 2006, p. 1); FIV does not
appear to be impacting lions in Kruger
National Park (Maas et al. 2012, p.
4,212), and no evidence of AIDS-like
illnesses or decreased lifespan has been
found in FIV lion populations in the
Serengeti (O’Brien et al. 2006, p. 263).
Infection with a single disease does
not appear to have detrimental impacts
on lions, although general body
condition, health, and lifespan may be
compromised. Co-infections, however,
could have synergistic effects that lead
to greater impacts on lions than a single
infection. Lions impacted by the 1994
CDV outbreak in Serengeti National
Park/Maasai Mara National Reserve may
have been more susceptible to CDV due
to depleted immunity caused by FIV
(O’Brien et al. 2006, p. 263). Troyer et
al. (2011, pp. 5–6) found that survival
during the CDV/Babesia outbreak in
Serengeti National Park/Maasai Mara
National Reserve was significantly less
for lions infected with FIV A and/or C
than FIV B. This finding suggests that
FIV A and C may predispose carriers to
CDV pathogenesis and may increase the
risk of mortality (O’Brien et al. 2012, p.
243). Additionally, certain
environmental conditions may
exacerbate the effects of an otherwise
innocuous infection. For example, as
discussed above, CDV and Babesia
infections generally have no measurable
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impacts on lion health, but climatic
conditions increased exposure of lions
to Babesia infections, which were
exacerbated by the immunosuppressive
effects of CDV and led to unusually high
mortality rates. Furthermore, species
with reduced genetic variation may be
less able to mount an effective immune
response against an emerging pathogen
(O’Brien et al. 2006, p. 255). Some lions
infected with bTB may remain
asymptomatic until conditions change
and they suffer from poor nutrition due
to low prey density, advancing age, or
become super-infected with other
diseases that may exacerbate the
infection (Renwick et al. 2007, p. 533).
Impacts of coinfections of FIV with
FCV, FPV, FHV, and FCoV on
individual lions are negligible and do
not endanger the lion population, at
least in the absence of other aggravating
cofactors (Hofmann-Lehmann et al.
1996, p. 561). Pathogen–pathogen
interactions may become more
important when lions are under
additional stress (e.g., increased parasite
load or low prey density) (Maas et al.
2012, p. 4,212).
Although disease is known in several
populations, the impacts are known in
only a couple of populations where
disease has been frequently studied.
Disease can be a factor in the decline of
lions when combined with other factors,
including environmental changes,
reduced prey density, and inbreeding
depression. However, this type of
impact has been observed in some small
populations that are at a higher risk, but
has not been observed at the species
population level. Therefore, we
conclude, based on the best scientific
and commercial information available,
that disease is not a significant threat to
the species.
Deleterious Effects Due to Small
Population Sizes
The risk of extinction is related to the
moment when a declining population
becomes a small population and is often
estimated using minimum viable
population (MVP) sizes (Traill et al.
2010, p. 28). The viability of a lion
population is complex, but it partly
depends on the number of prides and
ability of males to disperse and interact
with other prides, which affects
exchange of genetic material (Bjorklund
2003, p. 518). Without genetic exchange,
or variation, individual fitness is
reduced and species are less able to
adapt to environmental changes and
stress, increasing the risk of extinction
(Bijlsma and Loeschcke 2012, pp. 117,
119; Segelbacher et al. 2010, p. 2; Traill
et al. 2010, p. 31; Bjorklund 2003, p.
515).
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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
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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
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hunting quota, the country has halted
authorization of trophy hunting
pursuant to that quota until some later
date or until some further action is
taken, as prescribed by that country.
Therefore, you will see us refer to
countries like Zambia and Botswana,
each of which has hunting quotas and
bans in place. Trophy hunting is
currently banned in 12 countries:
Angola, Botswana, Cameroon,7 Congo,
Gabon, Ghana, Kenya, Malawi,
Mauritania, Niger, Nigeria, and Rwanda
(CITES 2014a, p.14; Lindsey et al.
2013a, entire; Lindsey 2013, pers.
comm.; Jackson 2013, pp. 7–8).
Botswana banned lion hunting between
2001 and 2004, and then again from
2008 to the present (Davison et al. 2011,
p. 114). Kenya banned all sport hunting
in 1977 (African Wildlife Foundation
1998, p. 3). Trophy hunting is restricted
to problem or dangerous animals in
Ethiopia and Uganda (Lindsey 2008, p.
42). Zambia banned all sport hunting in
January of 2013; while restrictions were
lifted from other trophy species in
August 2014, the ban on lions and
leopards remains in place (ABC News
2014, unpaginated; Flocken 2013,
unpaginated). In 2011, researchers in
Cameroon suggested that there should
be an immediate moratorium of at least
5 years on the hunting of lions in
Cameroon, during which lions are
allowed to recover and a management
plan for lion hunting is established
(Croes et al. 2011).
Quotas
A scientifically based ‘‘quota’’ is the
maximum number of a given species
that can be removed from a specific
population without damaging the
biological integrity and sustainability of
that population (World Wildlife Fund
(WWF) 1997, p. 9). For a quota to be
scientifically based, it must be based
upon available monitoring data of the
species. Although varying by country
and by economic resources, monitoring
data used to determine quotas have
included, but are not limited to, past
hunting off-take records, trophy quality
data, ground transect surveys, wildlife
ranger and safari operator input, the
species’ reproductive biology, and aerial
population census data, although
usually aerial data is limited to species
that can be easily observed from the air,
such as elephants and buffalo (Barnett &
Patterson 2005, p. 102). Generally, the
conservation principle behind
scientifically based quotas is to limit
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.
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offtake of the species to either equal or
slightly lower than the growth rate of
the target specimens (e.g., males vs.
female), provided the offtake does not
damage the integrity and sustainability
of that population.
In order for scientifically based quotas
to result in offtake less than the growth
rate of target specimens, many factors
are evaluated including the species’
biological factors (reproductive rate,
gender, age, and behavior), as well as
community and client objectives (WWF
1997, pp. 14–19). Each quota should be
then assigned to a geographical area
and/or population based on this
information. Thus, for lions, a
scientifically based quota defines the
specific number of lions that can be
removed from a specific geographical
area and population, for any purpose,
within a particular year. Scientifically
based quotas do not apply solely to
sport hunting, but set the limits for all
offtake for a particular year; other
potential offtake includes problemanimal control (to reduce humanwildlife conflict), translocation (to
expand conservation), culling (reducing
population pressures), and local hunting
(for protein/meat or employment) (WWF
1997, pp. 8–10).
While each of these uses offers
advantages and disadvantages, quotas
are typically utilized only for sport
hunting, as it may provide the highest
all-around benefits to local
communities. For example, a portion of
a quota could be used to kill a problem
animal; the benefits to the community
would then include the use of the
animal parts for meat or trade and it
would theoretically reduce the conflict.
However, this provides a more limited
economic benefit to the community than
would selling the same quota for trophy
hunting, which could potentially
eliminate the problem animal, provide
meat and parts for trade, and provide
revenue for the community (WWF 1997,
pp. 31–33).
There are two primary types of
quotas, ‘‘fixed’’ and ‘‘optional.’’ Trophy
fees for ‘‘optional’’ quotas are paid only
when the lion is shot, whereas, ‘‘fixed’’
quotas require the payment of a portion
(40–100 percent) of the lion trophy fee,
regardless of whether the hunt is
successful. Until 1999, male lions were
typically on ‘‘fixed’’ quotas, whereas
female lions were under ‘‘optional’’
quotas. Due to this approach, trophies
collected in the 1990’s were often of
lower quality, younger, less desirable
male lions, as operators and hunters had
no incentive to be selective (e.g. the
hunter had already paid for it).
Therefore, current recommendation for
all quotas is to be the ‘‘optional’’ type
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(Lindsey et al. 2013a, p. 9; Packer et al.
2006, pp. 5, 9).
Two primary concerns have been
raised by the scientific and international
community with regards to current lion
quotas. Specifically, that existing quotas
are set above sustainable levels and the
data used for setting quotas is
inconsistent and not scientifically based
(Hunter et al. 2013, unpaginated;
Lindsey et al. 2006, p. 284). For
example, recent quotas appear rarely to
address safeguards for sustainability or
establish a systematic approach to
setting lion quotas (Hunter et al. 2013,
p. 2; Lindsey et al. 2013b, p. 8).
Additionally, it has been noted that
previous quotas in Namibia,
Mozambique, and Zimbabwe may have
been influenced by human-lion conflict,
with higher quotas being allocated to
locations with reportedly higher humanlion conflict levels (Lindsey et al.
2013b, p. 4). Apparently, in recognition
of these inconsistencies, range countries
and conservationists have been working
to establish a set of best practices in
order to create a more consistent,
scientifically based approach to
determining quotas. The recommended
best practices include: (1) Establishing
processes and procedures that are
clearly outlined, transparent, and
accountable; (2) establishing processes
and procedures that are CITES
compliant;(3) demonstrating
management capacity; (4) standardizing
information sources; (5) establishing
monitoring systems for critical data; (6)
recording and analyzing trophy hunting
data; (7) conducting data collection and
analysis for each hunting block and
concession; and (8) establishing a
primary body who will approve quotas
(Burnett and Patterson 2005, p. 103). We
have no information on whether these
best practices have been implemented
by the lion range states. However, most
countries that allow trophy hunting of
lions appear to be reviewing their
trophy hunting practices (Jackson 2013,
pp. 2–3; White 2013, pp. 12–13). Benin
halved their quotas in 2002 after the
first population census of lions was
conducted and resulted in the current
quota of six lions every 2 years in
Pendjari and four lions every 2 years in
western Benin or one lion annually in
each of the five hunting zones. This was
largely due to impacts to lions from
habitat degradation and fragmentation
(particularly exacerbated by the increase
of human population), loss of prey by
poaching, trade (both legal and illegal),
and human-lion conflict. (CITES 2014a,
p. 5; Sogbohossou 2014, p. 1).
Throughout the countries in Africa,
most appear to have reduced their
offtake considerably since the 1990’s.
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According to Packer et al. (2006, pp. 2–
3), regardless of population estimates,
countries are allowing for only a small
proportion of their lion populations to
be hunted, with most countries ranging
from 2–4 percent annually (excluding
offtake from South Africa, where offtake
has been increasing from the trophy
hunting of primarily captive-born lions,
and Zimbabwe, where offtake was 2–3
percent higher than other countries from
1998–2004.
Regardless of these reductions, many
stakeholders consider the quota system
to be outdated and ineffective because it
does not address the biological and
social impacts of trophy hunting on lion
prides. Opponents also state that trophy
hunting affects the social structure of
the pride and results in increased
infanticide of lion cubs. This
supposition is inconclusive and not
well supported (CITES 2014a, p. 14;
Dagg 2000, pp. 831–835) (See
Infanticide and Age-based Hunting
Strategies). Regardless, since 2006,
researchers have recommended the
implementation of age-based hunting
strategies; these are discussed below
(Packer et al. 2006, pp. 6–8).
Five countries maintain quotas to
allow for approximately 6–15 lion
trophies to be taken per year: Benin,
Burkina Faso, Cameroon,7 Mozambique,
and Namibia. Tanzania allows the take
of approximately 50 lions annually, and
Zimbabwe allows approximately 70
animals annually to be taken (Jackson
2013 pp. 7–8, CITES WCMC–UNEP
trade database, accessed December
2013). In Ethiopia and Uganda, trophy
hunting is restricted to problem or
dangerous animals only (Lindsey 2008,
p. 42), and Botswana and Zambia
currently ban all trophy hunting (CITES
2014a, p.14). South Africa has not set a
quota for the take of wild lions since 99
percent of the trophy-hunted lions are
reportedly not of wild origin, but
captive-born (Hunter et al. 2013, p. 2;
RSA 2013, pp. 5, 7).
Below is a summary of estimated
annual hunting quotas for the African
lion:
TABLE 7—ANNUAL TROPHY QUOTAS
(APPROXIMATE) AS OF 2013
Country
Benin ...................................
Botswana (moratorium) ......
Burkina Faso ......................
Cameroon 7 .........................
Mozambique .......................
Namibia ...............................
Tanzania (as of 2012) ........
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Annual lion
trophy quotas
(Jackson 2013,
pp. 7–8)
6
30
6
6
15
10
50
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TABLE 7—ANNUAL TROPHY QUOTAS using the CITES Trade Database
(APPROXIMATE) AS OF 2013—Con- indicates that the outputs produced by
the CITES Trade Database can be easily
tinued
Country
Annual lion
trophy quotas
(Jackson 2013,
pp. 7–8)
Zambia (moratorium) ..........
Zimbabwe ...........................
50
70
Import/Export of Lion Trophies
Although each country has its own
method of regulating trophy hunting,
international trade of lion trophies must
adhere to CITES (see Conservation
Status). International trade of lion parts
and products (including trophies) are
reported by both the exporting and
importing countries and tracked by the
United Nations Environment
Programme World Conservation
Monitoring Centre (UNEP–WCMC). The
international trade data on the African
lion that has been compiled in the
CITES UNEP–WCMC Trade Database is
extensive. Therefore, it is likely that the
actual numbers of African lion parts and
products in international trade is
slightly smaller than what we have
reported using the UNEP–WCMC ‘‘gross
exports’’ report (CITES lion gross
exports, https://trade.cites.org, accessed
April 23, 2014).
In 2012, the most recent year for
which CITES trade data are available,
U.S. CITES Annual Report trade data
indicated that the United States allowed
the direct import of African lion
trophies from eight African countries, as
follows:
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Central African Republic = 1 trophy
Ethiopia = 1 trophy
Mozambique = 5 trophies
Namibia = 5 trophies
South Africa = 413 trophies (the majority of
which are reported to be of captive-born
origin)
Tanzania = 42 trophies
Zambia = 32 trophies
Zimbabwe = 49 trophies
According to the CITES UNEP–
WCMC database, between 2005 and
2012, exports of lion trophies have
demonstrated a decreasing trend when
exports of captive-born lions from South
Africa are excluded (CITES lion gross
exports, https://trade.cites.org, accessed
April 23, 2014). For example, in 2005
there were 874 lion trophy exports
reported in UNEP–WCMC, 521 if South
Africa were excluded; whereas in 2012,
there were 1,237 lion trophy exports
reported in UNEP–WCMC, 336 if South
Africa is excluded.
Here it should be noted that there are
limitations to interpreting the above
reported information. The 2004 guide to
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misinterpreted if one is not familiar
with it (CITES 2004b, p. 5). The number
of ‘‘trophies’’ reported does not
necessarily equate to the number of
lions hunted. Additionally, the number
of trophies reported for a given year in
the trade report does not equate directly
to the number of animals hunted in that
given year (CITES export permits may
be valid for 6 months, and a trophy
could in theory be exported the year
after it was hunted). The second
limitation to interpreting this
information is, although many permits
may indicate that an animal is of wild
origin (source code ‘‘W’’), these permits
may be incorrectly coded. This is true
for South Africa, where during the
period of 2000 to 2009, animals that
were captive-born and released into
private reserve systems were assigned
an incorrect source code of ‘‘wild.’’
South Africa has since requested their
provincial authorities to use the correct
source code for ‘‘captive bred’’ in order
to correctly reflect the source of sporthunted lion trophies; however, some
provinces are still not complying (RSA
2013, pp. 8–9). However, based on
South African trade data, the bulk of the
exports of lions and their parts and
products (including trophies) from
South Africa were from captive-born
lions (RSA 2013, p. 7).
Tanzania, with the highest lion
populations (Hamunyela et al. 2013, pp.
29, 283; Riggio et al. 2013, p. 32; Ikanda
2008, p. 4; Baldus 2004, pp. 5, 6), was
the largest exporter of wild-origin lion
trophies, but their exports have
decreased significantly since 2006. In
2008, approximately 138 lions had been
estimated to be killed in Tanzania as
trophies. In 2010, Tanzania’s numbers
declined to 128 exports, 55 in 2011, and
42 in 2012 (CITES lion gross exports,
https://trade.cites.org/, accessed April
25, 2014). In 2012, Tanzania established
an annual quota to limit trophy hunting
to no more than 50 animals (Jackson
2013, p. 7). Again, it should be noted
that there may be discrepancies between
the annual quota and the actual number
of trophies exported in a given year (see
https://www.cites.org/common/
resources/TradeDatabaseGuide.pdf for
additional information). Regardless, the
numbers of lion trophies exported by
Tanzania according to the UNEP–
WCMC database suggest a decreasing
trend.
In other areas within the range of the
African lion, the number of lions hunted
or authorized to be hunted annually has
remained fairly consistent. In Burkina
Faso, approximately 12 lions per year
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have been hunted over the past two
decades (IUCN 2009, pp. 36–37; Bauer
and Nowell 2004, p. 36), although their
current annual quota is 6 animals. In
Botswana, a quota of 30 lions per year
was authorized for nearly two decades;
however, Botswana has recently
implemented a hunting moratorium
(Jackson 2013, p. 8). (CITES lion gross
exports, https://trade.cites.org, accessed
April 23, 2014; CITES UNEP–WCMC
database, accessed January 8, 2014, and
August 16, 2013).
Potential Impacts of Trophy Hunting
Infanticide and Age-Based Hunting
Strategies
Tourist safari hunting of males has
been suggested by the petitioners to
increase infanticide rates (when males
kill young lion cubs sired by other
males) (Petition 2011, p. 24; Whitman et
al. 2004, p. 175), due in part to trophy
hunters taking males under a certain
age. Removing a younger male lion is
purported to allow another male to take
over the pride, and kill the former
patriarch’s cubs. This supposition is
inconclusive and not well supported
(CITES 2014a, p. 14; Dagg 2000, pp.
831–835). Infanticide is a common
practice among many species, including
lions (Hausfater et al. 1984, pp. 31, 145,
173, 487). When an adult male lion in
a pride is killed, surviving males who
form the pride’s coalition become
vulnerable to takeover by other male
coalitions, and this often results in
injury or death of the defeated males
(Davidson et al. 2011, p. 115). In some
cases, replacement males who take over
the pride will kill all cubs less than 9
months of age in the pride (Whitman et
al. 2004, p. 175). One range country
specifically addressed this issue; the
Republic of Namibia indicates that lion
populations reproduce at similar rates
in both harvested and non-harvested
populations, but it is unclear whether
cub survival is consistent in harvested
vs. non-harvested lion populations.
While utilizing individual-based
simulation models, Whitman et al.
(2004, pp. 175–177) found that if offtake
is restricted to males older than 6 years
of age, then trophy hunting will likely
have minimal impact on the pride’s
social structure and young (Packer et al.
2006, p. 6). This 6-year age restriction
approach for lion trophies is in the
process of being self-implemented,
along with other best practices, by
professional hunting guides, and is
being adopted by certain range states
(White 2013, p. 14; Davidson et al. 2011,
p. 114; Whitman et al. 2004, p. 176). It
involves conducting an age assessment
of male lions using identification
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techniques, such as mane development,
facial markings, nose pigmentation and
tooth-aging, to establish the relative age
of male lions. Tooth wear on incisors,
yellowing and chipping of teeth,
coupled with scars, head size, mane
length and color, and thinning hair on
the face, as well as other factors can be
an indicator of advanced age in lions
(Whitman and Packer 2006, entire).
Although these characteristics may be
subjective, as regional differences may
occur between lion populations, there
are clear attempts by the trophy hunting
community to establish and implement
best practices. Promoting the removal of
males 6 years of age or older,
theoretically allows younger males the
opportunity to remain resident long
enough to rear a cohort of cubs
(allowing their genes to enter the gene
pool; increasing the overall genetic
diversity). By removing males in a
manner that promotes healthy
population growth, the lion population
could yield more males in the long term
(Davidson et al. 2011, p. 114; Whitman
et al. 2004, p. 176). The governments of
Tanzania, western Zimbabwe,
Mozambique in the Niassa National
Reserve, Zambia, and most recently
Benin have instituted or are in the
process of instituting reforms such as 6year age restrictions on lion trophies to
increase the likelihood that trophy
hunting of lion is sustainable in those
countries (Van der Merwe 2013, p. 2;
Jackson 2013, p. 3; White 2013, p. 14;
Dallas Safari Club 2013, pp. 1–2; Hunter
et al. 2013, p. 2).
In addition to quota-setting,
moratoriums, and the 6 year age limit,
it has been reported that more protective
standards and guidelines are
implemented, such as the best practices
listed below (Jackson 2013, pp. 3, 8–10,
Dallas Safari Club 2013, pp. 1–2).
• Minimum trophy quality, sizes, and
standards;
• Wildlife hunting regulations
enacted and enforced;
• Professional hunting associations
formed;
• Professional hunting training
courses;
• Professional hunter standards
established;
• Quota-setting procedures;
• Compliance with CITES
demonstrated;
• Monitoring; and
• Information and data collection and
analysis.
While the supposition of increased
infanticide due to the remove of
established males from a pride is
inconclusive and not well supported, it
is clear that improved management
practices are beneficial to maintaining
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viable lion populations. Developing and
implementing best management
practices, while not categorically
establishing a direct correlation with
increased population numbers and
health, do appear to have practical
impacts on lion populations. Based on
the best available scientific and
commercial information, infanticide, as
a result of the removal of lions through
hunting, is not a threat to African lions.
Further, it is not likely to become a
threat in the foreseeable future since the
science is not well supported as to
whether infanticide resulting from
offtake due to trophy hunting is a
significant threat to the subspecies
(Whitman et al. 2004, pp. 175–176;
CITES 2014a, p. 14).
Corruption
Corruption is common in some areas
within the range of the African lion,
particularly in areas with extreme
poverty (Michler 2013, pp. 1–3; Kimati
2012, p. 1; Garnett et al. 2011, p. 1;
IUCN 2009, p. 89; Leader-Williams et al.
2009, p. 296–298; Kideghesho 2008, pp.
16–17; https://www.transparency.org).
Several of the range countries of African
lion have experienced political
instability for many years, which
appears to be a contributing factor in
intensifying levels of corruption.
Political instability results in war and
famine, which essentially halt
conservation efforts and the
enforcement of existing wildlife
protection laws (Barnett & Patterson
2005, p. 82). Corruption manifests itself
in several ways, including
embezzlement of funds and acceptance
of bribes to overlook illegal activities or
for political influence (Garnett et al.
2011, p. 1). Given the financial aspects
of sport hunting, it is reasonable to
assume that corruption and the inability
to control it could have a negative
impact on decisions made in lion
management by overriding biological
rationales with financial concerns.
Corruption has complex roots and
will not end immediately, but from all
appearances, it is being addressed in
many of the African lion range countries
where it has occurred in the past.
Countries throughout the range of the
African lion are putting tools in place to
combat corruption and create awareness
(https://www.transparency.org/cpi2012/
results, accessed June 20, 2013). In
recent years, in several African lion
range countries, leadership has taken
steps to address corruption, or activities
that facilitate corruption, associated
with wildlife management. For example,
in 2013, the Tourism Minister of Zambia
banned hunting in 19 game management
areas for 1 year due to corruption and
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malpractice among the hunting
companies and various government
departments. Some game management
areas and privately owned game ranches
were not included in the ban, but lion
hunting appears to be currently
prohibited throughout the country
(Michler 2013, pp. 1–3). According to
some authors (Martin 2012, pp. 4, 104;
Kimati 2012, p. 1; Kideghesho 2008, pp.
