Vol. 79 Wednesday, No. 209 October 29, 2014

Part II

Department of the Interior

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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DEPARTMENT OF THE INTERIOR FOR FURTHER INFORMATION CONTACT: (a) Genetics and taxonomy; Janine Van Norman, Chief, Branch of (b) Historical and current range, Fish and Wildlife Service Foreign Species, Ecological Services, including distribution; U.S. Fish and Wildlife Service, MS: ES, (c) Historical and current population 50 CFR Part 17 5275 Leesburg Pike, Falls Church, VA levels; 22041–3803; telephone, 703–358–2171; (d) Information pertaining to range [Docket No. FWS–R9–ES–2012–0025; 450 facsimile, 703–358–1735. If you use a countries’ regulatory mechanisms, 003 0115] telecommunications device for the deaf including specific laws and regulations RIN 1018–BA29 (TDD), call the Federal Information pertaining to loss of habitat, loss of prey Relay Service (FIRS) at 800–877–8339. base, and human-lion conflict. (e) Information pertaining to range Endangered and Threatened Wildlife SUPPLEMENTARY INFORMATION: and Plants; Listing the African Lion countries’ management plans, including Subspecies as Threatened With a Rule Executive Summary information on management and Under Section 4(d) of the ESA I. Purpose of the Regulatory Action implementation of hunting concessions, conservation measures in place for this AGENCY: Fish and Wildlife Service, Under the Act, a species may warrant subspecies and its habitat, community Interior. protection through listing if it is found education and outreach programs that ACTION: Proposed rule and 12-month to be an endangered or threatened address lion conservation, revenue finding. species throughout all or a significant gained from trophy hunting and how it portion of its range. Under the Act, if a is allocated, and any information SUMMARY: We, the U.S. Fish and species is determined to be endangered pertaining to long-term conservation of Wildlife Service (Service), announce a or threatened we are required to publish lions and their habitat and prey base; proposed rule and a 12-month finding in the Federal Register a proposed rule and on a petition to list the African lion to list the species. The purpose of this (f) Potential threats not already (Panthera leo leo) as endangered under proposed listing determination is to identified, such as extractive activities. the Endangered Species Act of 1973, as publish and seek comments on our 12- (2) The factors that are the basis for amended (Act). After review of the best month finding on a petition to add the making a listing determination for a available scientific and commercial African lion to the list of threatened and species or subspecies under section information, we find that listing the endangered species. 4(a)(1) of the Act (16 U.S.C. 1531 et subspecies Panthera leo leo as II. Major Provision of the Regulatory seq.), which are: threatened is warranted, and we Action (A) The present or threatened propose to list the subspecies as destruction, modification, or threatened. We are also proposing a rule After review of the best available curtailment of its habitat or range; under section 4(d) of the Act to provide scientific and commercial information, (B) Overutilization for commercial, for conservation measures for the we find that listing the African lion as recreational, scientific, or educational African lion. To ensure that subsequent threatened is warranted, and we purposes; rulemaking resulting from this proposed announce a proposed rule to list the (C) Disease or predation; rule is as accurate and effective as subspecies as threatened. We are also (D) The inadequacy of existing possible, we are soliciting information proposing a 4(d) rule to provide for regulatory mechanisms; or from the scientific community; other conservation measures for the African (E) Other natural or manmade factors governmental agencies, including those lion. affecting its continued existence. (3) The potential effects of climate within the range of the African lion; III. Costs and Benefits nongovernmental organizations; the change on the subspecies and its public; and any other interested parties. We have not analyzed the costs or habitat. benefits of this rulemaking action Please include sufficient information DATES: We will accept comments because the Act precludes consideration with your submission (such as scientific received or postmarked on or before of such impacts on listing and delisting journal articles or other publications) to January 27, 2015. We must receive determinations. Instead, listing and allow us to verify any scientific or requests for public hearings, in writing, delisting decisions are based solely on commercial information you include. at the address shown in FOR FURTHER the best scientific and commercial Submissions merely stating support for INFORMATION CONTACT by December 15, information available regarding the or opposition to the action under 2014. status of the subject species. consideration without providing ADDRESSES: You may submit comments supporting information, although noted, by one of the following methods: Information Requested will not be considered in making a (1) Electronically: Go to the Federal Section 4(b)(1)(A) of the Act directs determination. eRulemaking Portal: http:// that determinations as to whether any We request that you send comments www.regulations.gov. In the Search species is an endangered or threatened only by the methods described above in field, enter FWS–R9–ES–2012–0025, species must be made solely on the ADDRESSES. We will post all comments which is the docket number for this basis of the best scientific and on http://www.regulations.gov. If you rulemaking. Then, click the Search commercial data available. Therefore, submit information via http:// button. You may submit a comment by we request comments or information www.regulations.gov, your entire clicking on ‘‘Comment Now!’’ from other concerned governmental submission—including any personal (2) By hard copy: Submit by U.S. mail agencies, the scientific community, identifying information—will be posted or hand-delivery to: Public Comments industry, and any other interested on the Web site. If your submission is Processing, Attn: FWS–R9–ES–2012– parties concerning this proposed rule. made via a hardcopy that includes 0025, Division of Policy and Directives We particularly seek comments personal identifying information, you Management; U.S. Fish and Wildlife concerning: may request at the top of your document Service; MS: BPHC, 5275 Leesburg Pike, (1) The subspecies’ biology, range, that we withhold this information from Falls Church, VA 22041–3803. and population trends, including: public review. However, we cannot