16–17), corruption in the wildlife sector
has often been one of the most
discussed topics in Tanzania’s National
Assembly, which presumably would
indicate the awareness of and
willingness to address the corrupting
factors in the wildlife sector.
Provided that countries continue to
address corruption within the wildlife
sector, we conclude, based on the best
scientific and commercial information
available, that corruption, in and of
itself, does not currently pose a threat to
the species. However, if efforts to
address corruption do not continue, it
could become a threat to African lions
in the future.
Revenue From Trophy Hunting
The high value of lions makes them
one of the most expensive large game
species to hunt. The revenue derived
from lion hunting is substantial. Lions
are reported to generate the highest
daily rate of any mammal hunted (USD
$2,650 per day), the longest number of
days that must be booked, and the
highest trophy fee ($24,500) (Jackson
2013, p. 6; Lindsey et al. 2012a, p. 5).
According to Groom (2013, p. 4), a 21day lion hunt in Zimbabwe may be sold
for approximately $2,500 per day, with
an additional trophy fee of $10,000.
Depending on the country in which a
hunter visits, there may be several
different fees required, including game
fees, observer fees, conservation fees,
permit fees, trophy handling fees, and
government payments in terms of taxes,
as well as safari operator fees (Barnett &
Patterson 2005, p. 71). In the late 1990’s,
Tanzania reported annual revenue of
$29.9 million from all trophy hunting,
South Africa reported $28.4 million,
Zimbabwe reported $23.9 million from
all trophy hunting, Botswana reported
$12.6 million, and Namibia reported
$11.5 million; the revenue generated
solely from lion hunting was not broken
out (Barnett & Patterson 2005, p. iv). In
the past, government and private land
owners were the primary beneficiaries
of the revenue gained; however, a
portion of the revenue derived from
hunting, in some countries, is now
being distributed to local communities
as well, which benefits the livelihoods
of local people as well as contributes to
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national economies of African range
states (Barnett & Patterson 2005, p. vi).
Trophy Hunting as a Wildlife
Management Tool
The concept of using trophy hunting
to support lion conservation is complex
and counterintuitive to many. Many
range countries rely heavily on tourism
(predominantly ecotourism and safari
hunting) to provide funding for wildlife
management (IUCN 2006a, p. 24). The
countries that rely most on lion hunting
are proportionally the highest in
Mozambique, Tanzania, and Zambia
(Lindsey et al. 2012a, pp. 7–8). The
revenue generated from these industries
provides jobs for locals, such as game
guards, cooks, drivers, and security
personnel, and often brings in revenue
for local microbusinesses that sell art,
jewelry, and other native crafts.
Revenue generated from scientifically
based management program is used to
build and maintain fences, provide
security personnel with weapons and
vehicles, provide resources for antipoaching activities, and provides
resources for habitat acquisition and
management (Chardonnet et al. 2010,
pp. 33–34; Newmark 2008, p. 321).
Revenue from trophy hunting increases
the ability of many African countries to
manage wildlife populations both
within and adjacent to reserves; many of
these hunting areas are geographically
linked to national parks and reserves,
providing wildlife corridors and buffer
zones (Chardonnet et al. 2010, p. 34;
Newmark 2008, p. 321).
Proponents and most species experts
support trophy hunting as a
conservation tool for the African lion
(Hunter 2011, entire; van der Merwe
2013, entire; Hunter et al. 2013, entire)
because it provides: (1) Incentives for
the conservation of large tracts of prime
habitat, and (2) funding for park and
reserve management, anti-poaching, and
security activities. As habitat loss has
been identified as one of the primary
threats to lion populations, it is notable
that the total amount of land set aside
for hunting throughout Africa, although
not ameliorating the concerns about
habitat loss, exceeds the total area of the
national parks, accounting for
approximately half of the amount of
viable habitat currently available to
lions (Chardonnet et al. 2010, p. 34;
Packer et al. 2006, pp. 9–10). In
Tanzania, 25–33 percent of the total
area, encompassing 190 hunting units
and over 247,000 km2, has been set
aside for sport hunting purposes; this
has resulted in an area 5.1 times greater
than Tanzania’s fully protected and
gazetted parks (Jackson 2013, p. 6;
Barnett & Patterson 2005, p. 61).
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In Botswana, despite the current ban
on lion hunting, the country currently
has over 128,000 km2 of gazetted
wildlife management areas and
controlled hunting areas set aside for
hunting purposes, which equates to 22.1
percent of the country’s total area. This
is in addition to 111,000 km2 (or 19.1
percent) that has been set aside as
habitat in the form of National Parks,
Game Reserves, and Forest Reserves
(Barnett & Patterson 2005, p. 7).
Tanzania has land set aside for sport
hunting in the form of safari areas,
communal land, and privately owned
properties that make up 23.9 percent of
the total land base (Barnett & Patterson
2005, pp. 76–77). In 2000, five countries
in southern Africa (Botswana, Namibia,
South Africa, Tanzania, and Zimbabwe)
had set aside a combined 420,000 km2
of communal land, 188,000 km2 of
commercial land, and 420,089 km2 of
state land totaling over 1,028,000 km2
for sport hunting purposes (Barnett &
Patterson 2005, p. iii). As a species with
a considerable range (up to 1,000 km2)
(Packer et al. 2013 p. 636; Haas et al.
2005, p. 4), suitable habitat is important
to the survival of the species, and the
marked decline in suitable habitat is a
significant threat to the species (see
Habitat Loss). The land currently
designated for use in sport hunting has
helped to reduce, but not eliminate, the
impact of habitat loss for the African
lion.
Cost estimates for maintaining lion
populations range, from an annual
budget of $500 per km2 in smaller
fenced reserves to $2,000 per km2
annually for unfenced populations
(Packer et al. 2013, p. 640; Lindsey et al.
2012a, p. 9). This includes but is not
limited to costs associated with
permanent and temporary staff, fencing
installation and maintenance (fences
can cost $3,000 per km to install),
infrastructure maintenance, antipoaching activities such as surveillance
and snare/trap removal, wildlife
restocking fees (both for lions killed by
illegal poaching/snares as well as other
trophy species killed by lions on the
reserves), community outreach, and
compensation for loss of livestock in
surrounding communities (Packer et al.
2013, p. 640; Groom 2013, pp. 4–5;
Lindsey et al. 2012a, p. 9; Barnett &
Patterson 2005, p. 82). For example, in
´
the past, the Save Valley Conservancy in
Zimbabwe invested $546,000 annually
on anti-poaching activities and
employed 186 permanent scouts, while
operators in Coutada 16, Mozambique,
spent $60,000 annually on antipoaching (such as the removal of 5,000
gin traps) (Groom 2013, p. 5; Lindsey et
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al. 2012a, p. 9). According to Barnett
and Patterson (2005, p. 82), in
Zimbabwe:
Land invasions, resettlement and political
instability has had dire consequences for
wildlife occurring in the commercial sector.
Land invasions have affected all wildlife
management activities, and resulted in severe
habitat destruction, increased poaching and
infrastructure damage with thousands of
kilometers of fences being destroyed to make
wire snares . . . A typical questionnaire
response from an invaded 50,000 acre farm
in Masvingo Province . . . indicates
substantial poaching losses of up to
$1,819,040, with over 3,400 snares recovered
and 134 poachers arrested in just two
months.
Niassa National Reserve,
Mozambique, incurs annual costs of
approximately $1.9–2 million to
maintain a 42,000-km2 area (Lindsey et
al. 2012a, p. 9). As a single source of
revenue, the trophy hunting of lions
provides a substantial source of funds to
pay for the management of lion habitat.
According to Lindsey et al. (2012a, p. 5),
with the exception of rhinoceros and
exceptional elephant trophies, ‘‘lions
generate the highest revenue per hunt of
any species in Africa.’’ In Niassa
National Reserve, lion trophy hunting
has generated $380,000–400,000
annually (Lindsey et al. 2012a, p. 9). In
´
the Save Valley Conservancy, between
2005 and 2011, lion hunting in
Zimbabwe provided an estimated net
income (based on 26 lions) of
approximately $1,365,000 in per-night
charges and roughly $260,000 in trophy
fees (Groom 2013, p. 4).
Trophy hunting of lions, if part of a
scientifically based management
program, can provide direct benefits to
the species and its habitat, both at the
national and local level (See: Role of
Local Communities in Lion
Conservation). Trophy hunting and the
revenue generated from trophy hunting
are tools that range countries can use to
facilitate maintaining habitat to sustain
large ungulates and other lion prey,
protecting habitat for lions, supporting
the management of lion habitat, and
protecting both lions and their prey base
through anti-poaching efforts. While
hunting alone will not address all of the
issues that are contributing to the
declined status of the species, it can
provide benefits to the species.
Role of Local Communities in Lion
Conservation
Over the last few decades,
conservationists and range countries
have realized the integral role local
communities play in the conservation of
lions and their habitat; when
communities benefit from a species,
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they have incentive to protect it.
Therefore, utilizing the wildlife sector
as a land-use option and source of
income for rural populations has
increasingly been employed throughout
the range countries of the African lion.
Many of these countries are classified as
‘developing’ nations; specifically, seven
of the ten countries (we include
Cameroon here) where trophy hunting is
permitted have 27–64 percent of their
populations living in severe poverty
(United Nations Development
Programme’s (UNDP) Human
Development Report, https://
hdr.undp.org/en/data, accessed July 7,
2014; Barnett & Patterson 2005, p. iii).
These countries often have high
population growth, high
unemployment, limited industry, and a
Gross Domestic Product (GDP) per
capita lower than the poverty level
(Barnett & Patterson 2005, p. iii). These
combined challenges highlight the need
for innovative solutions.
Conservationists and range countries
recognize the value of the wildlife
sector; if managed sustainably, there is
high potential to contribute to rural
economic development while
simultaneously protecting the unique
ecological habitats and species
contained therein (Chardonnet et al.
2010, p. 33; Kiss [editor] 1990, pp. 1, 5–
15).
Studies have indicated that, in order
for species such as the African lion to
persist, the local communities must
benefit from or receive a percentage of
funds generated from tourism such as
wildlife viewing, photography, or
trophy hunting (White 2013, p. 21;
Martin 2012, p. 57; Kiss [editor] 1990,
pp. 1, 5–15). The economic value of a
species, such as lion, can encourage
range countries to develop management
and conservation programs that involve
local communities which would
ultimately discourage indiscriminate
killings by local communities (Groom
2013, pp. 3, 5; Hazzah et al. 2013, p. 1;
White 2013, p. 21; Martin 2012, p. 49).
If local communities see no beneficial
value of lions being present in their
communal areas, sustainable utilization
of lions as a land-use becomes less
competitive with other land-use
options, such as grazing and livestock
management, and local communities
become unwilling and unable to manage
their wildlife heritage (Barnett &
Patterson 2005, p. iii). When the value
of lions in areas outside of national
parks is diminished, those areas are
likely to be converted to forms of land
use less suitable for lions, such as
agriculture, livestock pastures, or areas
of resource extraction, making them
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even more vulnerable to expanding
human settlement (Van der Merwe
2013, p. 2).
Community conservancies that benefit
from trophy hunting have specifically
been formed as a way to protect wildlife
and habitat. As an example, in Namibia,
160,000 km2 (61,776 mi2) of community
conservancies were established in part
due to revenue from trophy hunting.
These conservancies benefit the local
communities, which in turn protect lion
´
habitat. For example, in 2012, the Save
Valley Conservancy (Zimbabwe)
‘‘provided over US$100,000 worth of
support to adjacent villages or farmers
in the resettled areas. Assistance
included drilling boreholes, maintaining
boreholes, dredging of dams, building
clinics and schools, assisting with
repairs, maintenance and materials for
schools, education initiatives, school
field trips, provision of computer
equipment in schools, and craft
programs’’ (Groom 2013, p. 5)
Connecting conservation to community
benefits can provide a value for wildlife,
including lions, where there was
previously resentment or indifference,
helping to instill a sense of importance
for lion conservation Additionally, an
estimated 125,000 kg of game meat is
provided annually to rural communities
by trophy hunters at an estimated value
of $250,000 per year, which is
considerable for rural locations where
severe poverty and malnutrition exists
(White 2013, p. 21), further providing a
value for wildlife, including lions.
Lastly, local communities benefit from
the trophy hunting industry by gaining
employment as cooks, drivers, game
guards, security, and anti-poaching
personnel, and they also obtain revenue
for items purchased by trophy hunters
such as jewelry, art, and native
handicrafts.
Trophy hunting as part of a
scientifically based management
program may provide direct economic
benefits to the local communities and
can create incentives for local
communities to conserve lions, reduce
the pressure on lion habitat, and end
retaliatory killing, primarily because
lions are viewed as having value.
Conversely, lack of incentives could
cause declines in lion populations
because lions are viewed as lacking
value and are perceived to kill livestock,
which do have value to communities
(see Human-lion Conflict).
Many range countries have realized
local communities must benefit from the
conservation of the species because
[why?] and have revised their land
management and ownership policies to
reflect this. Of the ten countries where
lion trophy hunting currently occurs
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(including Cameroon), seven have
developed National Poverty Reduction
Strategies in partnership with the
International Monetary Fund (for a
complete list, see https://www.imf.org/
external/np/prsp/prsp.aspx); each of
these has incorporated sustainable
natural resource development as a main
priority, and emphasized benefit
distribution and management to rural
communities (Benin 2000, unpaginated;
Burkina Faso 2000, unpaginated;
unpaginated; CAR 2000, p. 45;
Mozambique 2000, unpaginated;
Tanzania 2000, pp. 13, 21; Zambia 2000,
unpaginated). As a result, an increase in
participation by local communities in
managing natural resources that are
adjacent to reserves is occurring in
several areas.
Captive Lions
In analyzing threats to a species, the
Service focuses its analysis on threats
acting upon wild specimens within the
native range of the species, because the
goal of the Act is survival and recovery
of the species within its native
ecosystem. We do not separately
analyze ‘‘threats’’ to captive-held
specimens because the statutory five
factors under section 4 (16 U.S.C. 1533)
are not well-suited to consideration of
specimens in captivity and captive-held
specimens are not eligible for separate
consideration for listing. However, we
do consider the extent to which
specimens held in captivity create,
contribute to, reduce, or remove threats
to the species.
Captive-held African lions, including
those that are managed for trophy
hunting in South Africa and lions held
in captivity in zoos, are believed to
number between a few thousand and
5,000 worldwide (Republic of South
Africa 2013, p. 5; Barnett et al. 2006a,
p. 513). Captive lions in general are not
suitable for reintroduction due to their
uncertain origins (Barnett et al. 2006a,
p. 513; Hunter et al. 2012, p. 3),
potential maladaptive behaviors, and
higher failure risk compared to
translocated individuals (Hunter et al.
2012, pp. 2–3). There may be cases
where captive specimens provide a
benefit to the species under certain
circumstances. For example, the display
of Giant pandas in U.S. zoos has
generated considerable revenue that is
used for in-situ conservation of the
species in China. It may be possible that
captive lions could also serve a purpose
of generating revenue for in-situ
conservation.
Summary of Trophy Hunting
Although there is some indication
that trophy hunting could contribute to
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local declines in lion populations
through unsustainable quotas,
corruption, and possible disruption of
pride structure through infanticide and
take of males that are too young, we do
not find that any of these activities rises
to the level of a threat to the African
lion subspecies at this time. It appears
that most range countries that allow
trophy hunting of African lions restrict
offtake to approximately 2–4 percent of
their lion populations for trophy
hunting annually, excluding South
Africa, where offtake is from
predominantly captive-born animals,
and Zimbabwe, where offtake is 2–3
percent higher than in other countries
(Packer et al. (2006, pp. 2–3). Exports of
lion trophies have demonstrated a
decreasing trend when exports of likely
captive-born lions from South Africa are
excluded (CITES lion gross exports,
https://trade.cites.org, accessed April 23,
2014), and lions from South Africa are
likely captive-born (RSA 2013, p. 5).
Most of the range countries that allow
trophy hunting have quotas in place to
limit take. Tanzania, with a population
of approximately 16,000 lions, has a
quota of 50 animals per year. Many
other range countries have laws in effect
that address trophy hunting, and several
have moratoriums in place. The hunting
community is taking the lead in
developing best management practices
to address take of males that are under
6 years of age, and they are guiding the
development of scientifically based
tools for minimizing the impact of
trophy hunting on the social structure of
lion populations. This 6-year age
restriction on lion trophies is in the
process of being self-implemented by
professional hunting guides, and is
being adopted by certain range states,
such as Tanzania (White 2013, p. 14;
Whitman et al. 2004, p. 176).
Currently, most countries that allow
trophy hunting of lions appear to be
reviewing their trophy hunting practices
(Jackson 2013, pp. 2–3; White 2013, pp.
12–13). Range countries have
recognized the need to incorporate best
management practices, and have been
progressively updating the policies and
management systems in order to
implement them (Lindsey et al. 2013a,
pp. 4–10).
Finally, we found that, if trophy
hunting of lions is part of a scientifically
based management program, it could
provide considerable benefits to the
species, by reducing or removing
incentives by locals to kill lions in
retaliation for livestock losses, and by
reducing the conversion of lion habitat
to agriculture. Trophy hunting, if
managed well and with local
communities in mind, can bring in
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needed revenue, jobs, and a muchneeded protein source to local people,
demonstrating the value of lions to local
communities (Groom 2013, pp. 1–3;
Lindsey et al. 2006, pp. 283, 289). In
addition, the amount of habitat that has
been set aside by range countries
specifically for trophy hunting has
greatly increased the range and habitat
of lions and their prey base, which is
imperative given the current ongoing
rate of habitat destruction occurring in
Africa. The total amount of land set
aside for trophy hunting throughout
Africa exceeds the total area of the
national parks, providing half the
amount of viable lion habitat
(Chardonnet et al. 2010, p. 34; Packer et
al. 2006, pp. 9–10). However, expanding
protected areas without taking the
human population into consideration
could lead to more resentment and
retaliatory killing of lions (Nelson et al.
2009, p. 315).
Therefore, we conclude, based on the
best scientific and commercial
information available, that trophy
hunting is not a significant threat to the
species.
Traditional Use of Lion Parts and
Products
CITES (2014, p. 8) reports that many
African countries, including Somalia,
Nigeria, Burkina Faso, Kenya, and
Cameroon, maintain local markets in
lion products, which include teeth,
claws, fat, whiskers, bone, bile, testicles,
meat, and tails for use as talismans,
decorations, and in traditional African
medicine. In Ghana, lion parts and
products are used for ceremonial,
medicinal, and nutritional purposes
(Burton et al. 2010, p. 4). Skins and
claws of lions were observed for sale in
a market in Tamale, Ghana. Lions in and
around Mole National Park in Ghana
have been killed for traditional
consumptive purposes (Burton et al.
2010, p. 4). In some cases, lions (either
alive or dead) have been ‘‘laundered’’
through other countries so that their
country of origin is unknown. As an
example, lions have been found to be
shot in Zimbabwe and Mozambique and
declared as South African trophies (Lion
Aid 2011, p. 20). In other cases, there
have been reports of captive-born lions
being smuggled between Botswana and
South Africa and described as wild
(Mouton 2013, pp. 1–2). Lion products,
such as the trade in lion bone, seem to
be primarily byproducts of trophy
hunting; hunters are primarily
interested in the trophy and skin and,
therefore, the bones and other parts are
sold separately (CITES 2014a, p. 10).
However, since the reports of these
types of activities are primarily
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anecdotal in nature, based on the best
available scientific and commercial
information, we find that the sale of
these byproducts does not currently
pose a threat to the species. Further,
without a significant shift in the market,
it is not likely to become a threat in the
foreseeable future.
Conservation Measures in Place To
Protect Lions
There has been awareness for several
years that conservation strategies need
to be implemented for the African lion
due to the apparent decrease in its
population numbers (Hamunyela et al.
2013, p. 1; Henschel et al. 2010, p. 34;
Gebresenbet et al. 2009, p. 5; IUCN
2006a, b, entire). Prior to 2006,
institutional inconsistencies throughout
the African lion’s range resulted in poor
lion conservation policies and little to
no enforcement of existing laws (IUCN
2006b, p. 18). As mentioned, in 2005
and 2006, nongovernmental
organizations (NGOs) and several
governments at various levels organized
two regional lion conservation
workshops. Species specialists, wildlife
managers, and government officials
attended these regional workshops in
order to provide range country
governments with frameworks for
developing their own national action
plans for the conservation of lions. Over
50 lion specialists, representing all lion
range countries, participated in these
workshops (Henschel et al. 2010, p. 34).
During the workshops lion experts
collectively assessed what they believed
to be the then-current status of African
lions based on a variety of information,
and subsequently identified 86 African
LCUs. This information was then used
as a framework to identify lion areas,
strongholds, and potential strongholds
by Riggio et al. (2013, p. 32).
Many countries with very small lion
populations have developed or updated
their conservation plans for the African
lion. Some of these include Benin,
Cameroon, Uganda, and Malawi. Some
range countries participate in
transboundary conservation projects
and are collaborating on transboundary
lion conservation initiatives for shared
lion populations. Most range countries
have a national lion action plan or
strategies in place, particularly if there
are economic incentives for them to
have viable lion populations (Groom
2013; Nghidinwa et al. 2013, pp. 11–12;
Zambia Wildlife Authority 2012; Lion
Aid 2011, pp. 1–2; Mesochina et al.
2010; Government of Tanzania 2010;
Begg and Begg 2010). Range states have
also implemented a number of
conservation strategies designed to
conserve habitat, reduce human-lion
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conflict, and preserve the lion’s preybase.
Conservation Measures To Stem Habitat
Loss
Habitat loss represents one of the
main threats facing the African lion
(Bauer et al. 2008, unpaginated).
Attempts by range countries to address
this decline in habitat are manifested in
a number of ways, such as the creation
of protected areas and the establishment
of wildlife corridors to connect
fragmented habitats.
Two conservation tools utilized by
range countries for African lions include
the establishment of protected areas and
the enforcement of protections in these
areas (Mesochina et al. 2010a and b;
Treves et al. 2009, pp. 60, 64). Over the
past few decades, the effectiveness of
protected areas in protecting habitat has
been studied, particularly in Africa
(Pfeifer et al. 2012, p. 1; Craigie et al.
2010, pp. 2,221–2,222). A study
conducted by the Wildlife Conservation
Society in 2005 found that most lion
populations in protected areas of
southern and eastern Africa have been
essentially stable over the previous
three decades (Ray et al. 2005, p. 67).
However, several problems have
emerged. For example, certain landtenure systems do not recognize
community ownership of land and
wildlife and undermine the extent to
which benefits are converted into
incentives for conservation. Protectedarea ‘‘boundaries’’ are not always
visible. Additionally, law enforcement
in protected areas can be sporadic, and
parks are often understaffed (Pfeifer et
al. 2012, pp. 1, 7). Lastly, despite the
Wildlife Conservation Society’s
findings, more recent evidence suggests
that some protected areas are being
more commonly encroached upon as
human populations expand and search
for resources.
Despite encroachment, protected
areas are somewhat effective at
protecting wildlife and habitat as rates
of habitat loss tend to be lower in
protected areas than outside them
(Green et al. 2013, p. 70; Pfeifer et al.
2012, p. 2). African countries are
realizing the benefits of managing their
wildlife populations and parks for
tourism; however, conservation of vast
areas of land for megafauna such as the
African lion is not only complex, but
also expensive. As an example, the 28km (17-mi) elephant corridor,
completed in 2011 in Kenya, cost $1
million (The Nature Conservancy 2013,
unpaginated). Additionally, the overall
costs of anti-poaching and
compensation is expected to increase in
range states concurrently with growing
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human populations, declining
purchasing power of external funds, and
corruption (Garnett et al. 2011, pp. 1–2;
Wittemyer et al. 2008, pp. 123, 125).
Another mechanism for protecting
habitat is to reconnect fragmented
habitat across national boundaries.
Corridors are being restored, fences are
being removed, and protected areas are
being connected. Restoration of these
corridors allows wildlife to travel
between areas of suitable habitat (Jones
et al. 2012, pp. 469–470). In some areas,
fences have been constructed to protect
grazing resources for domestic livestock
as well as to provide barriers to disease
(Gadd 2012, pp. 153, 176). One aspect
of these fences is that they separate
lions from their prey. In southern
Africa, the trend now is to take down
fences to increase the size of connected
habitat and link it to reserves and
national parks (IUCN 2009, p. 101;
IUCN 2008, various). The Limpopo
Transfrontier Park is another example of
where this is being implemented
(Newmark 2008, p. 327). Boundary
fences along national borders that
separate many reserves are being
removed to form a 35,000-km2 park.
Limpopo National Park (formerly
known as Coutada 16) in Mozambique;
Kruger National Park in South Africa;
Gonarezhou National Park, Manjinji Pan
Sanctuary, and Malipati Safari Area in
Zimbabwe will all be connected, as will
be the area between Kruger and
Gonarezhou, and the Sengwe communal
land in Zimbabwe and the Makuleke
region in South Africa (Newmark 2008,
p. 327). However, in some locations,
areas that have previously been
designated as corridors have been
encroached upon by human settlements
and agriculture (Estes et al. 2012, pp.
258–261; Jones et al. 2012, p. 469).
Tanzania is an example of a country
attempting to reconnect habitat. As of
2002, the Tanzanian Government, with
donor and NGO support, was
reconnecting the nine largest blocks of
forest in the East Usambara Mountains
using wildlife corridors (Newmark 2002,
various). Additionally, the 2009
Wildlife Act of Tanzania allows the
Minister, in consultation with relevant
local authorities, to designate wildlife
corridors, dispersal areas, buffer zones,
and migratory routes. The 2010–2015
National Elephant Management Plan of
Tanzania indicates that corridors are the
primary objective of the plan, and
although primarily designed for
elephants, these corridors allow for
continuity of populations of other large
mammal species such as lions (Jones et
al. 2012, p. 470).
In 2011, Kenya (which neighbors
Tanzania to the North), completed a 28-
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km corridor through an area that had
been heavily impacted by humanwildlife conflict. The purpose of the
corridor was primarily to reduce
human-elephant conflict and appears to
have been successful (Mount Kenya
Trust 2011, p. 1). The corridor also
allows other wildlife such as lions to
disperse through habitat that otherwise
would have been unfavorable for
wildlife to travel through (Mount Kenya
Trust 2011, p. 1). It was an expensive
project, but recent reports indicate that
the effort has served its purpose:
Elephants are using the corridor on a
regular basis (particularly an underpass
under a highway), and humans are
reporting less human-wildlife conflict
(Mount Kenya Trust 2011, p. 1).
However, connectivity alone does not
ensure the dispersal of animals (Roever
et al. 2013, pp. 19–21). The Tanzania
Wildlife Research Institute (TAWIRI) is
a parastatal organization under
Tanzania’s Ministry of Natural
Resources and Tourism, and is
responsible for conducting and
coordinating wildlife research activities
in Tanzania (https://tawiri.or.tz/). In this
role, TAWIRI has been actively involved
in promoting the development of and
monitoring the use of wildlife corridors
in Tanzania (https://
www.tzwildlifecorridors.org). Surveys
conducted in 2009 and 2010 suggest
that the Nyanganje Corridor in Tanzania
is no longer being used by elephants
and other wildlife. This corridor is at a
narrow passage in the Kilombero Valley
and is the shortest distance for animals
to cross between the Udzungwa and
Selous ecosystems. Despite efforts in
place, much of the corridor is being
encroached upon by conversion of land
to rice farming and cattle grazing (Jones
et al. 2012, p. 469). Because these
activities often deter wildlife from
passing through, the corridor is
ineffective (Jones et al. 2012, p. 469).
TAWIRI reminds wildlife managers that
they need to continue to implement
steps to ensure that corridors are
functioning properly.
Conservation Measures in Place To
Stem the Loss of Prey Base
Lions, like most large carnivores, prey
upon a variety of species including
buffalo, plains zebra, wildebeest, giraffe,
gemsbok, kob, and warthog (Kenya
Wildlife Service 2013, p. 13; Niassa
National Reserve Technical Report
2011, p. 4; Nowell and Jackson 1996, p.
18). Depletion of these prey species due
to competition with humans represents
a threat to the lion (Chardonnet et al.
2005, pp. 8–9). As noted, the increase in
the human population in Africa is a
major contributor to the increase in the
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demand for bushmeat, which in turn
increases human encroachment into
wildlife lands (Lindsey et al. 2012b, p.
36). In addition to the increase in the
human population, lack of an
alternative livelihood, lack of alternate
food sources, and lack of clear rights
over land or wildlife are contributing
factors toward the increase in demand
for bushmeat (Lindsey et al. 2012b, pp.
36–41). The advent of automatic
weapons in the bushmeat trade impacts
the lion’s prey base, which is being
hunted at unsustainable levels.
Reconnecting fragmented habitat has
the additive effects of not only
conserving the biodiversity of the
African lion’s habitat, but also that of its
prey base (Lindsey et al. 2012b, p. 43).
These types of restoration practices
enhance the health of species by
allowing genetic interchange to occur
and, thus, conserve the genetic diversity
of all wildlife. Wildlife management
entities are linking many of the major
protected areas by removing boundary
fences along national borders that
separate many reserves in addition to
creating or improving corridors to link
good-quality habitat for wildlife (Gadd
2012, p. 179; Newmark 2008, pp. 323–
324). To address the increasing
consumption of bushmeat, host
countries have employed a variety of
different strategies, including the
development of alternative industries
for communities. Helping local
communities develop alternate
industries represents one of the ways
range countries can reduce their
dependence on bushmeat. Throughout
Africa, several ideas have been
attempted with varying levels of
success. For example, the Anne Kent
Taylor Fund (AKTF) helps local Maasai
women to buy beads and other supplies
to produce traditional items for the local
tourist industry (AKTF 2012, p. 7;
Lindsey et al. 2012b, p. 45; van Villet
2011, p. 17). In addition, AKTF helps
organize local men into anti-poaching
and de-snaring teams (AKTF 2012, p. 5;
van Villet 2011, p. 17). By creating
programs targeting both men and
women, AKTF creates an environment
that provides communities with
financial stability as well as direct
community interest in protecting local
wildlife. With 13 years assisting local
communities, the AKTF represents one
of the more successful attempts to
encourage locals to shift away from
relying on bushmeat.
Studies compiled by Huzzah 2013
(pp. 1, 8) have shown that local
communities who lived near protected
areas with more lenient policies have a
more positive attitude and relationship
with both the manager and the protected
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area as a whole. This open approach to
protected area management reflects a
trend in recent years to bring in local
communities to assist in the
management of protected areas (Lindsey
et al. 2012b, p. 53). Wildlife
management programs run by local
communities are defined by two goals:
Conserving wildlife and providing
economic aids to the community
(Bandyopadhyay et al. 2010, p. 5). With
regards to discouraging the
consumption of bushmeat, this new
approach is seen in the creation of
community-based wildlife management
programs (van Villet 2011, p. 26). The
purpose of these programs is to give the
local community a direct stake in the
management of wildlife areas. One use
for these areas is to turn them into game
ranches. These areas are used both for
legal bushmeat production as well as
trophy hunting and ecotourism.
One such program is the Chivaraidze
Game Ranch in Zimbabwe (van Villet
2011, pp. 28–29). The Chivaraidze Game
Ranch started in 1996 with the stated
goal of reducing poaching through
providing bushmeat at a reduced price.
However, internal infighting in the
organization over the devolution of
power to local communities, between
those in favor of devolution and a
powerful local interest group, limited
the effectiveness of the organization. In
the span of 8 years (between 2001 and
2009), the Chivaraidze Game Ranch has
had six different boards of directors
(Mombeshora and Le Bel 2010, p. 5).
Furthermore, a power shake-up in local
communities along party lines and
kinship affiliation limited the abilities
for communities to cooperate with each
other (van Villet 2011, pp. 28–29;
Mombeshora and Le Bel 2010, p. 7). The
result was that the cost of maintaining
the program exceeded the benefits to the
local community. The decline in
economic benefits to the local
community coincided with a resurgence
in poaching within areas of the park
(Mombeshora and Le Bel 2010, p. 3).
The result of the Chivaraidze Game
Ranch project reflects the difficulty in
shifting wildlife management from a
centralized national government
approach towards a more decentralized,
community-based approach.
Unlike the difficulties encountered in
Zimbabwe, Namibia has had greater
success in setting up community-run
conservancies. After gaining
independence in 1990, Namibia began
to turn over ownership of wildlife areas
to local communities (van Vliet 2011, p.
29; Bandyopadhyay et al. 2010, p. 6). By
2011, Namibia had 64 communities that
covered 17 percent of the country total
area (van Vliet 2011, p. 29; Connif 2011,
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npn; NASCO 2010, p. 4). The majority
of the incomes from these conservancies
come from ecotourism, followed by
trophy hunting (NASCO 2010, p. 22).
These incomes are then used to support
infrastructure improvement in the
community. In addition, legal bushmeat
acquired within conservancy lands is
distributed to local families (NASCO
2010, p. 25). The success of the program
in Namibia has been attributed to
Namibia’s unique characteristics,
including low population density and
favorable seasonal rain, which helps
prey species recover (van Vliet 2011, p.
30). Despite the successes in Namibia,
the country’s unique characteristics
mean that adapting Namibia’s success to
other, more densely populated countries
will be difficult.
Conservation Measures To Stem
Human-Lion Conflict
As the human population expands,
the potential for conflict with wildlife
increases. In Africa, conflict between
villagers and lions, who prey upon
livestock, represent a threat to the
species (Chardonnet et al. 2010, p. 12;
Moghari 2009, p. 14; IUCN 2006a, p.
23). In addition, habitat loss due to
conversion of land increases the chance
of villagers coming into direct contact
with lions (Chardonnet et al. 2010, p.
24). In an attempt to address these
problems, range countries have
employed a variety of different
strategies to help the lion. Such
strategies involve education, an effective
conservation plan, and interacting with
the local community.
Historically, range countries seek to
mitigate human-lion conflict through
controlling rather than conserving the
predator population. In countries such
as Malawi, for example, the Department
of Game, Fish and Tsetse Control would
shoot large carnivores that prey upon
livestock. The result of this policy was
that, between 1948 and 1961, over 560
predators (which include lions and
leopards) were killed in the country
(Mesochina et al. 2010b, p. 35). While
this department was disbanded in 1963
and jurisdiction shifted to the new
Department of Forestry, crop and
livestock protection still remains an
important part of its function. Despite
the department focusing on protecting
crops and livestock, the number of lions
killed in the country has declined.
Between 1977 and 1982, eight lions
were killed, whereas six lions were
killed between 1998 and 2007
(Mesochina et al. 2010b, p. 35). While
fewer lions are being killed than in the
previous decades, problems remain,
including lack of resources, lack of
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manpower, and corruption within the
range countries.
Current governmental management of
lions in countries such as Malawi,
Tanzania, and Zambia are managed by
the Problem Animal Control units
(Mesochina et al. 2010a, p. 41;
Mesochina et al. 2010b, p. 36). When
lion attack incidents occur, Problem
Animal Control dispatches officials to
investigate the problems. If the problem
lion is located, it is either removed or
eliminated. When properly funded, this
program has helped in reducing not
only conflicts between lions and
humans but also has driven down the
numbers of lions killed. Between 2005
and 2009, there were 116 reported cases
of lions killed, with the number of lions
killed being less than 50 per year in
Tanzania (Mesochina et al. 2010a, p.
41). However, limitations of resources
(including both manpower and funds)
have hampered the effectiveness of
these officials in responding to these
incidents. In addition, many Problem
Animal Control interventions resulted
in the death of the lion (Mesochina et
al. 2010a, p. 41; Chardonnet et al. 2009,
p. 36). Even in cases of translocation,
the lions that were being transported
often end up injured or continue to pose
problems to the community (Bauer et al.
2007, p. 91).
NGOs are also assisting in protecting
lions. Intervention by NGOs often takes
the form of interacting with the local
community (Winterbach et al. 2010, p.
98). Lion Guardians, which operate in
Kenya, recruits and educates local
young men. These men then monitor
and track lion movement and warn
herders of lion presence in the area,
thereby mitigating or preventing
possible lion-human conflict (Hazzah et
al. 2014, p. 853; Lion Guardians 2013,
p. 7; Lion Guardians 2012, p. 3). In
addition, Lion Guardians work with
tribal elders to dissuade young men
from killing lions for ceremonial
purposes. Historically, the killing of
lions through ritualized lion hunts
called ilmurran is rewarded with gifting
of cows and other rewards (Lion
Guardians 2012, p. 5; Goldman et al.
2010, p. 334). After introducing village
elders to the Lion Guardians program
first hand, many return home to their
village and give their blessings to the
project. This education led to significant
results; on August 11, 2013, two Lion
Guardians stopped a group of hunters
who were planning to hunt a lion in
retaliation for the lion preying on their
livestock. The local village elders fined
the potential hunters two cattle each for
going on a lion hunt, marking a gradual
but significant shift in the cultural
attitudes regarding the lion (Hazzah et
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al. 2014, p. 858; Lion Guardians 2013,
p. 20). Since its establishment in 2007,
only five lions had been killed in
territories where Lion Guardians
operates, in contrast to more than 100
lions killed in adjacent areas (Lion
Guardians 2013, p. 5). Furthermore,
reduced lion mortality was sustained
across multiple years, resulting in the
reserve having one of the highest lion
densities in Africa (Hazzah et al. 2014,
p. 857; Schuette et l. 2013, p. 149).
Despite the success of this program,
retaliatory as well as ceremonial killings
of lions outside the program areas
remain a threat to the species.
We found that many of the lion range
states are trying to address lion
conservation through the establishment
of protected areas, wildlife management
areas, wildlife corridors, and
reconnecting habitat. In some areas,
creating incentives for lion conservation
is occurring through community
conservation programs in range
countries. In other cases, participatory
strategies have been implemented to
enhance local tolerance for large
carnivores in Africa. An increasing
number of programs encourage local
communities to solve problems that
arise from human-lion conflict without
killing lions. However, the effectiveness
of these measures still ranges from
successful to unsuccessful, due in part
to lack of resources, political will, and
infighting. It is imperative that range
countries continue to recognize and
support the role that local communities
play in lion conservation. Greater
support by countries to address the
needs of local communities, and thereby
address the needs of lions, may be the
single-most important role these
countries can play in changing the
trajectory of lion declines.
Regulatory Mechanisms
Regulatory mechanisms in place to
provide protections to African lions
vary substantially throughout Africa. As
mentioned in the Conservation Status of
African Lions CITES section, lions are
listed in Appendix II under CITES, and
with the exception of South Sudan, all
of the lion range states are parties to
CITES. According to the draft CITES
Periodic Review of the Status of African
Lions (CITES 2014a, pp. 14–15) outside
of CITES, lions have no legal protections
in four countries: Burundi, Guinea
Bissau, Lesotho, and Swaziland.
However, CITES 2014a (p. 15), states
that most of the southern and eastern
lion range states have regulatory
mechanisms in place to protect lions.
We found that most of the range states
have national environmental legislation
to establish national parks and
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conservation areas, and to conserve and
regulate the take, hunting, and trade of
wildlife, including parts and products,
but could find no legislation specific to
lions, nor to the main threats affecting
lions: habitat loss, human-lion conflict,
and loss of prey base (See: Appendix A,
Ecolex information was accessed July 7–
10, 2014, at https://www.ecolex.org.8).
Our status review did not reveal
regulatory mechanisms in place that
specifically address the main threats
affecting lions. We are requesting
comments or information from lion
range states, other concerned
governmental agencies, the scientific
community, or any other interested
parties concerning regulatory
mechanisms that address the three main
threats to lions: habitat loss, human-lion
conflict, and loss of prey base.
Finding
Section 4 of the Act (16 U.S.C. 1533)
and its implementing regulations at 50
CFR 424 set forth the procedures for
adding a species to, and/or removing a
species from, the Federal Lists of
Endangered and Threatened Wildlife
and Plants. As noted in the Information
Requested section, a species may be
determined to be an endangered or
threatened species due to one or more
of the five factors set forth in section
4(a)(1) of the Act:
(A) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(C) Disease or predation;
(D) The inadequacy of existing
regulatory mechanisms;
(E) Other natural or manmade factors
affecting its continued existence.
In assessing whether the African lion
meets the definition of an endangered or
threatened species, we considered the
five factors in section 4(a)(1) of the Act.
A species is ‘‘endangered’’ for purposes
of the Act if it is in danger of extinction
throughout all or a significant portion of
its range and is ‘‘threatened’’ if it is
likely to become endangered within the
foreseeable future throughout all or a
significant portion of its range. The
‘‘foreseeable future’’ is the period of
time over which events or effects
reasonably can or should be anticipated,
or trends extrapolated.
8 ECOLEX is a comprehensive database on
environmental law, maintained by the International
Union for Conservation of Nature (IUCN), the
United Nations Environment Programme (UNEP),
and the Food and Agriculture Organization of the
United Nations (FAO). Our search terms used with
respect to wildlife laws were ‘‘African lion’’ and
‘‘country’’, e.g., ‘‘Angola’’, ‘‘Benin’’, etc. See
Appendix A.
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When considering what factors might
constitute threats to a species, we must
look beyond the mere exposure of the
species to a factor to evaluate whether
the species may respond to the factor in
a way that causes actual impacts to the
species. If there is exposure to a factor
and the species responds negatively, the
factor may be a threat and we attempt
to determine how significant a threat it
is. The threat is significant if it drives,
or contributes to, the risk of extinction
of the species such that the species may
warrant listing as endangered or
threatened as those terms are defined in
the Act. We conducted a review of the
best scientific and commercial data
available regarding the status of the
African lion and assessed whether the
African lion is endangered or threatened
throughout all of its range.
There is consensus within the
research community as well as lion
range states that the African lion is
impacted by a number of factors actively
contributing to its population decline
throughout Africa: habitat loss
(fragmentation and degradation) (Factor
A); decreased access to food prey
sources (aka loss of prey base) (Factor
B); retaliatory killing, snaring, and
poaching (both intentional and
unintentional), and deleterious effects
in its viability due to small populations
in some areas within its range (Factor E)
(Nyanganji et al. 2012, p. 12; Seguya et
al. 2010, p. 26).
We find three main threats, habitat
loss, loss of prey base, and human-lion
conflict, are impacting lions, alone and
in combination, such that the
subspecies is likely to become
endangered within the foreseeable
future throughout all of its range. In the
past several decades, the human
population has been expanding with
concomitant large decreases in lion
habitat and likely lion numbers,
resulting in an extremely large
reduction in the species’ range. As
human populations continue to rise in
sub-Saharan Africa, the amount of land
required to meet the expanding human
population’s needs is constantly
increasing. Lions are increasingly
limited to protected areas, and human
population growth rates around
protected areas in Africa tend to be
higher than the average rural growth
rate (Wittemyer et al. 2008, entire).
Considering the majority of the human
population in sub-Saharan Africa is
rural, and land supports the livelihood
of most of the population, loss and
degradation of lion habitat, loss of prey
base, and increased human-lion conflict
can reasonably be expected to
accompany the rapid growth in sub-
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Saharan Africa’s human population into
the foreseeable future.
Africa has the fastest population
growth rate in the world (UNEP 2012a,
p. 2). The majority of the population is
rural, and about 60–70 percent of the
population relies on agriculture and
livestock for their livelihood (UNEP
2006, pp. 82, 100, 106; IAASTD 2009, p.
2). As a result, a large portion of the
growing population will depend
directly on expansion of agriculture and
livestock grazing to survive in the
future. Between 2010 and 2050, the
population of sub-Saharan Africa is
projected to more than double to more
than 2 billion (from 831 million to 2.1
billion) (UN 2013, p. 9). During about
this same time period (2005 to 2050),
the area of cultivated land is projected
to increase by 51 million ha
(approximately 21 percent)
(Alexandratos and Bruinsma 2012, p.
107). However, this figure does not
include rangeland, and the majority of
agricultural land in Africa is devoted to
grazing (UNEP 2012b, p. 68), thus that
figure may be much larger. The number
of livestock (cattle, sheep, and goats) in
sub-Saharan Africa is projected to
increase about 73 percent, from 688
million to 1.2 billion, by 2050
(Alexandratos and Bruinsma 2012, p.
133). Therefore, in the case of African
lion, the best available scientific and
commercial data that we rely upon in
projecting future conditions for the
purpose of this listing determination
establish the foreseeable future to be
2050.
Human settlements and agricultural
and pastoral activities have expanded
into lion habitat and protected areas,
decreasing prey availability and
increasing exposure of livestock and
humans to lions. Human-lion conflict
and associated retaliatory killing of
lions will continue to play a major role
in the reduction of lion populations and
is the greatest current threat to
remaining lion populations. The lion’s
prey base has decreased in many parts
of its range in large part due to the
bushmeat trade
Bushmeat is the primary source of
protein for humans in much of the lion’s
range (Chardonnet et al. 2010, p. 27;
Mesochina et al. 2010a, p. 38; Abwe and
Morgan 2008, p. 26; Bennett et al. 2007,
p. 885; Fa et al. 2006, p. 507),
comprising between 6 percent (southern
Africa) and 55 percent (Central African
Republic) of a human’s diet (Chardonnet
et al. 2005, p. 9; IUCN 2006b, p. 19).
This reliance by humans on protein
obtained from bushmeat results in direct
competition for prey species between
humans and lions, and commercial
poaching of wildlife through the use of
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automatic weapons is a significant
threat to lion prey (Chardonnet et al.
2010, p. 27). Because many wildlife
species are being hunted at
unsustainable levels to meet this
demand within the range of the lion, its
prey base is becoming depleted in many
areas and has led to lion attacks on
livestock and humans (Hoppe-Dominik
et al. 2011, p. 452; Chardonnet et al.
2010, pp. 6, 13–14; Frank et al. 2006, p.
12). Given the rapid increase in humans
and livestock by 2050, we can
reasonably expect the conditions
described above to worsen. Also, as
livestock numbers increase and as
expansion of agricultural and pastoral
practices continue to deplete and
degrade the habitat that lion’s prey rely
on, the lion’s prey base is expected to
further decline. As the lion’s prey base
is hunted at unsustainable levels to
meet a growing demand for food,
livestock depredation and retributive
killing of lions through spearing,
shooting, trapping, and poisoning will
continue to occur, and will likely
increase (Dickman 2013, p. 379; HoppeDominik et al. 2011, p. 452; Chardonnet
et al. 2010, p. 19; Gebresenbet et al.