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guarantee that we will be able to do so. all the independent peer reviewers, and endangered criterion level of 2,500 Please include sufficient information that all information regarding peer individuals, it is classified by the IUCN with your comments to allow us to review be included in the as regionally endangered (Bauer and verify any scientific or commercial administrative record. All proposed Nowell 2004, entire). In the assessment information you include. listing rules must be peer reviewed for this classification, western Africa is Comments and materials we receive, according to this policy and to defined as consisting of Benin, Burkina as well as supporting documentation we applicable standards under the Service’s Faso, Cote d’Ivoire, Gambia (identified used in preparing this proposed rule, guidelines for implementing the as ‘‘Regionally Extinct’’ (RE)), Ghana, will be available for public inspection Information Quality Act and the Guinea, Guinea Bissau, Liberia (RE), on http://www.regulations.gov, or by December 15, 2004, Office of Mali, Mauritania (RE), Niger, Nigeria, appointment, during normal business Management and Budget Final Senegal, Sierra Leone (RE), and Togo hours, at the U.S. Fish and Wildlife Information Quality Bulletin for Peer (Bauer and Nowell 2004, p. 35). Service, Ecological Services, Branch of Review. Convention on International Trade in Foreign Species (see FOR FURTHER Endangered Species of Wild Fauna and INFORMATION CONTACT). Petition History and Previous Federal Action(s) Flora (CITES) Public Hearing On March 1, 2011, we received a The African lion is listed in Appendix At this time, we do not have a public petition dated the same day from the II of the Convention on International hearing scheduled for this proposed International Fund for Animal Welfare, Trade in Endangered Species of Wild rule. The main purpose of most public the Humane Society of the United Fauna and Flora (CITES). CITES (see hearings is to obtain public testimony or States, Humane Society International, http://www.cites.org) is an international comment. In most cases, it is sufficient the Born Free Foundation/Born Free agreement through which member to submit comments through the Federal USA, Defenders of Wildlife, and the countries work together to protect eRulemaking Portal, described above in Fund for Animals requesting that the against over-exploitation of animal and ADDRESSES. If you would like to request African lion subspecies be listed as plant species found in international a public hearing for this proposed rule, endangered under the Act. The petition trade. Parties regulate and monitor you must submit your request, in identified itself as such and included international trade in CITES-listed writing, to the person listed in FOR the information as required by 50 CFR species—that is, their import, export, FURTHER INFORMATION CONTACT by the 424.14(a). On November 27, 2012, we and reexport, and introduction from the date specified in DATES. published a ‘‘positive’’ 90-day finding sea—through a system of permits and Peer Review (77 FR 70727) indicating that we would certificates. CITES lists species in one of initiate a status review of the African three appendices—Appendix I, II, or III. In accordance with our policy Species such as the African lion that are published on July 1, 1994 (59 FR lion. This document consists of our proposed rule and our determination on listed in Appendix II of CITES may be 34270), we will solicit the expert commercially traded, subject to several opinions of at least three appropriate the status review for the African lion and publishes our finding. Our status restrictions. CITES Appendix II includes and independent specialists for peer species that are less vulnerable to review of this proposed rule. The review may be obtained at http:// www.regulations.gov under Docket No. extinction than species listed in purpose of such review is to ensure that Appendix I, and ‘‘although not FWS–R9–ES–2012–0025. decisions are based on scientifically necessarily now threatened with sound data, assumptions, and analysis. Conservation Status of the African Lion extinction, may become so unless trade We will send peer reviewers copies of in specimens of such species is subject U.S. Endangered Species Act this proposed rule immediately to strict regulation in order to avoid following publication in the Federal The African lion (Panthera leo leo) is utilization incompatible with their Register. We will invite peer reviewers currently not listed as either endangered survival.’’ The status of the African lion to comment, during the public comment or threatened under the Act, although with respect to CITES and how it is period, on the specific assumptions and the Asiatic lion (Panthera leo persica) affected by international trade is conclusions regarding the proposed has been listed as endangered since discussed in more detail below, in the listing status of threatened for the 1970 under the Act and its precursor, section titled Import/Export of Lion African lion subspecies. We will the Endangered Species Conservation Parts and Products. summarize the opinions of these Act of 1969. reviewers in the final decision Periodic Review Under CITES International Union for the document, and we will consider their In an attempt to increase CITES Conservation of Nature input and any additional information protections for the African lion, in 2004, we receive, as part of our process of In 2008, the International Union for Kenya submitted a proposal for making a final decision on the proposal. the Conservation of Nature (IUCN) consideration at the Thirteenth Meeting Peer review is an important tool at our classified the African lion as vulnerable of the Conference of the Parties to CITES disposal to help evaluate the quality of with a declining population trend, (CoP13) to change the listing of the the data and analyses we rely on in our which means the species is considered African lion from Appendix II of CITES decision making processes. The 1994 to be facing a high risk of extinction in to Appendix I (CoP13 Prop. 6; http:// peer review policy commits us to the wild (Bauer et al. 2008, www.cites.org/eng/cop/13/prop/E13- soliciting the expert opinions of unpaginated). This classification is P06.pdf). An Appendix-I listing ‘‘appropriate and independent based on a suspected reduction in its includes species threatened with specialists regarding pertinent scientific population of approximately 30 percent extinction whose trade is permitted only or commercial data and assumptions over the previous two decades (Bauer et under exceptional circumstances, which relating to taxonomy . . . for species al. 2008, unpaginated). Because the generally precludes commercial trade. under consideration for listing.’’ The regional lion population in western The import of specimens (both live and policy also requires that our final Africa is isolated and estimated to dead, as well as parts and products) of decision must document the opinions of number well below the IUCN an Appendix-I species generally