2009, p. 9; Hazzah and Dolrenry 2007,
p. 3).
Lion range countries are aware of the
threats affecting lions, and many are
working to address them. NGOs and
several governments at various levels
have organized regional lion
conservation workshops, which have
helped them to identify Lion
Conservation Units. Most range
countries have a national lion action
plan or strategy in place (Groom 2013;
Nghidinwa et al. 2013, pp. 11–12;
Zambia Wildlife Authority 2012; Lion
Aid 2011, pp. 1–2; Mesochina et al.
2010; Government of Tanzania 2010;
Begg and Begg 2010). Some range
countries participate in transboundary
conservation projects to create wildlife
corridors and reconnect habitat, and are
collaborating on transboundary lion
conservation initiatives for shared lion
populations. Reconnecting fragmented
habitat has the additive effects of not
only strengthening the biodiversity of
the African lion but also that of its prey
species (Lindsey et al. 2012b, p. 43).
Wildlife management entities are
linking many of the major protected
areas by removing boundary fences
along national borders that separate
many reserves, in addition to creating or
improving corridors to link good-quality
habitat for wildlife (Gadd 2012, p. 179;
Newmark 2008, pp. 323–324).
Range states have also implemented a
number of conservation strategies
designed to conserve habitat, reduce
human-lion conflict, and preserve lion
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prey-base. In order to address the
increasing consumption of bushmeat,
host countries have employed a variety
of different strategies, including the
development of alternative industries
for communities, which can reduce
their dependence on bushmeat. For
example, the Anne Kent Taylor Fund
(AKTF) helps local Maasai women to
buy beads and other supplies to produce
traditional items for the local tourist
industry (AKTF 2012, p. 7; Lindsey et
al. 2012b, p. 45; van Villet 2011, p. 17)
and has organized local men to
participate in anti-poaching and desnaring teams (AKTF 2012, p. 5; van
Villet 2011, p. 17). By targeting both
men and women in the community,
such programs provide communities
with financial stability as well as direct
community interest in protecting local
wildlife. African countries are realizing
the benefits of managing their wildlife
populations and parks for tourism;
however, conservation of vast areas of
land for megafauna such as the African
lion is expensive. The costs of antipoaching and compensation is expected
to increase in range states concurrently
with growing human populations,
declining purchasing power of external
funds, and corruption (Garnett et al.
2011, pp. 1–2; Wittemyer et al. 2008,
pp. 123, 125).
Studies have shown that local
communities who live near protected
areas (PAs) with community-based
conservation policies have more
positive attitudes and relationships with
both the park manager and the PA as a
whole (Huzzah 2013, pp. 1, 8). This
open approach to PA management
reflects a trend in recent years to bring
in local communities to assist in the
management of PAs (Lindsey et al.
2012b, p. 53). Wildlife management
programs run by local communities are
defined by two goals: conserving
wildlife and providing economic aids to
the community (Bandyopadhyay et al.
2010, p. 5). NGOs are also assisting in
protecting lions. Intervention by NGOs
often takes the form of interacting with
the local community (Winterbach et al.
2010, p. 98). For example, Lion
Guardians, which operates in Kenya,
has shown great success with its Lion
Guard program. Lion Guardians
educates local young men who monitor
and track lion movement and warn
herders of lion presence in the area,
thereby mitigating or preventing
possible lion-yhuman conflict (Hazzah
et al. 2014, p. 853; Lion Guardians 2013,
p. 7; Lion Guardians 2012, p. 3).
Outreach to tribal elders has
successfully helped elders to dissuade
young men from killing lions for
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ceremonial purposes. The result of such
programs has been a gradual change in
cultural attitudes towards lions (Hazzah
et al. 2014, p. 858; Lion Guardians 2013,
p. 20).
Finally, many range countries rely
heavily on tourism (predominantly
ecotourism and safari hunting) to
provide funding for wildlife
management (IUCN 2006a, p. 24). The
revenue generated from these industries
can be critical to fund wildlife
management programs in range states.
Tourism, through ecotourism and
trophy hunting, can provide jobs to
locals (such as game guards, cooks,
drivers, security personnel) and often
brings in revenue for local
microbusinesses that sell art, jewelry,
and other native crafts. Lions can
generate the highest daily rate of any
mammal hunted (USD $2,650 per day),
the longest number of days that must be
booked, and the highest trophy fee
($24,500) (Jackson 2013, p. 6; Lindsey et
al. 2012a, p. 5), thus generating
significant revenue for range countries.
Creating community-based incentives to
conserve lions from revenue derived
from trophy hunting may ameliorate the
human-lion conflict that arises from
lions and humans coexisting in the
same area.
Revenue from scientifically based
management programs that include
trophy hunting can increase the ability
of many African countries to manage
wildlife populations both within and
adjacent to reserves; many of these
hunting areas are geographically linked
to national parks and reserves,
providing wildlife corridors and buffer
zones (Chardonnet et al. 2010, p. 34;
Newmark 2008, p. 321). In the past,
government and private land owners
were the primary beneficiaries of the
revenue gained; however, a portion of
the revenue derived from hunting is
reportedly now being distributed to
local communities, creating a value for
lions that encourages their conservation
(Barnett & Patterson 2005, p. iv).
Revenue from trophy hunting is
purported to create: (1) Incentives for
countries to conserve large tracts of
prime habitat; and (2) funding for park
and reserve management, anti-poaching,
and security activities. Because habitat
loss has been identified as one of the
primary threats to lion populations, it is
notable that trophy hunting has
provided lion range states incentives to
set land aside for hunting throughout
Africa, and the land set aside exceeds
the total area of the national parks,
accounting for approximately half of the
amount of viable lion habitat
(Chardonnet et al. 2010, p. 34; Packer et
al. 2006, pp. 9–10).
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In Tanzania, which is home to 40
percent of all lions, land set aside for
sport hunting purposes has resulted in
an area 5.1 times greater than Tanzania’s
fully protected and gazetted parks
(Jackson 2013, p. 6; Barnett & Patterson
2005, p. 61). In Botswana, despite the
current ban on lion hunting, the country
currently has more than 128,000 km2 of
gazetted wildlife management areas and
controlled hunting areas set aside for
hunting purposes, which equates to 22.1
percent of the country’s total area; this
is in addition to 111,000 km2 (or 19.1
percent) that has been set aside as
habitat in the form of National Parks,
Game Reserves, and Forest Reserves
(Barnett & Patterson 2005, p. 7). In 2000,
five countries in southern Africa
(Botswana, Namibia, South Africa,
Tanzania, and Zimbabwe) had set aside
a combined 420,000 km2 of communal
land, 188,000 km2 of commercial land,
and 420,089 km2 of state land totaling
more than 1,028,000 km2 for sport
hunting purposes (Barnett & Patterson
2005, p. iii). As a species with a
considerable range (up to 1,000 km2)
(Packer et al. 2013 p. 636; Haas et al.
2005, p. 4), suitable habitat is important
to the survival of the species, and the
marked decline in suitable habitat is a
significant threat to the species. The
habitat currently preserved for use in
sport hunting has helped to reduce the
impact of habitat loss for the African
lion, but as discussed previously,
habitat loss remains a significant threat
to the species.
Within its current range, the African
lion exists in 10 stronghold populations
containing approximately 24,000 lions
(70 percent of the current African lion
population), 19,000 of which are in
protected areas, and in 7 potential
stronghold populations containing
another 4,000 lions. Reports from the
IUCN Species Survival Commission Cat
Specialist Group (IUN 2006a, b)
characterize the population as
increasing in 3 of those strongholds, as
stable in 6 of the strongholds, and as
decreasing in 1 stronghold. Most lion
populations in protected areas of
southern and eastern Africa have been
essentially stable over the last three
decades (Ray et al. 2005, p. 67). In
contrast to the stronghold or potential
stronghold populations, other African
lion populations, containing a total of
more than 6,000 individuals, have a
very high risk of local extinction (Reggio
et al. 2013, p. 33. During the 2005–2006
African lion workshops, lion experts
characterized lion populations in 36 (42
percent) of the 86 LCUs as decreasing.
In extensive surveys recently conducted
within 15 of the 20 LCUs in western and
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central Africa, Henschel et al. (2010,
entire) were able to confirm lion
presence in only four. The work of
Packer et al. (2013) suggests future
declines within a number of protected
areas. Craigie et al. (2010, entire)
provide evidence of declining large
mammal populations in Africa’s
protected areas, indicating that
protected areas in Africa have generally
failed to mitigate threats to large
mammal populations, including African
lion. Although Craigie et al. (2010, p.
2,225) found large regional differences
(from large declines in western Africa to
positive rates of change in southern
Africa), they found overall populations
decreased steadily from 1970 to 2005.
The best available scientific and
commercial information leads us to
conclude that the African lion is in
danger of extinction within the
foreseeable future throughout all of its
range. Accordingly, we find that listing
is warranted and we propose to list it as
a threatened species throughout its
range, wherever found.
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Significant Portion of Its Range
Under the Act and our implementing
regulations, a species may warrant
listing if it is endangered or threatened
throughout all or a significant portion of
its range. The term ‘‘species’’ includes
‘‘any subspecies of fish or wildlife or
plants, and any distinct population
segment [DPS] of any species of
vertebrate fish or wildlife which
interbreeds when mature.’’ We
published a final policy interpreting the
phrase ‘‘Significant Portion of its
Range’’ (SPR) (79 FR 37578, July 1,
2014). The final policy states that (1) if
a species is found to be endangered or
threatened throughout a significant
portion of its range, the entire species is
listed as endangered or threatened,
respectively, and the Act’s protections
apply to all individuals of the species
wherever found; (2) a portion of the
range of a species is ‘‘significant’’ if the
species is not currently endangered or
threatened throughout all of its range,
but the portion’s contribution to the
viability of the species is so important
that, without the members in that
portion, the species would be in danger
of extinction, or likely to become so in
the foreseeable future, throughout all of
its range; (3) the range of a species is
considered to be the general
geographical area within which that
species can be found at the time FWS
or NMFS makes any particular status
determination; and (4) if a vertebrate
species is endangered or threatened
throughout an SPR, and the population
in that significant portion is a valid
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DPS, we will list the DPS rather than the
entire taxonomic species or subspecies.
We found the African lion to be in
danger of extinction within the
foreseeable future throughout all of its
range. Therefore, no portions of the
species’ range are ‘‘significant’’ as
defined in our SPR policy and no
additional SPR analysis is required.
Proposed 4(d) Rule
The purposes of the ESA are to
provide a means whereby the
ecosystems upon which endangered
species and threatened species depend
may be conserved, to provide a program
for the conservation of such endangered
species and threatened species, and to
take such steps as may be appropriate to
achieve the purposes of the treaties and
conventions set forth in the ESA. When
a species is listed as endangered, certain
actions are prohibited under section 9 of
the ESA, as specified in 50 CFR 17.21.
These include, among others,
prohibitions on take within the United
States, within the territorial seas of the
United States, or upon the high seas;
import; export; and shipment in
interstate or foreign commerce in the
course of a commercial activity.
The ESA does not specify particular
prohibitions and exceptions to those
prohibitions for threatened species.
Instead, under section 4(d) of the ESA,
the Secretary, as well as the Secretary of
Commerce depending on the species,
was given the discretion to issue such
regulations as deemed necessary and
advisable to provide for the
conservation of such species. The
Secretary also has the discretion to
prohibit by regulation with respect to
any threatened species any act
prohibited under section 9(a)(1) of the
ESA. Exercising this discretion, the
Service has developed general
prohibitions (50 CFR 17.31) and
exceptions to those prohibitions (50
CFR 17.32) under the ESA that apply to
most threatened species. Under 50 CFR
17.32, permits may be issued to allow
persons to engage in otherwise
prohibited acts for certain purposes.
Under section 4(d) of the ESA, the
Secretary, who has delegated this
authority to the Service, may also
develop specific prohibitions and
exceptions tailored to the particular
conservation needs of a threatened
species. In such cases, the Service issues
a 4 (d) rule that may include some or all
of the prohibitions and authorizations
set out in 50 CFR 17.31 and 17.32 but
which also may be more or less
restrictive than the general provisions at
50 CFR 17.31 and 17.32. For the African
lion, the Service has determined that a
4(d) rule is appropriate.
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We propose to add a 4(d) (special)
rule for the African lion (Panthera leo
leo) at 50 CFR 17.40(n). This 4(d) rule
would maintain all of the prohibitions
and exceptions codified in 50 CFR 17.31
and 17.32 and would supersede with
regard to African lion the import
exemption found in 50 CFR 17.8 for
threatened wildlife listed in Appendix II
of CITES, such that a threatened species
import permit under 50 CFR 17.32
would be required for the importation of
all African lion specimens. Through the
promulgation of the proposed 4(d) rule,
the presumption of legality provided
under Section 9(c)(2) of the Act for the
otherwise lawful importation of wildlife
listed in Appendix II of CITES that is
not an endangered species listed
pursuant to section 4 of the Act would
not apply to this subspecies. Thus,
under the proposed 4(d) rule, all
otherwise prohibited activities,
including all imports of African lion
specimens, would require prior
authorization or permits under the Act.
Under our regulations, permits or
authorization to carry out an otherwise
prohibited activity could be issued for
scientific purposes, the enhancement of
propagation or survival of the species,
economic hardship, zoological
exhibitions, educational purposes, or
special purposes consistent with the
purposes of the Act. Applications for
these activities are available from
https://www.fws.gov/forms/3-200-37.pdf.
The intent of this proposed 4(d) rule
is to provide for the conservation of the
African lion consistent with the
purposes of the Act. Under the proposed
4(d) rule, the prohibitions would, in
part, make it illegal for any person
subject to the jurisdiction of the United
States to ‘‘take’’ (includes harass, harm,
pursue, hunt, shoot, wound, kill, trap,
capture, or to attempt any of these)
within the United States or upon the
high seas; import or export; deliver,
receive, carry, transport, or ship in
interstate or foreign commerce, by any
means whatsoever, in the course of
commercial activity; or sell or offer for
sale in interstate or foreign commerce
any lion specimens. It would also be
illegal to possess, sell, deliver, carry,
transport, or ship any such wildlife that
has been taken in violation of the Act.
We believe that these protections,
including the requirement for an import
permit for all African lion specimens,
will support and encourage
conservation actions for the African lion
and require that permitted activities
involving lions are carried out in a
manner that is consistent with the
purposes of the Act and our
implementing regulations.
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In connection with this proposed 4(d)
rule, the Service notes that the African
lion is listed in Appendix II of CITES,
and thus can be imported into the U.S.
pursuant to Section 9(c)(2) of the Act
and upon presentation of a proper
CITES export permit from the country of
origin. Section 9(c)(2) of the Act
provides that the otherwise lawful
importation of wildlife that is not an
endangered species listed pursuant to
section 4 of the Act, but that is listed in
Appendix II of CITES, shall be
presumed to be in compliance with
provisions of the Act and implementing
regulations. While there has been
question as to whether this provision of
the Act might automatically require
allowing the importation of a species
that is both listed as threatened and in
Appendix II, and preclude the issuance
of more restrictive 4 (d) rules covering
importation, the Service has concluded
that such 4 (d) rules may be issued to
provide for the conservation of the
involved species. Section 9(c)(2) does
not expressly refer to threatened species
or prevent the issuance of appropriate 4
(d) rules and could not logically have
been intended to allow the addition of
a species to an appendix of an
international convention to override the
needs of U.S. law, where there is
reliable evidence to affect the
presumption of validity. Finally, the
term ‘‘presumed’’ implies that the
established presumption is rebuttable
under certain circumstances, including
through the promulgation of a protective
regulation pursuant to section 4(d) of
the Act.
In the case of the African lion, there
are substantive grounds on which to
challenge the presumption. For the
import of sport-hunted trophies, while
there is evidence that many of the range
countries are implementing lion
management plans, we want to
encourage and support efforts by these
countries to develop plans that are
based on sound scientific information.
As noted, the proposed 4(d) rule for
African lion would provide for the
importation into the United States of
trophies taken legally in range countries
upon the issuance of a threatened
species import permit. While the
Service cannot control hunting of
foreign species such as African lion, we
can regulate their importation and
thereby require that U.S. imports of
sport-hunted African lion trophy
specimens are obtained in a manner that
is consistent with the purposes of the
Act and the conservation of the
subspecies in the wild, by allowing
importation from range countries that
have management plans that are based
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on scientifically sound data and are
being implemented to address the
threats that are facing lions within that
country.
Such management plans would be
expected to address, but are not limited
to, evaluating population levels and
trends; the biological needs of the
species; quotas; management practices;
legal protection; local community
involvement; and use of hunting fees for
conservation. In evaluating these
factors, we will work closely with the
range countries and interested parties to
obtain the best available scientific and
commercial data. By allowing entry into
the United States of African lion
trophies from range countries that have
scientifically based management plans,
the range countries would be
encouraged to adopt and financially
support the sustainable management of
lions that benefits both the species and
local communities. In addition to
addressing the biological needs of the
subspecies, a scientifically based
management plan would provide
economic incentives for local
communities to protect and expand
African lion habitat.
As stated, anyone wishing to conduct
any otherwise prohibited activity, such
as interstate commerce or imports, must
first obtain a permit under the current
permitting regulations found at 50 CFR
13 and 50 CFR 17. As will all permits,
the individual requesting authorization
to carry out an otherwise prohibited
activity under the Act must submit a
permit application to the Service with
specific information concerning the
proposed activity and the benefits/
impacts of the activity on the species. In
some cases, such as imports of sporthunted trophies, it is not always
possible for the applicant to provide all
of the necessary information needed by
the Service to make a positive
determination under the Act to
authorize the activity. For the import of
sport-hunted trophies, it is typical for
the Service to consult with the range
country and other interested parties to
obtain the necessary information. To
date, the Service typically has made the
required findings on sport-hunted
trophy imports on a country-wide basis,
although individual import permits are
issued for each applicant. While the
Service encourages the submission of
information from individual applicants,
we would primarily rely on information
from other sources when making a
permitting decision.
Effects of This Rule
This rule, if made final, would revise
50 CFR 17.11(h) to add the African lion
to the List of Endangered and
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Threatened Wildlife. This rule, if
adopted, would also establish a 4(d) rule
for the African lion, which implements
all of the prohibitions and exceptions
under 50 CFR 17.31 and 17.32 and
requires a threatened species import
permit under 50 CFR 17.32 for the
importation of all African lion
specimens. Under the proposed 4(d)
rule, the import exemption found in 50
CFR 17.8 for threatened wildlife listed
in Appendix II of CITES would not
apply to this subspecies. Through the
promulgation of the proposed 4(d) rule,
the presumption of legality provided
under Section 9(c)(2) of the Act for the
otherwise lawful importation of wildlife
listed in Appendix II of CITES that is
not an endangered species listed
pursuant to section 4 of the Act would
not apply to this subspecies. (See:
Proposed Special Rule section).
Available Conservation Measures
Conservation measures provided to
species listed as endangered or
threatened under the Act include
recognition of conservation status,
requirements for Federal protection, and
prohibitions against certain practices.
Recognition through listing encourages
and results in public awareness and
conservation actions by Federal and
State governments in the United States,
foreign governments, private agencies
and groups, and individuals.
Section 7(a) of the Act, as amended,
and as implemented by regulations at 50
CFR part 402, requires Federal agencies
to evaluate their actions that are to be
conducted within the United States or
upon the high seas, with respect to any
species that is proposed to be listed or
is listed as endangered or threatened.
Because the African lion is not native to
the United States, no critical habitat is
being proposed for designation with this
rule. Regulations implementing the
interagency cooperation provision of the
Act are codified at 50 CFR part 402.
Section 7(a)(2) of the Act requires
Federal agencies to ensure that activities
they authorize, fund, or carry out are not
likely to jeopardize the continued
existence of a listed species or to
destroy or adversely modify its critical
habitat. If a proposed Federal action
may adversely affect a listed species, the
responsible Federal agency must enter
into formal consultation with the
Service. Currently, with respect to the
African lion, no Federal activities are
known that would require consultation.
Section 8(a) of the Act authorizes the
provision of limited financial assistance
for the development and management of
programs that the Secretary of the
Interior determines to be necessary or
useful for the conservation of
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endangered or threatened species in
foreign countries. Sections 8(b) and 8(c)
of the Act authorize the Secretary to
encourage conservation programs for
foreign listed species, and to provide
assistance for such programs, in the
form of personnel and the training of
personnel.
Section 9 of the Act and its
implementing regulations at 50 CFR
17.31 set forth a series of general
prohibitions that apply to all threatened
wildlife, except where a 4(d) rule
applies, in which case the 4(d) rule will
contain all the applicable prohibitions
and exceptions. If the 4(d) rule is
adopted as proposed, these prohibitions
would apply to the African lion. These
prohibitions, in part, make it illegal for
any person subject to the jurisdiction of
the United States to ‘‘take’’ (includes
harass, harm, pursue, hunt, shoot,
wound, kill, trap, capture, or to attempt
any of these) within the United States or
upon the high seas; import or export;
deliver, receive, carry, transport, or ship
in interstate or foreign commerce, by
any means whatsoever, in the course of
commercial activity; or sell or offer for
sale in interstate or foreign commerce
any lion specimens. It also is illegal to
possess, sell, deliver, carry, transport, or
ship any such wildlife that has been
taken in violation of the Act. Permits
may be issued to carry out otherwise
prohibited activities involving
threatened wildlife species under
certain circumstances. Certain
exceptions apply to agents of the
Service and State conservation agencies.
Required Determinations
National Environmental Policy Act (42
U.S.C. 4321 et seq.)
We have determined that we do not
need to prepare an environmental
assessment, as defined under the
authority of the National Environmental
Policy Act of 1969, in connection with
regulations adopted under section 4(a)
of the Act. We published a notice
outlining our reasons for this
determination in the Federal Register
on October 25, 1983 (48 FR 49244).
References Cited
A list of all references cited in this
document is available at https://
www.regulations.gov at Docket No.
FWS–R9–ES–2012–0025, or upon
request from the U.S. Fish and Wildlife
Service, Endangered Species Program,
Species
Historic range
Common name
Vertebrate
population where
endangered or
threatened
*
Africa ......................
*
Entire ......................
Scientific name
Branch of Foreign Species (see FOR
FURTHER INFORMATION CONTACT).
Authors
The primary authors of this proposed
rule are staff of the Branch of Foreign
Species, Ecological Services, U.S. Fish
and Wildlife Service.
Proposed Regulation Promulgation
For the reasons described in the
preamble, we propose to amend part 17,
subchapter B of chapter I, title 50 of the
Code of Federal Regulations, as follows:
PART 17—[AMENDED]
1. The authority citation for part 17
continues to read as follows:
■
Authority: 16 U.S.C. 1361–1407; 1531–
1544; and 4201–4245; unless otherwise
noted.
2. In § 17.11(h), the List of Endangered
and Threatened Wildlife, add an entry
for ‘‘Lion, African’’ under Mammals to
read as follows:
■
§ 17.11 Endangered and threatened
wildlife.
*
*
*
(h) * * *
Status
*
When
listed
*
Critical
habitat
Special
rules
MAMMALS
*
Lion, African .............
*
*
Panthera leo leo .....
*
*
*
*
*
*
*
3. Amend § 17.40 by adding paragraph
(n) to read as follows:
■
§ 17.40
Special rules—mammals.
*
*
*
*
(n) African lion (Panthera leo leo).
(1) General requirements. All
prohibitions and provisions of §§ 17.31
and 17.32 of this part apply to this
subspecies.
mstockstill on DSK4VPTVN1PROD with PROPOSALS2
*
VerDate Sep<11>2014
16:29 Oct 28, 2014
Jkt 235001
*
*
T
*
(2) The import exemption found in
§ 17.8 of this part for threatened wildlife
listed in Appendix II of the Convention
on International Trade in Endangered
Species of Wild Fauna and Flora
(CITES) does not apply to this
subspecies. A threatened species import
permit under § 17.32 of this part is
required for the importation of all
African lion specimens.
PO 00000
Frm 00032
Fmt 4701
*
....................
Sfmt 9990
*
NA
*
17.40(n)
*
(3) All applicable provisions of 50
CFR parts 13, 14, 17, and 23 must be
met.
*
*
*
*
*
Dated: October 20, 2014.
Stephen Guertin,
Director, Fish and Wildlife Service.
[FR Doc. 2014–25731 Filed 10–28–14; 8:45 am]
BILLING CODE 4310–55–P
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[Federal Register Volume 79, Number 209 (Wednesday, October 29, 2014)]
[Proposed Rules]
[Pages 64471-64502]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2014-25731]
[[Page 64471]]
Vol. 79
Wednesday,
No. 209
October 29, 2014
Part II
Department of the Interior
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Fish and Wildlife Service
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50 CFR Part 17
Endangered and Threatened Wildlife and Plants; Listing the African Lion
Subspecies as Threatened With a Rule Under Section 4(d) of the ESA;
Proposed Rule
Federal Register / Vol. 79 , No. 209 / Wednesday, October 29, 2014 /
Proposed Rules
[[Page 64472]]
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DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R9-ES-2012-0025; 450 003 0115]
RIN 1018-BA29
Endangered and Threatened Wildlife and Plants; Listing 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: https://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 https://www.regulations.gov. If you submit information via https://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
[[Page 64473]]
guarantee that we will be able to do so. Please include sufficient
information with your comments to allow us to verify any scientific or
commercial information you include.
Comments and materials we receive, as well as supporting
documentation we used in preparing this proposed rule, will be
available for public inspection on https://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 https://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 https://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; https://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
[[Page 64474]]
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) (https://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)] (https://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 (https://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); https://www.cites.org/common/cop/16/sum/E-CoP16-Plen-06.pdf;
https://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
[[Page 64475]]
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.
[[Page 64476]]
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
----------------------------------------------------------------------------------------------------------------
Current range as percent of
Historic Current range historic range (percent of
Source Region of Africa range (km\2\) (km\2\) 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 + 12,080,000 3,915,000 32 percent.
Eastern.
--------------------------------
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 + 13,010,000 3,564,000 23 percent.
Eastern. (58 percent).
--------------------------------
Total............ 20,216,817 4,611,231 22 percent.
(38 percent).
Riggio 2013 \3\ (based on Western.......... .............. 133,784 ............................
estimates of savannah Central.......... 936,465
habitat):.
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
[[Page 64477]]
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]
--------------------------------------------------------------------------------------------------------------------------------------------------------
Western Africa Central Africa Eastern Africa Southern Africa
Source (percent of total) (percent of total) (percent of total) (percent of total) Total
--------------------------------------------------------------------------------------------------------------------------------------------------------
Ferreras & Cousins 1996 (estimate ...................... ...................... ..................... ..................... 75,800 (18,600 in
for lion abundance in 1980). protected areas).
Chardonnet 2002.................... 1,163 (3 percent)..... 2,815 (7 percent)..... 15,744 (40 percent).. 19,651 (50 percent).. 39,373.
Bauer & Van Der Merwe 2004......... 850 (4 percent)....... 950 (4 percent)....... 11,000 (48 percent).. 10,000 (44 percent).. 23,000.
IUCN 2006 \4\ (as calculated by 1,640 (5 percent)..... 2,410 (7 percent)..... 17,290 (52 percent).. 11,820 (37 percent).. 33,160.
Riggio et al. 2013).
Riggio 2013 (based on estimates of 480 (1 percent)....... 2,419 (7 percent)..... 19,972 (57 percent).. 12,036 (34 percent).. 34,907.
savannah habitat).
Henschel et al. 2014............... 406 (n/a).............
--------------------------------------------------------------------------------------------------------------------------------------------------------
In 2005-2006, in response to a growing concern that the African
lion was in decline, IUCN and the Wildlife Conservation Society
sponsored workshops to determine a lion conservation strategy. During
these workshops, lion experts collectively assessed what they believed
to be the then-current status of African lions based on a variety of
information, including professional opinion. During the workshops, lion
experts identified 86 African lion Conservation Units (LCUs). They
defined LCUs as areas of known, occasional, or possible lion range that
can be considered an ecological unit of importance for lion
conservation (IUCN 2006a, p. 14; IUCN 2006b, p. 17). Of the 86 LCUs, 20
are in western and central Africa and 66 are in southern and eastern
Africa (Table 4). Most (71 percent) have more than half their area
under some form of legal protection (Bauer et al. 2008, p. 19). Few (16
percent) were estimated to contain large populations (Table 4). This
was particularly the case for western and central Africa, where most
(13, or 65 percent) of LCUs were estimated to contain fewer than 50
lions (Table 4). The majority of those with large populations were in
southern and eastern Africa (Table 4). Only 23 of 86 LCUs (27 percent)
were considered to contain viable populations, though more than half
were thought to contain potentially viable populations (Table 4). Lion
populations within 42 percent of the 86 LCUs were considered to be
decreasing, whereas those in 9 percent were considered increasing. The
remaining were considered stable or of unknown trend (Table 4).
Table 4--Lion Conservation Units (LCUs) as Identified and Characterized in IUCN 2006a and IUCN 2006b
----------------------------------------------------------------------------------------------------------------
Eastern &
Number of LCUs Western & Southern All regions (percent)
Central Africa Africa
----------------------------------------------------------------------------------------------------------------
Total................................... 20 66 86.
Estimated to contain:
>500 lions.......................... 2 12 14 (16 percent).
50-500 lions........................ 5 28 33 (38 percent).
[[Page 64478]]
<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 Bj[ouml]rklund (2003) on
the number of prides needed to avoid the risk of inbreeding in lion
populations, and information in Bauer et al. (2008) on the average size
of lion prides. Bj[ouml]rklund (in Riggio et al. 2013, p. 32) assessed
the risk of inbreeding due to habitat loss and determined that, ``. . .
to sustain a large out-bred population of lions, a continuous
population of at least 50 prides, but preferably 100 prides, with no
limits to dispersal is required.'' Bauer et al. 2008 (in Riggio et al.
2013, p. 32) indicate the average lion pride as containing
approximately five adults.
The results of Riggio et al. indicate the size of the African lion
population to be about 35,000, which falls within the range of the
other recent estimates (Table 3). However, they state that ``Although
these numbers are similar to previous estimates, they are
geographically more comprehensive. There is abundant evidence of
widespread declines and local extinctions'' (Riggio et al. 2013, p.
18).
Riggio et al. identified lions as occurring in 67 areas (Table 5).
While a small portion (22 percent) of lion areas identified by Riggio
et al. contain large populations, the majority are small and isolated
(Riggio et al. 2013, p. 30; Table 5). Most (69 percent) contain fewer
than 250 lions. A considerable portion (39 percent) contains very small
populations of fewer than 50 lions. These include 63 percent of the
lion areas in western and central Africa, and 31 percent of those in e/
s Africa.
Table 5--Number of Lion Areas and Number of Areas Containing Lion Population Classes According to Riggio et al. 2013
--------------------------------------------------------------------------------------------------------------------------------------------------------
Number of lion areas Western Central Eastern Southern All regions (percent)
--------------------------------------------------------------------------------------------------------------------------------------------------------
Total....................................... 8 8 28 23 67.
# Estimated to contain:
>=500 lions............................. 0 1 7 7 15 (22 percent).
250-499 lions........................... 1 2 1 2 6 (9 percent).
50-249 lions............................ 0 2 12 6 20 (30 percent).
<50..................................... 7 3 8 8 26 (39 percent).
--------------------------------------------------------------------------------------------------------------------------------------------------------
Riggio et al. identify 10 lion strongholds (viable populations) and
7 potential strongholds (Table 6). According to Riggio et al. (2013, p.
29), the 10 strongholds contain approximately 24,000 lions, or about 70
percent of the current African lion population. Of those, most (about
19,000 lions) are in protected areas. Potential strongholds contain
about 4,000 lions. More than 6,000 lions are located in areas not
considered strongholds or potential strongholds and have a very high
risk of being extirpated (Riggio et al. 2013, p. 33).
[[Page 64479]]
Table 6--Lion Strongholds and Potential Strongholds Identified by Riggio et al. 2013
--------------------------------------------------------------------------------------------------------------------------------------------------------
Population
Lion size in IUCN (2006a, b)
Lion area Country Area (km\2\) Stronghold population protected Trend
size areas
--------------------------------------------------------------------------------------------------------------------------------------------------------
Western Africa
--------------------------------------------------------------------------------------------------------------------------------------------------------
W-Arly-Pendjari................... Benin, Burkina Faso, 29,403 Potential............. 350 350 Stable.
Niger.
--------------------------------------------------------------------------------------------------------------------------------------------------------
Central Africa
--------------------------------------------------------------------------------------------------------------------------------------------------------
SE Chad........................... Chad................. 133,408 Potential \5\......... 400 140 Stable.
E CAR............................. Central African 328,721 Potential \6\......... 1,244 148 Stable.
Republic.
--------------------------------------------------------------------------------------------------------------------------------------------------------
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 150,347 Stronghold............ 2,311 2,179 Increasing.
Africa, Zimbabwe.
Kgalagadi......................... Botswana, South 163,329 Stronghold............ 800 ~ 800 Stable.
Africa.
Luangwa........................... Malawi, Zambia....... 72,992 Stronghold............ 574 574 Stable.
Mid-Zambezi....................... Mozambique, Zambia, 64,672 Stronghold............ 755 ~ 350-650 Stable.
Zimbabwe.
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
[[Page 64480]]
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
[[Page 64481]]
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
[[Page 64482]]
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;
[[Page 64483]]
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
[[Page 64484]]
(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
[[Page 64485]]
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
[[Page 64486]]
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
[[Page 64487]]
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).
[[Page 64488]]
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
[[Page 64489]]
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. The recommended best practices
include: (1) Establishing processes and procedures that are clearly
outlined, transparent, and accountable; (2) establishing processes and
procedures that are CITES compliant;(3) demonstrating management
capacity; (4) standardizing information sources; (5) establishing
monitoring systems for critical data; (6) recording and analyzing
trophy hunting data; (7) conducting data collection and analysis for
each hunting block and concession; and (8) establishing a primary body
who will approve quotas (Burnett and Patterson 2005, p. 103). We have
no information on whether these best practices have been implemented by
the lion range states. However, most countries that allow trophy
hunting of lions appear to be reviewing their trophy hunting practices
(Jackson 2013, pp. 2-3; White 2013, pp. 12-13). Benin halved their
quotas in 2002 after the first population census of lions was conducted
and resulted in the current quota of six lions every 2 years in
Pendjari and four lions every 2 years in western Benin or one lion
annually in each of the five hunting zones. This was largely due to
impacts to lions from habitat degradation and fragmentation
(particularly exacerbated by the increase of human population), loss of
prey by poaching, trade (both legal and illegal), and human-lion
conflict. (CITES 2014a, p. 5; Sogbohossou 2014, p. 1).
Throughout the countries in Africa, most appear to have reduced
their offtake considerably since the 1990's. According to Packer et al.
(2006, pp. 2-3), regardless of population estimates, countries are
allowing for only a small proportion of their lion populations to be
hunted, with most countries ranging from 2-4 percent annually
(excluding offtake from South Africa, where offtake has been increasing
from the trophy hunting of primarily captive-born lions, and Zimbabwe,
where offtake was 2-3 percent higher than other countries from 1998-
2004.
Regardless of these reductions, many stakeholders consider the
quota system to be outdated and ineffective because it does not address
the biological and social impacts of trophy hunting on lion prides.
Opponents also state that trophy hunting affects the social structure
of the pride and results in increased infanticide of lion cubs. This
supposition is inconclusive and not well supported (CITES 2014a, p. 14;
Dagg 2000, pp. 831-835) (See Infanticide and Age-based Hunting
Strategies). Regardless, since 2006, researchers have recommended the
implementation of age-based hunting strategies; these are discussed
below (Packer et al. 2006, pp. 6-8).
Five countries maintain quotas to allow for approximately 6-15 lion
trophies to be taken per year: Benin, Burkina Faso, Cameroon,\7\
Mozambique, and Namibia. Tanzania allows the take of approximately 50
lions annually, and Zimbabwe allows approximately 70 animals annually
to be taken (Jackson 2013 pp. 7-8, CITES WCMC-UNEP trade database,
accessed December 2013). In Ethiopia and Uganda, trophy hunting is
restricted to problem or dangerous animals only (Lindsey 2008, p. 42),
and Botswana and Zambia currently ban all trophy hunting (CITES 2014a,
p.14). South Africa has not set a quota for the take of wild lions
since 99 percent of the trophy-hunted lions are reportedly not of wild
origin, but captive-born (Hunter et al. 2013, p. 2; RSA 2013, pp. 5,
7).
Below is a summary of estimated annual hunting quotas for the
African lion:
Table 7--Annual Trophy Quotas (Approximate) as of 2013
------------------------------------------------------------------------
Annual lion
trophy quotas
Country (Jackson 2013,
pp. 7-8)
------------------------------------------------------------------------
Benin.................................................. 6
Botswana (moratorium).................................. 30
Burkina Faso........................................... 6
Cameroon \7\........................................... 6
Mozambique............................................. 15
Namibia................................................ 10
Tanzania (as of 2012).................................. 50
[[Page 64490]]
Zambia (moratorium).................................... 50
Zimbabwe............................................... 70
------------------------------------------------------------------------
Import/Export of Lion Trophies
Although each country has its own method of regulating trophy
hunting, international trade of lion trophies must adhere to CITES (see
Conservation Status). International trade of lion parts and products
(including trophies) are reported by both the exporting and importing
countries and tracked by the United Nations Environment Programme World
Conservation Monitoring Centre (UNEP-WCMC). The international trade
data on the African lion that has been compiled in the CITES UNEP-WCMC
Trade Database is extensive. Therefore, it is likely that the actual
numbers of African lion parts and products in international trade is
slightly smaller than what we have reported using the UNEP-WCMC ``gross
exports'' report (CITES lion gross exports, https://trade.cites.org,
accessed April 23, 2014).
In 2012, the most recent year for which CITES trade data are
available, U.S. CITES Annual Report trade data indicated that the
United States allowed the direct import of African lion trophies from
eight African countries, as follows:
Central African Republic = 1 trophy
Ethiopia = 1 trophy
Mozambique = 5 trophies
Namibia = 5 trophies
South Africa = 413 trophies (the majority of which are reported to
be of captive-born origin)
Tanzania = 42 trophies
Zambia = 32 trophies
Zimbabwe = 49 trophies
According to the CITES UNEP-WCMC database, between 2005 and 2012,
exports of lion trophies have demonstrated a decreasing trend when
exports of captive-born lions from South Africa are excluded (CITES
lion gross exports, https://trade.cites.org, accessed April 23, 2014).
For example, in 2005 there were 874 lion trophy exports reported in
UNEP-WCMC, 521 if South Africa were excluded; whereas in 2012, there
were 1,237 lion trophy exports reported in UNEP-WCMC, 336 if South
Africa is excluded.
Here it should be noted that there are limitations to interpreting
the above reported information. The 2004 guide to using the CITES Trade
Database indicates that the outputs produced by the CITES Trade
Database can be easily misinterpreted if one is not familiar with it
(CITES 2004b, p. 5). The number of ``trophies'' reported does not
necessarily equate to the number of lions hunted. Additionally, the
number of trophies reported for a given year in the trade report does
not equate directly to the number of animals hunted in that given year
(CITES export permits may be valid for 6 months, and a trophy could in
theory be exported the year after it was hunted). The second limitation
to interpreting this information is, although many permits may indicate
that an animal is of wild origin (source code ``W''), these permits may
be incorrectly coded. This is true for South Africa, where during the
period of 2000 to 2009, animals that were captive-born and released
into private reserve systems were assigned an incorrect source code of
``wild.'' South Africa has since requested their provincial authorities
to use the correct source code for ``captive bred'' in order to
correctly reflect the source of sport-hunted lion trophies; however,
some provinces are still not complying (RSA 2013, pp. 8-9). However,
based on South African trade data, the bulk of the exports of lions and
their parts and products (including trophies) from South Africa were
from captive-born lions (RSA 2013, p. 7).
Tanzania, with the highest lion populations (Hamunyela et al. 2013,
pp. 29, 283; Riggio et al. 2013, p. 32; Ikanda 2008, p. 4; Baldus 2004,
pp. 5, 6), was the largest exporter of wild-origin lion trophies, but
their exports have decreased significantly since 2006. In 2008,
approximately 138 lions had been estimated to be killed in Tanzania as
trophies. In 2010, Tanzania's numbers declined to 128 exports, 55 in
2011, and 42 in 2012 (CITES lion gross exports, https://trade.cites.org/
, accessed April 25, 2014). In 2012, Tanzania established an annual
quota to limit trophy hunting to no more than 50 animals (Jackson 2013,
p. 7). Again, it should be noted that there may be discrepancies
between the annual quota and the actual number of trophies exported in
a given year (see https://www.cites.org/common/resources/TradeDatabaseGuide.pdf for additional information). Regardless, the
numbers of lion trophies exported by Tanzania according to the UNEP-
WCMC database suggest a decreasing trend.
In other areas within the range of the African lion, the number of
lions hunted or authorized to be hunted annually has remained fairly
consistent. In Burkina Faso, approximately 12 lions per year have been
hunted over the past two decades (IUCN 2009, pp. 36-37; Bauer and
Nowell 2004, p. 36), although their current annual quota is 6 animals.
In Botswana, a quota of 30 lions per year was authorized for nearly two
decades; however, Botswana has recently implemented a hunting
moratorium (Jackson 2013, p. 8). (CITES lion gross exports, https://trade.cites.org, accessed April 23, 2014; CITES UNEP-WCMC database,
accessed January 8, 2014, and August 16, 2013).
Potential Impacts of Trophy Hunting
Infanticide and Age-Based Hunting Strategies
Tourist safari hunting of males has been suggested by the
petitioners to increase infanticide rates (when males kill young lion
cubs sired by other males) (Petition 2011, p. 24; Whitman et al. 2004,
p. 175), due in part to trophy hunters taking males under a certain
age. Removing a younger male lion is purported to allow another male to
take over the pride, and kill the former patriarch's cubs. This
supposition is inconclusive and not well supported (CITES 2014a, p. 14;
Dagg 2000, pp. 831-835). Infanticide is a common practice among many
species, including lions (Hausfater et al. 1984, pp. 31, 145, 173,
487). When an adult male lion in a pride is killed, surviving males who
form the pride's coalition become vulnerable to takeover by other male
coalitions, and this often results in injury or death of the defeated
males (Davidson et al. 2011, p. 115). In some cases, replacement males
who take over the pride will kill all cubs less than 9 months of age in
the pride (Whitman et al. 2004, p. 175). One range country specifically
addressed this issue; the Republic of Namibia indicates that lion
populations reproduce at similar rates in both harvested and non-
harvested populations, but it is unclear whether cub survival is
consistent in harvested vs. non-harvested lion populations.
While utilizing individual-based simulation models, Whitman et al.
(2004, pp. 175-177) found that if offtake is restricted to males older
than 6 years of age, then trophy hunting will likely have minimal
impact on the pride's social structure and young (Packer et al. 2006,
p. 6). This 6-year age restriction approach for lion trophies is in the
process of being self-implemented, along with other best practices, by
professional hunting guides, and is being adopted by certain range
states (White 2013, p. 14; Davidson et al. 2011, p. 114; Whitman et al.
2004, p. 176). It involves conducting an age assessment of male lions
using identification
[[Page 64491]]
techniques, such as mane development, facial markings, nose
pigmentation and tooth-aging, to establish the relative age of male
lions. Tooth wear on incisors, yellowing and chipping of teeth, coupled
with scars, head size, mane length and color, and thinning hair on the
face, as well as other factors can be an indicator of advanced age in
lions (Whitman and Packer 2006, entire). Although these characteristics
may be subjective, as regional differences may occur between lion
populations, there are clear attempts by the trophy hunting community
to establish and implement best practices. Promoting the removal of
males 6 years of age or older, theoretically allows younger males the
opportunity to remain resident long enough to rear a cohort of cubs
(allowing their genes to enter the gene pool; increasing the overall
genetic diversity). By removing males in a manner that promotes healthy
population growth, the lion population could yield more males in the
long term (Davidson et al. 2011, p. 114; Whitman et al. 2004, p. 176).
The governments of Tanzania, western Zimbabwe, Mozambique in the Niassa
National Reserve, Zambia, and most recently Benin have instituted or
are in the process of instituting reforms such as 6-year age
restrictions on lion trophies to increase the likelihood that trophy
hunting of lion is sustainable in those countries (Van der Merwe 2013,
p. 2; Jackson 2013, p. 3; White 2013, p. 14; Dallas Safari Club 2013,
pp. 1-2; Hunter et al. 2013, p. 2).
In addition to quota-setting, moratoriums, and the 6 year age
limit, it has been reported that more protective standards and
guidelines are implemented, such as the best practices listed below
(Jackson 2013, pp. 3, 8-10, Dallas Safari Club 2013, pp. 1-2).
Minimum trophy quality, sizes, and standards;
Wildlife hunting regulations enacted and enforced;
Professional hunting associations formed;
Professional hunting training courses;
Professional hunter standards established;
Quota-setting procedures;
Compliance with CITES demonstrated;
Monitoring; and
Information and data collection and analysis.
While the supposition of increased infanticide due to the remove of
established males from a pride is inconclusive and not well supported,
it is clear that improved management practices are beneficial to
maintaining viable lion populations. Developing and implementing best
management practices, while not categorically establishing a direct
correlation with increased population numbers and health, do appear to
have practical impacts on lion populations. Based on the best available
scientific and commercial information, infanticide, as a result of the
removal of lions through hunting, is not a threat to African lions.
Further, it is not likely to become a threat in the foreseeable future
since the science is not well supported as to whether infanticide
resulting from offtake due to trophy hunting is a significant threat to
the subspecies (Whitman et al. 2004, pp. 175-176; CITES 2014a, p. 14).
Corruption
Corruption is common in some areas within the range of the African
lion, particularly in areas with extreme poverty (Michler 2013, pp. 1-
3; Kimati 2012, p. 1; Garnett et al. 2011, p. 1; IUCN 2009, p. 89;
Leader-Williams et al. 2009, p. 296-298; Kideghesho 2008, pp. 16-17;
https://www.transparency.org). Several of the range countries of African
lion have experienced political instability for many years, which
appears to be a contributing factor in intensifying levels of
corruption. Political instability results in war and famine, which
essentially halt conservation efforts and the enforcement of existing
wildlife protection laws (Barnett & Patterson 2005, p. 82). Corruption
manifests itself in several ways, including embezzlement of funds and
acceptance of bribes to overlook illegal activities or for political
influence (Garnett et al. 2011, p. 1). Given the financial aspects of
sport hunting, it is reasonable to assume that corruption and the
inability to control it could have a negative impact on decisions made
in lion management by overriding biological rationales with financial
concerns.