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requires the issuance of both an import the Animals Committee, at its 25th Animals Committee took note of the and export permit under CITES. Import Meeting (AC25) (Geneva, Switzerland, upcoming Red List Assessment and permits are issued only if findings are July 2011), agreed to include the African requested Namibia and Kenya to made that the import would be for lion in the Periodic Review of Felidae incorporate this information into their purposes that are not detrimental to the [Decision 13.93 (Rev. CoP15)] (http:// Periodic Review and prepare a revised survival of the species in the wild and www.cites.org/eng/dec/valid15/E15- document for consideration at the 28th that the specimen will not be used for Dec.pdf) under the Animals Committee Meeting of the Animals Committee. primarily commercial purposes. For live periodic review of the appendices. Further, the Animals Committee made specimens, a finding must also be made Kenya and Namibia offered to lead the plans to continue seeking information that the recipient must be suitably review as a high priority with range from lion range states that had not yet equipped to house and care for the country consultation (http:// responded to requests for information specimens (CITES Article III(3)). Export www.cites.org/eng/com/ac/25/sum/E25- on the species. Finally, the Animals permits are issued only if findings are SumRec.pdf). At CoP16 in March 2013, Committee took note of the recent made that the specimen was legally the Parties adopted a revised Decision information concerning changes in the acquired and the export is not [Decision 13.93 (Rev. CoP16); http:// nomenclature of lion subspecies and detrimental to the survival of the www.cites.org/common/cop/16/sum/E- requested that the nomenclature expert species in the wild, and that a living CoP16-Plen-06.pdf; http:// of the Animals Committee review the specimen will be so prepared and www.cites.org/eng/dec/valid16/13_93_ information (CITES 2014b, p. 3). shipped as to minimize the risk of CoP16.php], directing the Animals Regions in Which African Lions Occur injury, damage to health, or cruel Committee to complete its Review of the treatment. (CITES Article III(2)). Appendices for Felidae and to provide The literature on African lion often Although Kenya had submitted its a report at CoP17 on the result of the includes reference to the following proposal to CoP13 for consideration, it review of all Felidae. Kenya and broad geographic regions: northern, withdrew its proposal due to the lack of Namibia recently submitted a report of western, central, southern, and eastern regional consensus on the proposal. their work on the Periodic Review of the Africa. The boundaries of these regions Furthermore, plans were under way at African lion for discussion at the 27th vary somewhat among authors, based on that time for convening a regional Meeting of the Animals Committee the nature and result of the studies workshop on lion management in 2005, (AC27, Veracruz, Mexico, 28 April–3 undertaken. the results of which would be reported May 2014) (CITES 2014a, entire). During As reflected in the literature reviewed to the CITES Animals Committee discussion of this document at AC27, a for this proposed rule, the lion (Animals Committee) (http:// representative of the IUCN informed the conservation community generally www.cites.org/eng/cop/13/rep/E13- committee that the IUCN would be works in the context of the regions of ComIRep13.pdf). completing an updated Red List Africa as they are described in Table 1. Recognizing that lion workshops and Assessment of the lion in 2015. In The regions as described in Table 1 may other research had been completed, addition, she suggested potential vary somewhat from the descriptions of producing updated information on the nomenclature changes to lion the regions that may be found in conservation and status of this species, subspecies (see Taxonomy). The 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.23 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 selection, regional environmental Characteristics include sharp, retractile adaptations, and gene flow (Mazak claws, a short neck, a broad face with The lion is the second-largest extant 2010, p. 194). These include, among prominent whiskers, rounded ears and a cat species (second in size only to the others, variation in size, coat color and muscular body. Lions are typically a tawny tiger) and the largest carnivore in Africa. color with black on the backs of the ears and It has a broad geographical range, thickness, mane color and form, and skull characteristics (Mazak 2010, p. white on the abdomen and inner legs. Males historically distributed throughout usually have a mane around the head, neck 194, citing several sources; Hollister Africa (Ray et al. 2005, p. 67). As with and chest. Lions are sexually dimorphic, other widely distributed large cats, there 1917, in Dubach 2005, p. 15). They are with males weighing about 20–27 percent is considerable morphological variation described by CITES (2014, p. 3) as more than females. Adult males, on average, within the species as a result of sexual follows: weigh about 188 kg with the heaviest male