Corruption has complex roots and will not end immediately, but from
all appearances, it is being addressed in many of the African lion
range countries where it has occurred in the past. Countries throughout
the range of the African lion are putting tools in place to combat
corruption and create awareness (https://www.transparency.org/cpi2012/results, accessed June 20, 2013). In recent years, in several African
lion range countries, leadership has taken steps to address corruption,
or activities that facilitate corruption, associated with wildlife
management. For example, in 2013, the Tourism Minister of Zambia banned
hunting in 19 game management areas for 1 year due to corruption and
malpractice among the hunting companies and various government
departments. Some game management areas and privately owned game
ranches were not included in the ban, but lion hunting appears to be
currently prohibited throughout the country (Michler 2013, pp. 1-3).
According to some authors (Martin 2012, pp. 4, 104; Kimati 2012, p. 1;
Kideghesho 2008, pp. 16-17), corruption in the wildlife sector has
often been one of the most discussed topics in Tanzania's National
Assembly, which presumably would indicate the awareness of and
willingness to address the corrupting factors in the wildlife sector.
Provided that countries continue to address corruption within the
wildlife sector, we conclude, based on the best scientific and
commercial information available, that corruption, in and of itself,
does not currently pose a threat to the species. However, if efforts to
address corruption do not continue, it could become a threat to African
lions in the future.
Revenue From Trophy Hunting
The high value of lions makes them one of the most expensive large
game species to hunt. The revenue derived from lion hunting is
substantial. Lions are reported to generate the highest daily rate of
any mammal hunted (USD $2,650 per day), the longest number of days that
must be booked, and the highest trophy fee ($24,500) (Jackson 2013, p.
6; Lindsey et al. 2012a, p. 5). According to Groom (2013, p. 4), a 21-
day lion hunt in Zimbabwe may be sold for approximately $2,500 per day,
with an additional trophy fee of $10,000. Depending on the country in
which a hunter visits, there may be several different fees required,
including game fees, observer fees, conservation fees, permit fees,
trophy handling fees, and government payments in terms of taxes, as
well as safari operator fees (Barnett & Patterson 2005, p. 71). In the
late 1990's, Tanzania reported annual revenue of $29.9 million from all
trophy hunting, South Africa reported $28.4 million, Zimbabwe reported
$23.9 million from all trophy hunting, Botswana reported $12.6 million,
and Namibia reported $11.5 million; the revenue generated solely from
lion hunting was not broken out (Barnett & Patterson 2005, p. iv). In
the past, government and private land owners were the primary
beneficiaries of the revenue gained; however, a portion of the revenue
derived from hunting, in some countries, is now being distributed to
local communities as well, which benefits the livelihoods of local
people as well as contributes to
[[Page 64492]]
national economies of African range states (Barnett & Patterson 2005,
p. vi).
Trophy Hunting as a Wildlife Management Tool
The concept of using trophy hunting to support lion conservation is
complex and counterintuitive to many. Many range countries rely heavily
on tourism (predominantly ecotourism and safari hunting) to provide
funding for wildlife management (IUCN 2006a, p. 24). The countries that
rely most on lion hunting are proportionally the highest in Mozambique,
Tanzania, and Zambia (Lindsey et al. 2012a, pp. 7-8). The revenue
generated from these industries provides jobs for locals, such as game
guards, cooks, drivers, and security personnel, and often brings in
revenue for local microbusinesses that sell art, jewelry, and other
native crafts. Revenue generated from scientifically based management
program is used to build and maintain fences, provide security
personnel with weapons and vehicles, provide resources for anti-
poaching activities, and provides resources for habitat acquisition and
management (Chardonnet et al. 2010, pp. 33-34; Newmark 2008, p. 321).
Revenue from trophy hunting increases the ability of many African
countries to manage wildlife populations both within and adjacent to
reserves; many of these hunting areas are geographically linked to
national parks and reserves, providing wildlife corridors and buffer
zones (Chardonnet et al. 2010, p. 34; Newmark 2008, p. 321).
Proponents and most species experts support trophy hunting as a
conservation tool for the African lion (Hunter 2011, entire; van der
Merwe 2013, entire; Hunter et al. 2013, entire) because it provides:
(1) Incentives for the conservation of large tracts of prime habitat,
and (2) funding for park and reserve management, anti-poaching, and
security activities. As habitat loss has been identified as one of the
primary threats to lion populations, it is notable that the total
amount of land set aside for hunting throughout Africa, although not
ameliorating the concerns about habitat loss, exceeds the total area of
the national parks, accounting for approximately half of the amount of
viable habitat currently available to lions (Chardonnet et al. 2010, p.
34; Packer et al. 2006, pp. 9-10). In Tanzania, 25-33 percent of the
total area, encompassing 190 hunting units and over 247,000 km\2\, has
been set aside for sport hunting purposes; this has resulted in an area
5.1 times greater than Tanzania's fully protected and gazetted parks
(Jackson 2013, p. 6; Barnett & Patterson 2005, p. 61).
In Botswana, despite the current ban on lion hunting, the country
currently has over 128,000 km\2\ of gazetted wildlife management areas
and controlled hunting areas set aside for hunting purposes, which
equates to 22.1 percent of the country's total area. This is in
addition to 111,000 km\2\ (or 19.1 percent) that has been set aside as
habitat in the form of National Parks, Game Reserves, and Forest
Reserves (Barnett & Patterson 2005, p. 7). Tanzania has land set aside
for sport hunting in the form of safari areas, communal land, and
privately owned properties that make up 23.9 percent of the total land
base (Barnett & Patterson 2005, pp. 76-77). In 2000, five countries in
southern Africa (Botswana, Namibia, South Africa, Tanzania, and
Zimbabwe) had set aside a combined 420,000 km\2\ of communal land,
188,000 km\2\ of commercial land, and 420,089 km\2\ of state land
totaling over 1,028,000 km\2\ for sport hunting purposes (Barnett &
Patterson 2005, p. iii). As a species with a considerable range (up to
1,000 km\2\) (Packer et al. 2013 p. 636; Haas et al. 2005, p. 4),
suitable habitat is important to the survival of the species, and the
marked decline in suitable habitat is a significant threat to the
species (see Habitat Loss). The land currently designated for use in
sport hunting has helped to reduce, but not eliminate, the impact of
habitat loss for the African lion.
Cost estimates for maintaining lion populations range, from an
annual budget of $500 per km\2\ in smaller fenced reserves to $2,000
per km\2\ annually for unfenced populations (Packer et al. 2013, p.
640; Lindsey et al. 2012a, p. 9). This includes but is not limited to
costs associated with permanent and temporary staff, fencing
installation and maintenance (fences can cost $3,000 per km to
install), infrastructure maintenance, anti-poaching activities such as
surveillance and snare/trap removal, wildlife restocking fees (both for
lions killed by illegal poaching/snares as well as other trophy species
killed by lions on the reserves), community outreach, and compensation
for loss of livestock in surrounding communities (Packer et al. 2013,
p. 640; Groom 2013, pp. 4-5; Lindsey et al. 2012a, p. 9; Barnett &
Patterson 2005, p. 82). For example, in the past, the Sav[eacute]
Valley Conservancy in Zimbabwe invested $546,000 annually on anti-
poaching activities and employed 186 permanent scouts, while operators
in Coutada 16, Mozambique, spent $60,000 annually on anti-poaching
(such as the removal of 5,000 gin traps) (Groom 2013, p. 5; Lindsey et
al. 2012a, p. 9). According to Barnett and Patterson (2005, p. 82), in
Zimbabwe:
Land invasions, resettlement and political instability has had
dire consequences for wildlife occurring in the commercial sector.
Land invasions have affected all wildlife management activities, and
resulted in severe habitat destruction, increased poaching and
infrastructure damage with thousands of kilometers of fences being
destroyed to make wire snares . . . A typical questionnaire response
from an invaded 50,000 acre farm in Masvingo Province . . .
indicates substantial poaching losses of up to $1,819,040, with over
3,400 snares recovered and 134 poachers arrested in just two months.
Niassa National Reserve, Mozambique, incurs annual costs of
approximately $1.9-2 million to maintain a 42,000-km\2\ area (Lindsey
et al. 2012a, p. 9). As a single source of revenue, the trophy hunting
of lions provides a substantial source of funds to pay for the
management of lion habitat. According to Lindsey et al. (2012a, p. 5),
with the exception of rhinoceros and exceptional elephant trophies,
``lions generate the highest revenue per hunt of any species in
Africa.'' In Niassa National Reserve, lion trophy hunting has generated
$380,000-400,000 annually (Lindsey et al. 2012a, p. 9). In the
Sav[eacute] Valley Conservancy, between 2005 and 2011, lion hunting in
Zimbabwe provided an estimated net income (based on 26 lions) of
approximately $1,365,000 in per-night charges and roughly $260,000 in
trophy fees (Groom 2013, p. 4).
Trophy hunting of lions, if part of a scientifically based
management program, can provide direct benefits to the species and its
habitat, both at the national and local level (See: Role of Local
Communities in Lion Conservation). Trophy hunting and the revenue
generated from trophy hunting are tools that range countries can use to
facilitate maintaining habitat to sustain large ungulates and other
lion prey, protecting habitat for lions, supporting the management of
lion habitat, and protecting both lions and their prey base through
anti-poaching efforts. While hunting alone will not address all of the
issues that are contributing to the declined status of the species, it
can provide benefits to the species.
Role of Local Communities in Lion Conservation
Over the last few decades, conservationists and range countries
have realized the integral role local communities play in the
conservation of lions and their habitat; when communities benefit from
a species,
[[Page 64493]]
they have incentive to protect it. Therefore, utilizing the wildlife
sector as a land-use option and source of income for rural populations
has increasingly been employed throughout the range countries of the
African lion. Many of these countries are classified as `developing'
nations; specifically, seven of the ten countries (we include Cameroon
here) where trophy hunting is permitted have 27-64 percent of their
populations living in severe poverty (United Nations Development
Programme's (UNDP) Human Development Report, https://hdr.undp.org/en/data, accessed July 7, 2014; Barnett & Patterson 2005, p. iii). These
countries often have high population growth, high unemployment, limited
industry, and a Gross Domestic Product (GDP) per capita lower than the
poverty level (Barnett & Patterson 2005, p. iii). These combined
challenges highlight the need for innovative solutions.
Conservationists and range countries recognize the value of the
wildlife sector; if managed sustainably, there is high potential to
contribute to rural economic development while simultaneously
protecting the unique ecological habitats and species contained therein
(Chardonnet et al. 2010, p. 33; Kiss [editor] 1990, pp. 1, 5-15).
Studies have indicated that, in order for species such as the
African lion to persist, the local communities must benefit from or
receive a percentage of funds generated from tourism such as wildlife
viewing, photography, or trophy hunting (White 2013, p. 21; Martin
2012, p. 57; Kiss [editor] 1990, pp. 1, 5-15). The economic value of a
species, such as lion, can encourage range countries to develop
management and conservation programs that involve local communities
which would ultimately discourage indiscriminate killings by local
communities (Groom 2013, pp. 3, 5; Hazzah et al. 2013, p. 1; White
2013, p. 21; Martin 2012, p. 49). If local communities see no
beneficial value of lions being present in their communal areas,
sustainable utilization of lions as a land-use becomes less competitive
with other land-use options, such as grazing and livestock management,
and local communities become unwilling and unable to manage their
wildlife heritage (Barnett & Patterson 2005, p. iii). When the value of
lions in areas outside of national parks is diminished, those areas are
likely to be converted to forms of land use less suitable for lions,
such as agriculture, livestock pastures, or areas of resource
extraction, making them even more vulnerable to expanding human
settlement (Van der Merwe 2013, p. 2).
Community conservancies that benefit from trophy hunting have
specifically been formed as a way to protect wildlife and habitat. As
an example, in Namibia, 160,000 km\2\ (61,776 mi\2\) of community
conservancies were established in part due to revenue from trophy
hunting. These conservancies benefit the local communities, which in
turn protect lion habitat. For example, in 2012, the Sav[eacute] Valley
Conservancy (Zimbabwe) ``provided over US$100,000 worth of support to
adjacent villages or farmers in the resettled areas. Assistance
included drilling boreholes, maintaining boreholes, dredging of dams,
building clinics and schools, assisting with repairs, maintenance and
materials for schools, education initiatives, school field trips,
provision of computer equipment in schools, and craft programs'' (Groom
2013, p. 5) Connecting conservation to community benefits can provide a
value for wildlife, including lions, where there was previously
resentment or indifference, helping to instill a sense of importance
for lion conservation Additionally, an estimated 125,000 kg of game
meat is provided annually to rural communities by trophy hunters at an
estimated value of $250,000 per year, which is considerable for rural
locations where severe poverty and malnutrition exists (White 2013, p.
21), further providing a value for wildlife, including lions. Lastly,
local communities benefit from the trophy hunting industry by gaining
employment as cooks, drivers, game guards, security, and anti-poaching
personnel, and they also obtain revenue for items purchased by trophy
hunters such as jewelry, art, and native handicrafts.
Trophy hunting as part of a scientifically based management program
may provide direct economic benefits to the local communities and can
create incentives for local communities to conserve lions, reduce the
pressure on lion habitat, and end retaliatory killing, primarily
because lions are viewed as having value. Conversely, lack of
incentives could cause declines in lion populations because lions are
viewed as lacking value and are perceived to kill livestock, which do
have value to communities (see Human-lion Conflict).
Many range countries have realized local communities must benefit
from the conservation of the species because [why?] and have revised
their land management and ownership policies to reflect this. Of the
ten countries where lion trophy hunting currently occurs (including
Cameroon), seven have developed National Poverty Reduction Strategies
in partnership with the International Monetary Fund (for a complete
list, see https://www.imf.org/external/np/prsp/prsp.aspx); each of these
has incorporated sustainable natural resource development as a main
priority, and emphasized benefit distribution and management to rural
communities (Benin 2000, unpaginated; Burkina Faso 2000, unpaginated;
unpaginated; CAR 2000, p. 45; Mozambique 2000, unpaginated; Tanzania
2000, pp. 13, 21; Zambia 2000, unpaginated). As a result, an increase
in participation by local communities in managing natural resources
that are adjacent to reserves is occurring in several areas.
Captive Lions
In analyzing threats to a species, the Service focuses its analysis
on threats acting upon wild specimens within the native range of the
species, because the goal of the Act is survival and recovery of the
species within its native ecosystem. We do not separately analyze
``threats'' to captive-held specimens because the statutory five
factors under section 4 (16 U.S.C. 1533) are not well-suited to
consideration of specimens in captivity and captive-held specimens are
not eligible for separate consideration for listing. However, we do
consider the extent to which specimens held in captivity create,
contribute to, reduce, or remove threats to the species.
Captive-held African lions, including those that are managed for
trophy hunting in South Africa and lions held in captivity in zoos, are
believed to number between a few thousand and 5,000 worldwide (Republic
of South Africa 2013, p. 5; Barnett et al. 2006a, p. 513). Captive
lions in general are not suitable for reintroduction due to their
uncertain origins (Barnett et al. 2006a, p. 513; Hunter et al. 2012, p.
3), potential maladaptive behaviors, and higher failure risk compared
to translocated individuals (Hunter et al. 2012, pp. 2-3). There may be
cases where captive specimens provide a benefit to the species under
certain circumstances. For example, the display of Giant pandas in U.S.
zoos has generated considerable revenue that is used for in-situ
conservation of the species in China. It may be possible that captive
lions could also serve a purpose of generating revenue for in-situ
conservation.
Summary of Trophy Hunting
Although there is some indication that trophy hunting could
contribute to
[[Page 64494]]
local declines in lion populations through unsustainable quotas,
corruption, and possible disruption of pride structure through
infanticide and take of males that are too young, we do not find that
any of these activities rises to the level of a threat to the African
lion subspecies at this time. It appears that most range countries that
allow trophy hunting of African lions restrict offtake to approximately
2-4 percent of their lion populations for trophy hunting annually,
excluding South Africa, where offtake is from predominantly captive-
born animals, and Zimbabwe, where offtake is 2-3 percent higher than in
other countries (Packer et al. (2006, pp. 2-3). Exports of lion
trophies have demonstrated a decreasing trend when exports of likely
captive-born lions from South Africa are excluded (CITES lion gross
exports, https://trade.cites.org, accessed April 23, 2014), and lions
from South Africa are likely captive-born (RSA 2013, p. 5). Most of the
range countries that allow trophy hunting have quotas in place to limit
take. Tanzania, with a population of approximately 16,000 lions, has a
quota of 50 animals per year. Many other range countries have laws in
effect that address trophy hunting, and several have moratoriums in
place. The hunting community is taking the lead in developing best
management practices to address take of males that are under 6 years of
age, and they are guiding the development of scientifically based tools
for minimizing the impact of trophy hunting on the social structure of
lion populations. This 6-year age restriction on lion trophies is in
the process of being self-implemented by professional hunting guides,
and is being adopted by certain range states, such as Tanzania (White
2013, p. 14; Whitman et al. 2004, p. 176).
Currently, most countries that allow trophy hunting of lions appear
to be reviewing their trophy hunting practices (Jackson 2013, pp. 2-3;
White 2013, pp. 12-13). Range countries have recognized the need to
incorporate best management practices, and have been progressively
updating the policies and management systems in order to implement them
(Lindsey et al. 2013a, pp. 4-10).
Finally, we found that, if trophy hunting of lions is part of a
scientifically based management program, it could provide considerable
benefits to the species, by reducing or removing incentives by locals
to kill lions in retaliation for livestock losses, and by reducing the
conversion of lion habitat to agriculture. Trophy hunting, if managed
well and with local communities in mind, can bring in needed revenue,
jobs, and a much-needed protein source to local people, demonstrating
the value of lions to local communities (Groom 2013, pp. 1-3; Lindsey
et al. 2006, pp. 283, 289). In addition, the amount of habitat that has
been set aside by range countries specifically for trophy hunting has
greatly increased the range and habitat of lions and their prey base,
which is imperative given the current ongoing rate of habitat
destruction occurring in Africa. The total amount of land set aside for
trophy hunting throughout Africa exceeds the total area of the national
parks, providing half the amount of viable lion habitat (Chardonnet et
al. 2010, p. 34; Packer et al. 2006, pp. 9-10). However, expanding
protected areas without taking the human population into consideration
could lead to more resentment and retaliatory killing of lions (Nelson
et al. 2009, p. 315).
Therefore, we conclude, based on the best scientific and commercial
information available, that trophy hunting is not a significant threat
to the species.
Traditional Use of Lion Parts and Products
CITES (2014, p. 8) reports that many African countries, including
Somalia, Nigeria, Burkina Faso, Kenya, and Cameroon, maintain local
markets in lion products, which include teeth, claws, fat, whiskers,
bone, bile, testicles, meat, and tails for use as talismans,
decorations, and in traditional African medicine. In Ghana, lion parts
and products are used for ceremonial, medicinal, and nutritional
purposes (Burton et al. 2010, p. 4). Skins and claws of lions were
observed for sale in a market in Tamale, Ghana. Lions in and around
Mole National Park in Ghana have been killed for traditional
consumptive purposes (Burton et al. 2010, p. 4). In some cases, lions
(either alive or dead) have been ``laundered'' through other countries
so that their country of origin is unknown. As an example, lions have
been found to be shot in Zimbabwe and Mozambique and declared as South
African trophies (Lion Aid 2011, p. 20). In other cases, there have
been reports of captive-born lions being smuggled between Botswana and
South Africa and described as wild (Mouton 2013, pp. 1-2). Lion
products, such as the trade in lion bone, seem to be primarily
byproducts of trophy hunting; hunters are primarily interested in the
trophy and skin and, therefore, the bones and other parts are sold
separately (CITES 2014a, p. 10). However, since the reports of these
types of activities are primarily anecdotal in nature, based on the
best available scientific and commercial information, we find that the
sale of these byproducts does not currently pose a threat to the
species. Further, without a significant shift in the market, it is not
likely to become a threat in the foreseeable future.
Conservation Measures in Place To Protect Lions
There has been awareness for several years that conservation
strategies need to be implemented for the African lion due to the
apparent decrease in its population numbers (Hamunyela et al. 2013, p.
1; Henschel et al. 2010, p. 34; Gebresenbet et al. 2009, p. 5; IUCN
2006a, b, entire). Prior to 2006, institutional inconsistencies
throughout the African lion's range resulted in poor lion conservation
policies and little to no enforcement of existing laws (IUCN 2006b, p.
18). As mentioned, in 2005 and 2006, nongovernmental organizations
(NGOs) and several governments at various levels organized two regional
lion conservation workshops. Species specialists, wildlife managers,
and government officials attended these regional workshops in order to
provide range country governments with frameworks for developing their
own national action plans for the conservation of lions. Over 50 lion
specialists, representing all lion range countries, participated in
these workshops (Henschel et al. 2010, p. 34). During the workshops
lion experts collectively assessed what they believed to be the then-
current status of African lions based on a variety of information, and
subsequently identified 86 African LCUs. This information was then used
as a framework to identify lion areas, strongholds, and potential
strongholds by Riggio et al. (2013, p. 32).
Many countries with very small lion populations have developed or
updated their conservation plans for the African lion. Some of these
include Benin, Cameroon, Uganda, and Malawi. Some range countries
participate in transboundary conservation projects and are
collaborating on transboundary lion conservation initiatives for shared
lion populations. Most range countries have a national lion action plan
or strategies in place, particularly if there are economic incentives
for them to have viable lion populations (Groom 2013; Nghidinwa et al.
2013, pp. 11-12; Zambia Wildlife Authority 2012; Lion Aid 2011, pp. 1-
2; Mesochina et al. 2010; Government of Tanzania 2010; Begg and Begg
2010). Range states have also implemented a number of conservation
strategies designed to conserve habitat, reduce human-lion
[[Page 64495]]
conflict, and preserve the lion's prey-base.
Conservation Measures To Stem Habitat Loss
Habitat loss represents one of the main threats facing the African
lion (Bauer et al. 2008, unpaginated). Attempts by range countries to
address this decline in habitat are manifested in a number of ways,
such as the creation of protected areas and the establishment of
wildlife corridors to connect fragmented habitats.
Two conservation tools utilized by range countries for African
lions include the establishment of protected areas and the enforcement
of protections in these areas (Mesochina et al. 2010a and b; Treves et
al. 2009, pp. 60, 64). Over the past few decades, the effectiveness of
protected areas in protecting habitat has been studied, particularly in
Africa (Pfeifer et al. 2012, p. 1; Craigie et al. 2010, pp. 2,221-
2,222). A study conducted by the Wildlife Conservation Society in 2005
found that most lion populations in protected areas of southern and
eastern Africa have been essentially stable over the previous three
decades (Ray et al. 2005, p. 67). However, several problems have
emerged. For example, certain land-tenure systems do not recognize
community ownership of land and wildlife and undermine the extent to
which benefits are converted into incentives for conservation.
Protected-area ``boundaries'' are not always visible. Additionally, law
enforcement in protected areas can be sporadic, and parks are often
understaffed (Pfeifer et al. 2012, pp. 1, 7). Lastly, despite the
Wildlife Conservation Society's findings, more recent evidence suggests
that some protected areas are being more commonly encroached upon as
human populations expand and search for resources.
Despite encroachment, protected areas are somewhat effective at
protecting wildlife and habitat as rates of habitat loss tend to be
lower in protected areas than outside them (Green et al. 2013, p. 70;
Pfeifer et al. 2012, p. 2). African countries are realizing the
benefits of managing their wildlife populations and parks for tourism;
however, conservation of vast areas of land for megafauna such as the
African lion is not only complex, but also expensive. As an example,
the 28-km (17-mi) elephant corridor, completed in 2011 in Kenya, cost
$1 million (The Nature Conservancy 2013, unpaginated). Additionally,
the overall costs of anti-poaching and compensation is expected to
increase in range states concurrently with growing human populations,
declining purchasing power of external funds, and corruption (Garnett
et al. 2011, pp. 1-2; Wittemyer et al. 2008, pp. 123, 125).
Another mechanism for protecting habitat is to reconnect fragmented
habitat across national boundaries. Corridors are being restored,
fences are being removed, and protected areas are being connected.
Restoration of these corridors allows wildlife to travel between areas
of suitable habitat (Jones et al. 2012, pp. 469-470). In some areas,
fences have been constructed to protect grazing resources for domestic
livestock as well as to provide barriers to disease (Gadd 2012, pp.
153, 176). One aspect of these fences is that they separate lions from
their prey. In southern Africa, the trend now is to take down fences to
increase the size of connected habitat and link it to reserves and
national parks (IUCN 2009, p. 101; IUCN 2008, various). The Limpopo
Transfrontier Park is another example of where this is being
implemented (Newmark 2008, p. 327). Boundary fences along national
borders that separate many reserves are being removed to form a 35,000-
km\2\ park. Limpopo National Park (formerly known as Coutada 16) in
Mozambique; Kruger National Park in South Africa; Gonarezhou National
Park, Manjinji Pan Sanctuary, and Malipati Safari Area in Zimbabwe will
all be connected, as will be the area between Kruger and Gonarezhou,
and the Sengwe communal land in Zimbabwe and the Makuleke region in
South Africa (Newmark 2008, p. 327). However, in some locations, areas
that have previously been designated as corridors have been encroached
upon by human settlements and agriculture (Estes et al. 2012, pp. 258-
261; Jones et al. 2012, p. 469).
Tanzania is an example of a country attempting to reconnect
habitat. As of 2002, the Tanzanian Government, with donor and NGO
support, was reconnecting the nine largest blocks of forest in the East
Usambara Mountains using wildlife corridors (Newmark 2002, various).
Additionally, the 2009 Wildlife Act of Tanzania allows the Minister, in
consultation with relevant local authorities, to designate wildlife
corridors, dispersal areas, buffer zones, and migratory routes. The
2010-2015 National Elephant Management Plan of Tanzania indicates that
corridors are the primary objective of the plan, and although primarily
designed for elephants, these corridors allow for continuity of
populations of other large mammal species such as lions (Jones et al.
2012, p. 470).
In 2011, Kenya (which neighbors Tanzania to the North), completed a
28-km corridor through an area that had been heavily impacted by human-
wildlife conflict. The purpose of the corridor was primarily to reduce
human-elephant conflict and appears to have been successful (Mount
Kenya Trust 2011, p. 1). The corridor also allows other wildlife such
as lions to disperse through habitat that otherwise would have been
unfavorable for wildlife to travel through (Mount Kenya Trust 2011, p.
1). It was an expensive project, but recent reports indicate that the
effort has served its purpose: Elephants are using the corridor on a
regular basis (particularly an underpass under a highway), and humans
are reporting less human-wildlife conflict (Mount Kenya Trust 2011, p.
1).
However, connectivity alone does not ensure the dispersal of
animals (Roever et al. 2013, pp. 19-21). The Tanzania Wildlife Research
Institute (TAWIRI) is a parastatal organization under Tanzania's
Ministry of Natural Resources and Tourism, and is responsible for
conducting and coordinating wildlife research activities in Tanzania
(https://tawiri.or.tz/). In this role, TAWIRI has been actively involved
in promoting the development of and monitoring the use of wildlife
corridors in Tanzania (https://www.tzwildlifecorridors.org). Surveys
conducted in 2009 and 2010 suggest that the Nyanganje Corridor in
Tanzania is no longer being used by elephants and other wildlife. This
corridor is at a narrow passage in the Kilombero Valley and is the
shortest distance for animals to cross between the Udzungwa and Selous
ecosystems. Despite efforts in place, much of the corridor is being
encroached upon by conversion of land to rice farming and cattle
grazing (Jones et al. 2012, p. 469). Because these activities often
deter wildlife from passing through, the corridor is ineffective (Jones
et al. 2012, p. 469). TAWIRI reminds wildlife managers that they need
to continue to implement steps to ensure that corridors are functioning
properly.
Conservation Measures in Place To Stem the Loss of Prey Base
Lions, like most large carnivores, prey upon a variety of species
including buffalo, plains zebra, wildebeest, giraffe, gemsbok, kob, and
warthog (Kenya Wildlife Service 2013, p. 13; Niassa National Reserve
Technical Report 2011, p. 4; Nowell and Jackson 1996, p. 18). Depletion
of these prey species due to competition with humans represents a
threat to the lion (Chardonnet et al. 2005, pp. 8-9). As noted, the
increase in the human population in Africa is a major contributor to
the increase in the
[[Page 64496]]
demand for bushmeat, which in turn increases human encroachment into
wildlife lands (Lindsey et al. 2012b, p. 36). In addition to the
increase in the human population, lack of an alternative livelihood,
lack of alternate food sources, and lack of clear rights over land or
wildlife are contributing factors toward the increase in demand for
bushmeat (Lindsey et al. 2012b, pp. 36-41). The advent of automatic
weapons in the bushmeat trade impacts the lion's prey base, which is
being hunted at unsustainable levels.
Reconnecting fragmented habitat has the additive effects of not
only conserving the biodiversity of the African lion's habitat, but
also that of its prey base (Lindsey et al. 2012b, p. 43). These types
of restoration practices enhance the health of species by allowing
genetic interchange to occur and, thus, conserve the genetic diversity
of all wildlife. Wildlife management entities are linking many of the
major protected areas by removing boundary fences along national
borders that separate many reserves in addition to creating or
improving corridors to link good-quality habitat for wildlife (Gadd
2012, p. 179; Newmark 2008, pp. 323-324). To address the increasing
consumption of bushmeat, host countries have employed a variety of
different strategies, including the development of alternative
industries for communities. Helping local communities develop alternate
industries represents one of the ways range countries can reduce their
dependence on bushmeat. Throughout Africa, several ideas have been
attempted with varying levels of success. For example, the Anne Kent
Taylor Fund (AKTF) helps local Maasai women to buy beads and other
supplies to produce traditional items for the local tourist industry
(AKTF 2012, p. 7; Lindsey et al. 2012b, p. 45; van Villet 2011, p. 17).
In addition, AKTF helps organize local men into anti-poaching and de-
snaring teams (AKTF 2012, p. 5; van Villet 2011, p. 17). By creating
programs targeting both men and women, AKTF creates an environment that
provides communities with financial stability as well as direct
community interest in protecting local wildlife. With 13 years
assisting local communities, the AKTF represents one of the more
successful attempts to encourage locals to shift away from relying on
bushmeat.
Studies compiled by Huzzah 2013 (pp. 1, 8) have shown that local
communities who lived near protected areas with more lenient policies
have a more positive attitude and relationship with both the manager
and the protected area as a whole. This open approach to protected area
management reflects a trend in recent years to bring in local
communities to assist in the management of protected areas (Lindsey et
al. 2012b, p. 53). Wildlife management programs run by local
communities are defined by two goals: Conserving wildlife and providing
economic aids to the community (Bandyopadhyay et al. 2010, p. 5). With
regards to discouraging the consumption of bushmeat, this new approach
is seen in the creation of community-based wildlife management programs
(van Villet 2011, p. 26). The purpose of these programs is to give the
local community a direct stake in the management of wildlife areas. One
use for these areas is to turn them into game ranches. These areas are
used both for legal bushmeat production as well as trophy hunting and
ecotourism.
One such program is the Chivaraidze Game Ranch in Zimbabwe (van
Villet 2011, pp. 28-29). The Chivaraidze Game Ranch started in 1996
with the stated goal of reducing poaching through providing bushmeat at
a reduced price. However, internal infighting in the organization over
the devolution of power to local communities, between those in favor of
devolution and a powerful local interest group, limited the
effectiveness of the organization. In the span of 8 years (between 2001
and 2009), the Chivaraidze Game Ranch has had six different boards of
directors (Mombeshora and Le Bel 2010, p. 5). Furthermore, a power
shake-up in local communities along party lines and kinship affiliation
limited the abilities for communities to cooperate with each other (van
Villet 2011, pp. 28-29; Mombeshora and Le Bel 2010, p. 7). The result
was that the cost of maintaining the program exceeded the benefits to
the local community. The decline in economic benefits to the local
community coincided with a resurgence in poaching within areas of the
park (Mombeshora and Le Bel 2010, p. 3). The result of the Chivaraidze
Game Ranch project reflects the difficulty in shifting wildlife
management from a centralized national government approach towards a
more decentralized, community-based approach.
Unlike the difficulties encountered in Zimbabwe, Namibia has had
greater success in setting up community-run conservancies. After
gaining independence in 1990, Namibia began to turn over ownership of
wildlife areas to local communities (van Vliet 2011, p. 29;
Bandyopadhyay et al. 2010, p. 6). By 2011, Namibia had 64 communities
that covered 17 percent of the country total area (van Vliet 2011, p.
29; Connif 2011, npn; NASCO 2010, p. 4). The majority of the incomes
from these conservancies come from ecotourism, followed by trophy
hunting (NASCO 2010, p. 22). These incomes are then used to support
infrastructure improvement in the community. In addition, legal
bushmeat acquired within conservancy lands is distributed to local
families (NASCO 2010, p. 25). The success of the program in Namibia has
been attributed to Namibia's unique characteristics, including low
population density and favorable seasonal rain, which helps prey
species recover (van Vliet 2011, p. 30). Despite the successes in
Namibia, the country's unique characteristics mean that adapting
Namibia's success to other, more densely populated countries will be
difficult.
Conservation Measures To Stem Human-Lion Conflict
As the human population expands, the potential for conflict with
wildlife increases. In Africa, conflict between villagers and lions,
who prey upon livestock, represent a threat to the species (Chardonnet
et al. 2010, p. 12; Moghari 2009, p. 14; IUCN 2006a, p. 23). In
addition, habitat loss due to conversion of land increases the chance
of villagers coming into direct contact with lions (Chardonnet et al.
2010, p. 24). In an attempt to address these problems, range countries
have employed a variety of different strategies to help the lion. Such
strategies involve education, an effective conservation plan, and
interacting with the local community.
Historically, range countries seek to mitigate human-lion conflict
through controlling rather than conserving the predator population. In
countries such as Malawi, for example, the Department of Game, Fish and
Tsetse Control would shoot large carnivores that prey upon livestock.
The result of this policy was that, between 1948 and 1961, over 560
predators (which include lions and leopards) were killed in the country
(Mesochina et al. 2010b, p. 35). While this department was disbanded in
1963 and jurisdiction shifted to the new Department of Forestry, crop
and livestock protection still remains an important part of its
function. Despite the department focusing on protecting crops and
livestock, the number of lions killed in the country has declined.
Between 1977 and 1982, eight lions were killed, whereas six lions were
killed between 1998 and 2007 (Mesochina et al. 2010b, p. 35). While
fewer lions are being killed than in the previous decades, problems
remain, including lack of resources, lack of
[[Page 64497]]
manpower, and corruption within the range countries.
Current governmental management of lions in countries such as
Malawi, Tanzania, and Zambia are managed by the Problem Animal Control
units (Mesochina et al. 2010a, p. 41; Mesochina et al. 2010b, p. 36).
When lion attack incidents occur, Problem Animal Control dispatches
officials to investigate the problems. If the problem lion is located,
it is either removed or eliminated. When properly funded, this program
has helped in reducing not only conflicts between lions and humans but
also has driven down the numbers of lions killed. Between 2005 and
2009, there were 116 reported cases of lions killed, with the number of
lions killed being less than 50 per year in Tanzania (Mesochina et al.
2010a, p. 41). However, limitations of resources (including both
manpower and funds) have hampered the effectiveness of these officials
in responding to these incidents. In addition, many Problem Animal
Control interventions resulted in the death of the lion (Mesochina et
al. 2010a, p. 41; Chardonnet et al. 2009, p. 36). Even in cases of
translocation, the lions that were being transported often end up
injured or continue to pose problems to the community (Bauer et al.
2007, p. 91).
NGOs are also assisting in protecting lions. Intervention by NGOs
often takes the form of interacting with the local community
(Winterbach et al. 2010, p. 98). Lion Guardians, which operate in
Kenya, recruits and educates local young men. These men then monitor
and track lion movement and warn herders of lion presence in the area,
thereby mitigating or preventing possible lion-human conflict (Hazzah
et al. 2014, p. 853; Lion Guardians 2013, p. 7; Lion Guardians 2012, p.
3). In addition, Lion Guardians work with tribal elders to dissuade
young men from killing lions for ceremonial purposes. Historically, the
killing of lions through ritualized lion hunts called ilmurran is
rewarded with gifting of cows and other rewards (Lion Guardians 2012,
p. 5; Goldman et al. 2010, p. 334). After introducing village elders to
the Lion Guardians program first hand, many return home to their
village and give their blessings to the project. This education led to
significant results; on August 11, 2013, two Lion Guardians stopped a
group of hunters who were planning to hunt a lion in retaliation for
the lion preying on their livestock. The local village elders fined the
potential hunters two cattle each for going on a lion hunt, marking a
gradual but significant shift in the cultural attitudes regarding the
lion (Hazzah et al. 2014, p. 858; Lion Guardians 2013, p. 20). Since
its establishment in 2007, only five lions had been killed in
territories where Lion Guardians operates, in contrast to more than 100
lions killed in adjacent areas (Lion Guardians 2013, p. 5).
Furthermore, reduced lion mortality was sustained across multiple
years, resulting in the reserve having one of the highest lion
densities in Africa (Hazzah et al. 2014, p. 857; Schuette et l. 2013,
p. 149). Despite the success of this program, retaliatory as well as
ceremonial killings of lions outside the program areas remain a threat
to the species.
We found that many of the lion range states are trying to address
lion conservation through the establishment of protected areas,
wildlife management areas, wildlife corridors, and reconnecting
habitat. In some areas, creating incentives for lion conservation is
occurring through community conservation programs in range countries.
In other cases, participatory strategies have been implemented to
enhance local tolerance for large carnivores in Africa. An increasing
number of programs encourage local communities to solve problems that
arise from human-lion conflict without killing lions. However, the
effectiveness of these measures still ranges from successful to
unsuccessful, due in part to lack of resources, political will, and
infighting. It is imperative that range countries continue to recognize
and support the role that local communities play in lion conservation.
Greater support by countries to address the needs of local communities,
and thereby address the needs of lions, may be the single-most
important role these countries can play in changing the trajectory of
lion declines.
Regulatory Mechanisms
Regulatory mechanisms in place to provide protections to African
lions vary substantially throughout Africa. As mentioned in the
Conservation Status of African Lions CITES section, lions are listed in
Appendix II under CITES, and with the exception of South Sudan, all of
the lion range states are parties to CITES. According to the draft
CITES Periodic Review of the Status of African Lions (CITES 2014a, pp.
14-15) outside of CITES, lions have no legal protections in four
countries: Burundi, Guinea Bissau, Lesotho, and Swaziland. However,
CITES 2014a (p. 15), states that most of the southern and eastern lion
range states have regulatory mechanisms in place to protect lions. We
found that most of the range states have national environmental
legislation to establish national parks and conservation areas, and to
conserve and regulate the take, hunting, and trade of wildlife,
including parts and products, but could find no legislation specific to
lions, nor to the main threats affecting lions: habitat loss, human-
lion conflict, and loss of prey base (See: Appendix A, Ecolex
information was accessed July 7-10, 2014, at https://www.ecolex.org.\8\).
---------------------------------------------------------------------------
\8\ ECOLEX is a comprehensive database on environmental law,
maintained by the International Union for Conservation of Nature
(IUCN), the United Nations Environment Programme (UNEP), and the
Food and Agriculture Organization of the United Nations (FAO). Our
search terms used with respect to wildlife laws were ``African
lion'' and ``country'', e.g., ``Angola'', ``Benin'', etc. See
Appendix A.
---------------------------------------------------------------------------
Our status review did not reveal regulatory mechanisms in place
that specifically address the main threats affecting lions. We are
requesting comments or information from lion range states, other
concerned governmental agencies, the scientific community, or any other
interested parties concerning regulatory mechanisms that address the
three main threats to lions: habitat loss, human-lion conflict, and
loss of prey base.
Finding
Section 4 of the Act (16 U.S.C. 1533) and its implementing
regulations at 50 CFR 424 set forth the procedures for adding a species
to, and/or removing a species from, the Federal Lists of Endangered and
Threatened Wildlife and Plants. As noted in the Information Requested
section, a species may be determined to be an endangered or threatened
species due to one or more of the five factors set forth in section
4(a)(1) of the Act:
(A) The present or threatened destruction, modification, or
curtailment of its habitat or range;
(B) Overutilization for commercial, recreational, scientific, or
educational purposes;
(C) Disease or predation;
(D) The inadequacy of existing regulatory mechanisms;
(E) Other natural or manmade factors affecting its continued
existence.
In assessing whether the African lion meets the definition of an
endangered or threatened species, we considered the five factors in
section 4(a)(1) of the Act. A species is ``endangered'' for purposes of
the Act if it is in danger of extinction throughout all or a
significant portion of its range and is ``threatened'' if it is likely
to become endangered within the foreseeable future throughout all or a
significant portion of its range. The ``foreseeable future'' is the
period of time over which events or effects reasonably can or should be
anticipated, or trends extrapolated.
[[Page 64498]]
When considering what factors might constitute threats to a
species, we must look beyond the mere exposure of the species to a
factor to evaluate whether the species may respond to the factor in a
way that causes actual impacts to the species. If there is exposure to
a factor and the species responds negatively, the factor may be a
threat and we attempt to determine how significant a threat it is. The
threat is significant if it drives, or contributes to, the risk of
extinction of the species such that the species may warrant listing as
endangered or threatened as those terms are defined in the Act. We
conducted a review of the best scientific and commercial data available
regarding the status of the African lion and assessed whether the
African lion is endangered or threatened throughout all of its range.
There is consensus within the research community as well as lion
range states that the African lion is impacted by a number of factors
actively contributing to its population decline throughout Africa:
habitat loss (fragmentation and degradation) (Factor A); decreased
access to food prey sources (aka loss of prey base) (Factor B);
retaliatory killing, snaring, and poaching (both intentional and
unintentional), and deleterious effects in its viability due to small
populations in some areas within its range (Factor E) (Nyanganji et al.
2012, p. 12; Seguya et al. 2010, p. 26).
We find three main threats, habitat loss, loss of prey base, and
human-lion conflict, are impacting lions, alone and in combination,
such that the subspecies is likely to become endangered within the
foreseeable future throughout all of its range. In the past several
decades, the human population has been expanding with concomitant large
decreases in lion habitat and likely lion numbers, resulting in an
extremely large reduction in the species' range. As human populations
continue to rise in sub-Saharan Africa, the amount of land required to
meet the expanding human population's needs is constantly increasing.
Lions are increasingly limited to protected areas, and human population
growth rates around protected areas in Africa tend to be higher than
the average rural growth rate (Wittemyer et al. 2008, entire).
Considering the majority of the human population in sub-Saharan Africa
is rural, and land supports the livelihood of most of the population,
loss and degradation of lion habitat, loss of prey base, and increased
human-lion conflict can reasonably be expected to accompany the rapid
growth in sub-Saharan Africa's human population into the foreseeable
future.
Africa has the fastest population growth rate in the world (UNEP
2012a, p. 2). The majority of the population is rural, and about 60-70
percent of the population relies on agriculture and livestock for their
livelihood (UNEP 2006, pp. 82, 100, 106; IAASTD 2009, p. 2). As a
result, a large portion of the growing population will depend directly
on expansion of agriculture and livestock grazing to survive in the
future. Between 2010 and 2050, the population of sub-Saharan Africa is
projected to more than double to more than 2 billion (from 831 million
to 2.1 billion) (UN 2013, p. 9). During about this same time period
(2005 to 2050), the area of cultivated land is projected to increase by
51 million ha (approximately 21 percent) (Alexandratos and Bruinsma
2012, p. 107). However, this figure does not include rangeland, and the
majority of agricultural land in Africa is devoted to grazing (UNEP
2012b, p. 68), thus that figure may be much larger. The number of
livestock (cattle, sheep, and goats) in sub-Saharan Africa is projected
to increase about 73 percent, from 688 million to 1.2 billion, by 2050
(Alexandratos and Bruinsma 2012, p. 133). Therefore, in the case of
African lion, the best available scientific and commercial data that we
rely upon in projecting future conditions for the purpose of this
listing determination establish the foreseeable future to be 2050.
Human settlements and agricultural and pastoral activities have
expanded into lion habitat and protected areas, decreasing prey
availability and increasing exposure of livestock and humans to lions.
Human-lion conflict and associated retaliatory killing of lions will
continue to play a major role in the reduction of lion populations and
is the greatest current threat to remaining lion populations. The
lion's prey base has decreased in many parts of its range in large part
due to the bushmeat trade
Bushmeat is the primary source of protein for humans in much of the
lion's range (Chardonnet et al. 2010, p. 27; Mesochina et al. 2010a, p.
38; Abwe and Morgan 2008, p. 26; Bennett et al. 2007, p. 885; Fa et al.
2006, p. 507), comprising between 6 percent (southern Africa) and 55
percent (Central African Republic) of a human's diet (Chardonnet et al.
2005, p. 9; IUCN 2006b, p. 19). This reliance by humans on protein
obtained from bushmeat results in direct competition for prey species
between humans and lions, and commercial poaching of wildlife through
the use of automatic weapons is a significant threat to lion prey
(Chardonnet et al. 2010, p. 27). Because many wildlife species are
being hunted at unsustainable levels to meet this demand within the
range of the lion, its prey base is becoming depleted in many areas and
has led to lion attacks on livestock and humans (Hoppe-Dominik et al.
2011, p. 452; Chardonnet et al. 2010, pp. 6, 13-14; Frank et al. 2006,
p. 12). Given the rapid increase in humans and livestock by 2050, we
can reasonably expect the conditions described above to worsen. Also,
as livestock numbers increase and as expansion of agricultural and
pastoral practices continue to deplete and degrade the habitat that
lion's prey rely on, the lion's prey base is expected to further
decline. As the lion's prey base is hunted at unsustainable levels to
meet a growing demand for food, livestock depredation and retributive
killing of lions through spearing, shooting, trapping, and poisoning
will continue to occur, and will likely increase (Dickman 2013, p. 379;
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).