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on record weighing 272 kg. Females are international organizations and to them (Barnett et al. 2014, pp. 1, 8). smaller, weighing, on average, 126 kg. The governing bodies. As a result, this is the Recently, Mazak (2010, entire) male body length, not including the tail, classification on which the conservation examined morphological characteristics ranges from 1.7 m to 2.5 m with a tail from of the species is largely based. However, of 255 skulls of wild lions and found 0.9 m to 1 m (Nowell & Jackson, 1996). results of recent genetic research call considerable variation throughout the Taxonomy into question this classification. species’ range, with variation being In recent years, several genetic studies The lion (Panthera leo) was first greater within populations than between have provided evidence of an described by Linnaeus (1758, in Haas et them. However, according to Dubach et evolutionary division within lions in al. 2005, p. 1), who gave it the name al. (2013, p. 742), the genetic distinction Africa (see Barnett et al. 2014, p. 6; Felis leo. It was later placed in the genus of lions in southern and eastern Africa Dubach et al. 2013, p. 746; Bertola et al. Panthera (Pocock 1930, in Haas et al. from those elsewhere in the species’ 2011 (entire); Antunes et al. 2008 2005, p. 1). Although the classification range is confirmed by results of studies (entire); Barnett et al. 2006a, pp. 511– by Antunes et al. (2008, entire) which, of the modern lion as Panthera leo is 512). These studies include analysis of in addition to analysis of mtDNA, also accepted within the scientific DNA samples from all major regions of included analysis of nDNA sequence community, there is a lack of consensus the species’ range, though some regions and microsatellite variation. regarding lion intraspecific taxonomy are represented by few samples. Results The recent results of genetic research (Mazak 2010, p. 194; Barnett et. al. of analysis indicate that a major genetic have renewed debate on lion taxonomy 2006b, p. 2,120). subdivision among lions occurs in among the experts. For this reason, the Based on morphology, traditional Africa, with lions in southern and IUCN Species Survival Commission Cat classifications recognize anywhere from eastern Africa being genetically distinct Specialist Group has commissioned a zero subspecies (classifying lions as one from and more genetically diverse than Cat Classification Task Force from monotypic species) up to nine lions elsewhere (western and central among its expert members to determine subspecies (Mazak 2010, p. 194, citing western and central Africa and Asia). a consensus taxonomy for the group. several sources). The most widely Evidence indicates that lions in western Until then, we conclude that the referenced of the morphology-based and central Africa (as well as now- taxonomy of the species is currently taxonomies is an eight-subspecies (six extinct north African lions) are more unresolved. However, as required by the extant) classification provided by closely related to lions in India than to Act, we base this status review on the Hemmer (1974, in Nowell and Jackson lions in southern and eastern Africa best available scientific and commercial 1996, p. 312; Barnett et al. 2006a, p. 507; (Barnett et al. 2014, pp. 4–8; Dubach et information, which is the most recent Barnett et al. 2006b, p. 2,120), which is al. 2013, pp. 741, 746–747, 750–751; taxonomy that is the most widely recognized by the Integrated Taxonomic Bertola et al. 2011, entire). According to recognized by taxonomic experts: P. leo Information System (ITIS) (ITIS 2013, Dubach et al. (2013, p. 753) leo (Africa) and P. leo persica (India). www.itis.gov, accessed June 6, 2013). It contemporary range collapse and Consequently, in this document we divides the lion species into: Panthera fragmentation is too recent a review the status of the petitioned leo persica (India); P. l. leo, commonly phenomenon to explain the lower entity, the African lion, P. leo leo. referred to as the Barbary lion (Morocco genetic variability in these regions. Range through Tunisia, extinct); P. l. Rather, the low genetic diversity in and senegalensis (West Africa east to the between western and central African Historically, lions occupied most of Central African Republic); P. l. azandica lion populations suggests they have a the African continent except the West (northern Zaire); P. l. bleyenberghi shorter evolutionary history than the African coastal rainforest zone, the (southern Zaire and presumably more genetically diverse lions in Congo Basin rainforest zone, and the neighboring areas of Zambia and southern and eastern Africa (Bertola et inner Sahara Desert (Bauer 2003, in Ray Angola); P. l. nubica (East Africa); P. l. al. 2011, p. 1362). Several authors argue et al. 2005, p. 67; IUCN 2006a, p. 10; krugeri (Kalahari region east to the that the origin of these genetically IUCN 2006b, p. 10). Ray et al. (2005, p. Transvaal and Natal regions of South distinct groups may be the result of 52) estimate lion historical range in Africa), and P. l. melanochaita, also regional extinctions and recolonizations Africa (at about 150 years prior to their called the Cape lion (Cape region of during major climate (and consequently study) to be roughly 22.2 million square South Africa, extinct) (Nowell and biome) fluctuations during the kilometers (km2), while IUCN (2006a, p. Jackson 1996, p. 312). Pleistocene Epoch (Barnett et al. 2014, 12; 2006b, p. 13) estimates lion In 1987, O’Brien (1987a, entire; pp. 5–8; Bertola et al. 2011, pp. 1,362– historical range in sub-Saharan Africa to 1987b, entire) reported the first results 1,364). be 19.3 million km2 (Table 2). of genetic studies conducted on lion These genetic studies on lion are Depending on the study and methods samples from some, but not all, regions based primarily on analysis of used, the species’ range is reported to of the species’ range using early genetic mitochondrial DNA (mtDNA), which is currently cover between 3.0 million and techniques. Results indicated that lions inherited only from the mother. Because 5.0 million km2 (Table 2). The most in India differed from lions in Africa, lions display sex-biased dispersal, in recent range-wide study was based on a supporting a two-subspecies which males leave their natal range and review of all of the most current classification for extant lions: P. leo leo females tend to remain in their natal available estimates of lion populations and P. leo persica, the African and range, one would expect gene flow in (up through 2012) (Riggio et al, p. 21), Asian lion, respectively (Ellerman et al. females to be lower than in males, combined with satellite imagery of 1953, Meester and Setzer 1971, O’Brien resulting in greater geographic savannah habitat, and provided et al. 1987, in Dubach 2005, p. 16). differentiation in females (Mazak 2010, estimates of current lion range to be 3.4 According to Dubach (2005, p. 16), most p. 204). Consequently, some authors million km2 (Riggio et al. 2013, p. 26), taxonomic authorities recognize this state that results of mtDNA analyses or about 25 percent of the subspecies’ two-subspecies taxonomy. This should be backed up by studies on historic range in savannah habitat. taxonomy is also recognized by the nuclear DNA (nDNA, inherited from According to Chardonnet (2002, pp. 24– IUCN (Bauer et al. 2012, unpaginated) both parents) and morphological traits 25), about half the range of the African and, consequently, by several before assigning taxonomic importance lion falls within protected areas.