Lion range countries are aware of the threats affecting lions, and
many are working to address them. NGOs and several governments at
various levels have organized regional lion conservation workshops,
which have helped them to identify Lion Conservation Units. Most range
countries have a national lion action plan or strategy in place (Groom
2013; Nghidinwa et al. 2013, pp. 11-12; Zambia Wildlife Authority 2012;
Lion Aid 2011, pp. 1-2; Mesochina et al. 2010; Government of Tanzania
2010; Begg and Begg 2010). Some range countries participate in
transboundary conservation projects to create wildlife corridors and
reconnect habitat, and are collaborating on transboundary lion
conservation initiatives for shared lion populations. Reconnecting
fragmented habitat has the additive effects of not only strengthening
the biodiversity of the African lion but also that of its prey species
(Lindsey et al. 2012b, p. 43). Wildlife management entities are linking
many of the major protected areas by removing boundary fences along
national borders that separate many reserves, in addition to creating
or improving corridors to link good-quality habitat for wildlife (Gadd
2012, p. 179; Newmark 2008, pp. 323-324).
Range states have also implemented a number of conservation
strategies designed to conserve habitat, reduce human-lion conflict,
and preserve lion
[[Page 64499]]
prey-base. In order to address the increasing consumption of bushmeat,
host countries have employed a variety of different strategies,
including the development of alternative industries for communities,
which can reduce their dependence on bushmeat. For example, the Anne
Kent Taylor Fund (AKTF) helps local Maasai women to buy beads and other
supplies to produce traditional items for the local tourist industry
(AKTF 2012, p. 7; Lindsey et al. 2012b, p. 45; van Villet 2011, p. 17)
and has organized local men to participate in anti-poaching and de-
snaring teams (AKTF 2012, p. 5; van Villet 2011, p. 17). By targeting
both men and women in the community, such programs provide communities
with financial stability as well as direct community interest in
protecting local wildlife. African countries are realizing the benefits
of managing their wildlife populations and parks for tourism; however,
conservation of vast areas of land for megafauna such as the African
lion is expensive. The costs of anti-poaching and compensation is
expected to increase in range states concurrently with growing human
populations, declining purchasing power of external funds, and
corruption (Garnett et al. 2011, pp. 1-2; Wittemyer et al. 2008, pp.
123, 125).
Studies have shown that local communities who live near protected
areas (PAs) with community-based conservation policies have more
positive attitudes and relationships with both the park manager and the
PA as a whole (Huzzah 2013, pp. 1, 8). This open approach to PA
management reflects a trend in recent years to bring in local
communities to assist in the management of PAs (Lindsey et al. 2012b,
p. 53). Wildlife management programs run by local communities are
defined by two goals: conserving wildlife and providing economic aids
to the community (Bandyopadhyay et al. 2010, p. 5). NGOs are also
assisting in protecting lions. Intervention by NGOs often takes the
form of interacting with the local community (Winterbach et al. 2010,
p. 98). For example, Lion Guardians, which operates in Kenya, has shown
great success with its Lion Guard program. Lion Guardians educates
local young men who monitor and track lion movement and warn herders of
lion presence in the area, thereby mitigating or preventing possible
lion-yhuman conflict (Hazzah et al. 2014, p. 853; Lion Guardians 2013,
p. 7; Lion Guardians 2012, p. 3). Outreach to tribal elders has
successfully helped elders to dissuade young men from killing lions for
ceremonial purposes. The result of such programs has been a gradual
change in cultural attitudes towards lions (Hazzah et al. 2014, p. 858;
Lion Guardians 2013, p. 20).
Finally, many range countries rely heavily on tourism
(predominantly ecotourism and safari hunting) to provide funding for
wildlife management (IUCN 2006a, p. 24). The revenue generated from
these industries can be critical to fund wildlife management programs
in range states. Tourism, through ecotourism and trophy hunting, can
provide jobs to locals (such as game guards, cooks, drivers, security
personnel) and often brings in revenue for local microbusinesses that
sell art, jewelry, and other native crafts. Lions can generate the
highest daily rate of any mammal hunted (USD $2,650 per day), the
longest number of days that must be booked, and the highest trophy fee
($24,500) (Jackson 2013, p. 6; Lindsey et al. 2012a, p. 5), thus
generating significant revenue for range countries. Creating community-
based incentives to conserve lions from revenue derived from trophy
hunting may ameliorate the human-lion conflict that arises from lions
and humans coexisting in the same area.
Revenue from scientifically based management programs that include
trophy hunting can increase the ability of many African countries to
manage wildlife populations both within and adjacent to reserves; many
of these hunting areas are geographically linked to national parks and
reserves, providing wildlife corridors and buffer zones (Chardonnet et
al. 2010, p. 34; Newmark 2008, p. 321). In the past, government and
private land owners were the primary beneficiaries of the revenue
gained; however, a portion of the revenue derived from hunting is
reportedly now being distributed to local communities, creating a value
for lions that encourages their conservation (Barnett & Patterson 2005,
p. iv). Revenue from trophy hunting is purported to create: (1)
Incentives for countries to conserve large tracts of prime habitat; and
(2) funding for park and reserve management, anti-poaching, and
security activities. Because habitat loss has been identified as one of
the primary threats to lion populations, it is notable that trophy
hunting has provided lion range states incentives to set land aside for
hunting throughout Africa, and the land set aside exceeds the total
area of the national parks, accounting for approximately half of the
amount of viable lion habitat (Chardonnet et al. 2010, p. 34; Packer et
al. 2006, pp. 9-10).
In Tanzania, which is home to 40 percent of all lions, land set
aside for sport hunting purposes has resulted in an area 5.1 times
greater than Tanzania's fully protected and gazetted parks (Jackson
2013, p. 6; Barnett & Patterson 2005, p. 61). In Botswana, despite the
current ban on lion hunting, the country currently has more than
128,000 km\2\ of gazetted wildlife management areas and controlled
hunting areas set aside for hunting purposes, which equates to 22.1
percent of the country's total area; this is in addition to 111,000
km\2\ (or 19.1 percent) that has been set aside as habitat in the form
of National Parks, Game Reserves, and Forest Reserves (Barnett &
Patterson 2005, p. 7). In 2000, five countries in southern Africa
(Botswana, Namibia, South Africa, Tanzania, and Zimbabwe) had set aside
a combined 420,000 km\2\ of communal land, 188,000 km\2\ of commercial
land, and 420,089 km\2\ of state land totaling more than 1,028,000
km\2\ for sport hunting purposes (Barnett & Patterson 2005, p. iii). As
a species with a considerable range (up to 1,000 km\2\) (Packer et al.
2013 p. 636; Haas et al. 2005, p. 4), suitable habitat is important to
the survival of the species, and the marked decline in suitable habitat
is a significant threat to the species. The habitat currently preserved
for use in sport hunting has helped to reduce the impact of habitat
loss for the African lion, but as discussed previously, habitat loss
remains a significant threat to the species.
Within its current range, the African lion exists in 10 stronghold
populations containing approximately 24,000 lions (70 percent of the
current African lion population), 19,000 of which are in protected
areas, and in 7 potential stronghold populations containing another
4,000 lions. Reports from the IUCN Species Survival Commission Cat
Specialist Group (IUN 2006a, b) characterize the population as
increasing in 3 of those strongholds, as stable in 6 of the
strongholds, and as decreasing in 1 stronghold. Most lion populations
in protected areas of southern and eastern Africa have been essentially
stable over the last three decades (Ray et al. 2005, p. 67). In
contrast to the stronghold or potential stronghold populations, other
African lion populations, containing a total of more than 6,000
individuals, have a very high risk of local extinction (Reggio et al.
2013, p. 33. During the 2005-2006 African lion workshops, lion experts
characterized lion populations in 36 (42 percent) of the 86 LCUs as
decreasing. In extensive surveys recently conducted within 15 of the 20
LCUs in western and
[[Page 64500]]
central Africa, Henschel et al. (2010, entire) were able to confirm
lion presence in only four. The work of Packer et al. (2013) suggests
future declines within a number of protected areas. Craigie et al.
(2010, entire) provide evidence of declining large mammal populations
in Africa's protected areas, indicating that protected areas in Africa
have generally failed to mitigate threats to large mammal populations,
including African lion. Although Craigie et al. (2010, p. 2,225) found
large regional differences (from large declines in western Africa to
positive rates of change in southern Africa), they found overall
populations decreased steadily from 1970 to 2005.
The best available scientific and commercial information leads us
to conclude that the African lion is in danger of extinction within the
foreseeable future throughout all of its range. Accordingly, we find
that listing is warranted and we propose to list it as a threatened
species throughout its range, wherever found.
Significant Portion of Its Range
Under the Act and our implementing regulations, a species may
warrant listing if it is endangered or threatened throughout all or a
significant portion of its range. The term ``species'' includes ``any
subspecies of fish or wildlife or plants, and any distinct population
segment [DPS] of any species of vertebrate fish or wildlife which
interbreeds when mature.'' We published a final policy interpreting the
phrase ``Significant Portion of its Range'' (SPR) (79 FR 37578, July 1,
2014). The final policy states that (1) if a species is found to be
endangered or threatened throughout a significant portion of its range,
the entire species is listed as endangered or threatened, respectively,
and the Act's protections apply to all individuals of the species
wherever found; (2) a portion of the range of a species is
``significant'' if the species is not currently endangered or
threatened throughout all of its range, but the portion's contribution
to the viability of the species is so important that, without the
members in that portion, the species would be in danger of extinction,
or likely to become so in the foreseeable future, throughout all of its
range; (3) the range of a species is considered to be the general
geographical area within which that species can be found at the time
FWS or NMFS makes any particular status determination; and (4) if a
vertebrate species is endangered or threatened throughout an SPR, and
the population in that significant portion is a valid DPS, we will list
the DPS rather than the entire taxonomic species or subspecies.
We found the African lion to be in danger of extinction within the
foreseeable future throughout all of its range. Therefore, no portions
of the species' range are ``significant'' as defined in our SPR policy
and no additional SPR analysis is required.
Proposed 4(d) Rule
The purposes of the ESA are to provide a means whereby the
ecosystems upon which endangered species and threatened species depend
may be conserved, to provide a program for the conservation of such
endangered species and threatened species, and to take such steps as
may be appropriate to achieve the purposes of the treaties and
conventions set forth in the ESA. When a species is listed as
endangered, certain actions are prohibited under section 9 of the ESA,
as specified in 50 CFR 17.21. These include, among others, prohibitions
on take within the United States, within the territorial seas of the
United States, or upon the high seas; import; export; and shipment in
interstate or foreign commerce in the course of a commercial activity.
The ESA does not specify particular prohibitions and exceptions to
those prohibitions for threatened species. Instead, under section 4(d)
of the ESA, the Secretary, as well as the Secretary of Commerce
depending on the species, was given the discretion to issue such
regulations as deemed necessary and advisable to provide for the
conservation of such species. The Secretary also has the discretion to
prohibit by regulation with respect to any threatened species any act
prohibited under section 9(a)(1) of the ESA. Exercising this
discretion, the Service has developed general prohibitions (50 CFR
17.31) and exceptions to those prohibitions (50 CFR 17.32) under the
ESA that apply to most threatened species. Under 50 CFR 17.32, permits
may be issued to allow persons to engage in otherwise prohibited acts
for certain purposes.
Under section 4(d) of the ESA, the Secretary, who has delegated
this authority to the Service, may also develop specific prohibitions
and exceptions tailored to the particular conservation needs of a
threatened species. In such cases, the Service issues a 4 (d) rule that
may include some or all of the prohibitions and authorizations set out
in 50 CFR 17.31 and 17.32 but which also may be more or less
restrictive than the general provisions at 50 CFR 17.31 and 17.32. For
the African lion, the Service has determined that a 4(d) rule is
appropriate.
We propose to add a 4(d) (special) rule for the African lion
(Panthera leo leo) at 50 CFR 17.40(n). This 4(d) rule would maintain
all of the prohibitions and exceptions codified in 50 CFR 17.31 and
17.32 and would supersede with regard to African lion the import
exemption found in 50 CFR 17.8 for threatened wildlife listed in
Appendix II of CITES, such that a threatened species import permit
under 50 CFR 17.32 would be required for the importation of all African
lion specimens. Through the promulgation of the proposed 4(d) rule, the
presumption of legality provided under Section 9(c)(2) of the Act for
the otherwise lawful importation of wildlife listed in Appendix II of
CITES that is not an endangered species listed pursuant to section 4 of
the Act would not apply to this subspecies. Thus, under the proposed
4(d) rule, all otherwise prohibited activities, including all imports
of African lion specimens, would require prior authorization or permits
under the Act. Under our regulations, permits or authorization to carry
out an otherwise prohibited activity could be issued for scientific
purposes, the enhancement of propagation or survival of the species,
economic hardship, zoological exhibitions, educational purposes, or
special purposes consistent with the purposes of the Act. Applications
for these activities are available from https://www.fws.gov/forms/3-200-37.pdf.
The intent of this proposed 4(d) rule is to provide for the
conservation of the African lion consistent with the purposes of the
Act. Under the proposed 4(d) rule, the prohibitions would, in part,
make it illegal for any person subject to the jurisdiction of the
United States to ``take'' (includes harass, harm, pursue, hunt, shoot,
wound, kill, trap, capture, or to attempt any of these) within the
United States or upon the high seas; import or export; deliver,
receive, carry, transport, or ship in interstate or foreign commerce,
by any means whatsoever, in the course of commercial activity; or sell
or offer for sale in interstate or foreign commerce any lion specimens.
It would also be illegal to possess, sell, deliver, carry, transport,
or ship any such wildlife that has been taken in violation of the Act.
We believe that these protections, including the requirement for an
import permit for all African lion specimens, will support and
encourage conservation actions for the African lion and require that
permitted activities involving lions are carried out in a manner that
is consistent with the purposes of the Act and our implementing
regulations.
[[Page 64501]]
In connection with this proposed 4(d) rule, the Service notes that
the African lion is listed in Appendix II of CITES, and thus can be
imported into the U.S. pursuant to Section 9(c)(2) of the Act and upon
presentation of a proper CITES export permit from the country of
origin. Section 9(c)(2) of the Act provides that the otherwise lawful
importation of wildlife that is not an endangered species listed
pursuant to section 4 of the Act, but that is listed in Appendix II of
CITES, shall be presumed to be in compliance with provisions of the Act
and implementing regulations. While there has been question as to
whether this provision of the Act might automatically require allowing
the importation of a species that is both listed as threatened and in
Appendix II, and preclude the issuance of more restrictive 4 (d) rules
covering importation, the Service has concluded that such 4 (d) rules
may be issued to provide for the conservation of the involved species.
Section 9(c)(2) does not expressly refer to threatened species or
prevent the issuance of appropriate 4 (d) rules and could not logically
have been intended to allow the addition of a species to an appendix of
an international convention to override the needs of U.S. law, where
there is reliable evidence to affect the presumption of validity.
Finally, the term ``presumed'' implies that the established presumption
is rebuttable under certain circumstances, including through the
promulgation of a protective regulation pursuant to section 4(d) of the
Act.
In the case of the African lion, there are substantive grounds on
which to challenge the presumption. For the import of sport-hunted
trophies, while there is evidence that many of the range countries are
implementing lion management plans, we want to encourage and support
efforts by these countries to develop plans that are based on sound
scientific information. As noted, the proposed 4(d) rule for African
lion would provide for the importation into the United States of
trophies taken legally in range countries upon the issuance of a
threatened species import permit. While the Service cannot control
hunting of foreign species such as African lion, we can regulate their
importation and thereby require that U.S. imports of sport-hunted
African lion trophy specimens are obtained in a manner that is
consistent with the purposes of the Act and the conservation of the
subspecies in the wild, by allowing importation from range countries
that have management plans that are based on scientifically sound data
and are being implemented to address the threats that are facing lions
within that country.
Such management plans would be expected to address, but are not
limited to, evaluating population levels and trends; the biological
needs of the species; quotas; management practices; legal protection;
local community involvement; and use of hunting fees for conservation.
In evaluating these factors, we will work closely with the range
countries and interested parties to obtain the best available
scientific and commercial data. By allowing entry into the United
States of African lion trophies from range countries that have
scientifically based management plans, the range countries would be
encouraged to adopt and financially support the sustainable management
of lions that benefits both the species and local communities. In
addition to addressing the biological needs of the subspecies, a
scientifically based management plan would provide economic incentives
for local communities to protect and expand African lion habitat.
As stated, anyone wishing to conduct any otherwise prohibited
activity, such as interstate commerce or imports, must first obtain a
permit under the current permitting regulations found at 50 CFR 13 and
50 CFR 17. As will all permits, the individual requesting authorization
to carry out an otherwise prohibited activity under the Act must submit
a permit application to the Service with specific information
concerning the proposed activity and the benefits/impacts of the
activity on the species. In some cases, such as imports of sport-hunted
trophies, it is not always possible for the applicant to provide all of
the necessary information needed by the Service to make a positive
determination under the Act to authorize the activity. For the import
of sport-hunted trophies, it is typical for the Service to consult with
the range country and other interested parties to obtain the necessary
information. To date, the Service typically has made the required
findings on sport-hunted trophy imports on a country-wide basis,
although individual import permits are issued for each applicant. While
the Service encourages the submission of information from individual
applicants, we would primarily rely on information from other sources
when making a permitting decision.
Effects of This Rule
This rule, if made final, would revise 50 CFR 17.11(h) to add the
African lion to the List of Endangered and Threatened Wildlife. This
rule, if adopted, would also establish a 4(d) rule for the African
lion, which implements all of the prohibitions and exceptions under 50
CFR 17.31 and 17.32 and requires a threatened species import permit
under 50 CFR 17.32 for the importation of all African lion specimens.
Under the proposed 4(d) rule, the import exemption found in 50 CFR 17.8
for threatened wildlife listed in Appendix II of CITES would not apply
to this subspecies. Through the promulgation of the proposed 4(d) rule,
the presumption of legality provided under Section 9(c)(2) of the Act
for the otherwise lawful importation of wildlife listed in Appendix II
of CITES that is not an endangered species listed pursuant to section 4
of the Act would not apply to this subspecies. (See: Proposed Special
Rule section).
Available Conservation Measures
Conservation measures provided to species listed as endangered or
threatened under the Act include recognition of conservation status,
requirements for Federal protection, and prohibitions against certain
practices. Recognition through listing encourages and results in public
awareness and conservation actions by Federal and State governments in
the United States, foreign governments, private agencies and groups,
and individuals.
Section 7(a) of the Act, as amended, and as implemented by
regulations at 50 CFR part 402, requires Federal agencies to evaluate
their actions that are to be conducted within the United States or upon
the high seas, with respect to any species that is proposed to be
listed or is listed as endangered or threatened. Because the African
lion is not native to the United States, no critical habitat is being
proposed for designation with this rule. Regulations implementing the
interagency cooperation provision of the Act are codified at 50 CFR
part 402. Section 7(a)(2) of the Act requires Federal agencies to
ensure that activities they authorize, fund, or carry out are not
likely to jeopardize the continued existence of a listed species or to
destroy or adversely modify its critical habitat. If a proposed Federal
action may adversely affect a listed species, the responsible Federal
agency must enter into formal consultation with the Service. Currently,
with respect to the African lion, no Federal activities are known that
would require consultation.
Section 8(a) of the Act authorizes the provision of limited
financial assistance for the development and management of programs
that the Secretary of the Interior determines to be necessary or useful
for the conservation of
[[Page 64502]]
endangered or threatened species in foreign countries. Sections 8(b)
and 8(c) of the Act authorize the Secretary to encourage conservation
programs for foreign listed species, and to provide assistance for such
programs, in the form of personnel and the training of personnel.
Section 9 of the Act and its implementing regulations at 50 CFR
17.31 set forth a series of general prohibitions that apply to all
threatened wildlife, except where a 4(d) rule applies, in which case
the 4(d) rule will contain all the applicable prohibitions and
exceptions. If the 4(d) rule is adopted as proposed, these prohibitions
would apply to the African lion. These prohibitions, in part, make it
illegal for any person subject to the jurisdiction of the United States
to ``take'' (includes harass, harm, pursue, hunt, shoot, wound, kill,
trap, capture, or to attempt any of these) within the United States or
upon the high seas; import or export; deliver, receive, carry,
transport, or ship in interstate or foreign commerce, by any means
whatsoever, in the course of commercial activity; or sell or offer for
sale in interstate or foreign commerce any lion specimens. It also is
illegal to possess, sell, deliver, carry, transport, or ship any such
wildlife that has been taken in violation of the Act. Permits may be
issued to carry out otherwise prohibited activities involving
threatened wildlife species under certain circumstances. Certain
exceptions apply to agents of the Service and State conservation
agencies.
Required Determinations
National Environmental Policy Act (42 U.S.C. 4321 et seq.)
We have determined that we do not need to prepare an environmental
assessment, as defined under the authority of the National
Environmental Policy Act of 1969, in connection with regulations
adopted under section 4(a) of the Act. We published a notice outlining
our reasons for this determination in the Federal Register on October
25, 1983 (48 FR 49244).
References Cited
A list of all references cited in this document is available at
https://www.regulations.gov at Docket No. FWS-R9-ES-2012-0025, or upon
request from the U.S. Fish and Wildlife Service, Endangered Species
Program, Branch of Foreign Species (see FOR FURTHER INFORMATION
CONTACT).
Authors
The primary authors of this proposed rule are staff of the Branch
of Foreign Species, Ecological Services, U.S. Fish and Wildlife
Service.
Proposed Regulation Promulgation
For the reasons described in the preamble, we propose to amend part
17, subchapter B of chapter I, title 50 of the Code of Federal
Regulations, as follows:
PART 17--[AMENDED]
0
1. The authority citation for part 17 continues to read as follows:
Authority: 16 U.S.C. 1361-1407; 1531-1544; and 4201-4245; unless
otherwise noted.
0
2. In Sec. 17.11(h), the List of Endangered and Threatened Wildlife,
add an entry for ``Lion, African'' under Mammals to read as follows:
Sec. 17.11 Endangered and threatened wildlife.
* * * * *
(h) * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
Species Vertebrate
-------------------------------------------------------- population where Critical Special
Historic range endangered or Status When listed habitat rules
Common name Scientific name threatened
--------------------------------------------------------------------------------------------------------------------------------------------------------
Mammals
* * * * * * *
Lion, African.................... Panthera leo leo.... Africa............. Entire............. T ........... NA 17.40(n)
* * * * * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
* * * * *
0
3. Amend Sec. 17.40 by adding paragraph (n) to read as follows:
Sec. 17.40 Special rules--mammals.
* * * * *
(n) African lion (Panthera leo leo).
(1) General requirements. All prohibitions and provisions of
Sec. Sec. 17.31 and 17.32 of this part apply to this subspecies.
(2) The import exemption found in Sec. 17.8 of this part for
threatened wildlife listed in Appendix II of the Convention on
International Trade in Endangered Species of Wild Fauna and Flora
(CITES) does not apply to this subspecies. A threatened species import
permit under Sec. 17.32 of this part is required for the importation
of all African lion specimens.
(3) All applicable provisions of 50 CFR parts 13, 14, 17, and 23
must be met.
* * * * *
Dated: October 20, 2014.
Stephen Guertin,
Director, Fish and Wildlife Service.
[FR Doc. 2014-25731 Filed 10-28-14; 8:45 am]
BILLING CODE 4310-55-P