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The African lion is now believed to be 2), the species’ range has declined by an 16). More recently, Henschel et al. extirpated from between 75 and 83 estimated 91 percent in western Africa, (2014, p. 5) estimate the confirmed lion percent of its former range (Table 2). 79 percent in central Africa, and 68 range in western Africa, based on data The subspecies has been extirpated from percent in eastern/southern Africa collected between 2006 and 2012, to be all of its former range in northern Africa (Table 2), with lion occurrence 49,000 km2, or an estimated 1.1 percent (Black et al. 2013, p. 1). In addition, unknown in an additional 38 percent of of the species’ former range in the according to IUCN (2006a,b; see Table the historical range (Bauer et al 2008, p. region.

TABLE 2—ESTIMATES OF THE AFRICAN LION RANGE

Current range as percent of historic Historic Current range Source Region of Africa range range 2 2 (percent of historic (km ) (km ) range w/unknown lion presence)

Ray et al. 2005: ...... Continent-wide ...... 22,200,000 3,800,000 17 percent. Chardonnet 2002: ...... Western ...... 121,980 Central ...... 651,970 Eastern ...... 1,137,205 Southern ...... 1,039,212

Total ...... 2,950,367 IUCN 2006a, b: 1 ...... Western ...... 3,814,576 331,749 9 percent. Central ...... 3,392,241 715,482 21 percent. Western + Central ...... 7,206,817 1,047,231 15 percent. Southern + Eastern ...... 12,080,000 3,915,000 32 percent.

Total ...... 19,286,817 4,962,231 26 percent. Bauer et al. 2008: 1 thnsp;2 ...... Western + Central ...... 7,206,817 1,047,231 15 percent. (0 percent). Southern + Eastern ...... 13,010,000 3,564,000 23 percent. (58 percent).

Total ...... 20,216,817 4,611,231 22 percent. (38 percent). Riggio 2013 3 (based on estimates of sa- Western ...... 133,784 vannah habitat):. Central ...... 936,465 Eastern ...... 780,401 Southern ...... 1,540,171 Total ...... 13,500,000 3,390,821 25 percent.

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

The historical range of the African recently, during surveys of 21 large of southern Chad and northern Central lion included most current continental protected areas in western Africa, African Republic, are virtually absent from African countries (Chardonnet 2002, pp. Henschel et al. (2014, p. 4) considered unprotected areas (Bauer 2003). 25–28). Currently, the subspecies occurs lions to be absent from protected areas Estimates of lion abundance on a large only in sub-Saharan Africa. Within this in 5 of these 10 countries (Table 1). geographical scale are few in number. region, Chardonnet (2002, p. 27) Distribution and Abundance For a variety of reasons—including low described lions as present in 34 range densities, large ranges, cryptic The general distribution of lions in states (35 with South Sudan, which coloration, nocturnal and wary habits— Africa is summarized by Ray et al. gained its independence as a country in lions are difficult to count (Bauer et al. (2005, p. 67) as follows: July 2011) and recently extirpated from 2005, p. 6; Riggio et al. 2013, p. 31). 6 range countries (Chardonnet 2002, p. Lions formerly occupied most of the There are large areas of the species’ 27) (Table 1). The 34 sub-Saharan African continent except for equatorial forest range in which no data are available on African range countries in which and the inner-Sahara. Today, they are extinct lion occurrence or abundance (IUCN in North Africa and have undergone dramatic Chardonnet considered lions present 2006b, pp. 12–13). Species experts included 10 in western Africa. More range retraction at the limits of their historical distribution. Currently, lions are recognize that estimating the size of the African lion population is an ambitious 1 Current range includes occasional and probable restricted mainly to protected areas and range. surrounding conservancies or ‘game task, involving many uncertainties 2 Bauer et al (2008) provides a synthesis of the management areas,’ with the largest (IUCN 2012, p. 2). Estimates, efforts from which the IUCN (2006a, b) estimates populations in East and southern Africa. particularly range-wide or broad region- were generated, providing somewhat different Where protection is poor, particularly wide estimates, tend to rely to a numbers for southern and eastern Africa. Also, outside protected areas, range loss or considerable extent on expert opinion or current range is range where lion occurrence is population decreases can be significant. known, and in approximately 38 percent of inference (Riggio et al. 2013, p. 21; historical range, the occurrence of lion is unknown. Declines have been most severe in West and Central Africa, with only small, isolated Chardonnet 2002, p. 19). Consequently, 3 Riggio et al. (2013) calculate estimates for there is a large degree of uncertainty in savannah habitat, defined as areas that receive populations scattered chiefly through the between 300 and 1,500 mm of rain annually and Sahel. Lions in the region are declining in these estimates. In addition, to date all which includes most of sub-Saharan Africa. some protected areas and, with the exception efforts to estimate the size of the African

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lion population have used different Specialist Group members made educated eastern Africa (Table 3). According to methods; the results of earlier estimates ‘‘guesstimates’’ of 30,000 to 100,000 for the Riggio et al. (2013, p. 27), only nine cannot be directly compared to those of African Lion population (Nowell and Jackson countries contain resident populations 1996). later estimates to determine population of at least 1,000 free-ranging lions trend. The earliest estimates of lion Ferreras and Cousins (1996, entire) (Central African Republic, Kenya, abundance in Africa were educated provided the first quantitatively derived Tanzania, Mozambique, Zambia, guesses made during the latter half of estimate using a GIS-based model Zimbabwe, South Africa, Botswana, and the 20th Century. Bauer et al. (2008, calibrated with information obtained possibly Angola). Approximately 40 unpaginated) summarize the from lion experts. Ferreras and Cousins percent of all lions are found in information as follows: predicted African lion abundance in Tanzania (Riggio et al. 2013, p. 27). There have been few efforts in the past to 1980 to be 75,800. Later, four additional Only about 10 percent of all lions occur estimate the number of lions in Africa. Myers efforts—Chardonnet (2002), Bauer and in western and central Africa (Table 3). (1975) wrote, ‘‘Since 1950, their [lion] Van Der Merwe (2004), IUCN (2006a, According to the most recent survey numbers may well have been cut in half, 2006b), and Riggio et al. 2013— effort, numbers in western Africa are perhaps to as low as 200,000 in all or even estimated lion population sizes ranging extremely low. Henschel et al. (2014, p. less.’’ Later, Myers (1986) wrote, ‘‘In light of from 23,000 to 40,000 (Table 3). evidence from all the main countries of its 5) estimate that only 400 lions in the range, the lion has been undergoing decline Currently, about 90 percent of all African lions occur in southern and entire region, with most (about 350, or in both range and numbers, often an 88 percent) concentrated in a single accelerating decline, during the past two eastern Africa (Table 3). According to decades’’. In the early 1990s, IUCN SSC Cat most studies, most African lions are in population.

TABLE 3—ESTIMATES OF AFRICAN LION ABUNDANCE [Rows may not tally due to rounding]

Western Central Source Africa Africa Eastern Africa Southern Africa Total (percent of total) (percent of total) (percent of total) (percent of total)

Ferreras & Cousins 1996 (estimate for ...... 75,800 (18,600 in lion abundance in 1980). protected areas). Chardonnet 2002 ...... 1,163 (3 percent) 2,815 (7 percent) 15,744 (40 per- 19,651 (50 per- 39,373. cent). cent). Bauer & Van Der Merwe 2004 ...... 850 (4 percent) .... 950 (4 percent) .... 11,000 (48 per- 10,000 (44 per- 23,000. cent). cent). IUCN 2006 4 (as calculated by Riggio 1,640 (5 percent) 2,410 (7 percent) 17,290 (52 per- 11,820 (37 per- 33,160. et al. 2013). cent). cent). Riggio 2013 (based on estimates of 480 (1 percent) .... 2,419 (7 percent) 19,972 (57 per- 12,036 (34 per- 34,907. savannah habitat). cent). cent). Henschel et al. 2014 ...... 406 (n/a).

In 2005–2006, in response to a range that can be considered an The majority of those with large growing concern that the African lion ecological unit of importance for lion populations were in southern and was in decline, IUCN and the Wildlife conservation (IUCN 2006a, p. 14; IUCN eastern Africa (Table 4). Only 23 of 86 Conservation Society sponsored 2006b, p. 17). Of the 86 LCUs, 20 are in LCUs (27 percent) were considered to workshops to determine a lion western and central Africa and 66 are in contain viable populations, though more conservation strategy. During these southern and eastern Africa (Table 4). than half were thought to contain workshops, lion experts collectively Most (71 percent) have more than half potentially viable populations (Table 4). assessed what they believed to be the their area under some form of legal Lion populations within 42 percent of then-current status of African lions protection (Bauer et al. 2008, p. 19). Few the 86 LCUs were considered to be based on a variety of information, (16 percent) were estimated to contain decreasing, whereas those in 9 percent including professional opinion. During large populations (Table 4). This was were considered increasing. The the workshops, lion experts identified particularly the case for western and 86 African lion Conservation Units central Africa, where most (13, or 65 remaining were considered stable or of (LCUs). They defined LCUs as areas of percent) of LCUs were estimated to unknown trend (Table 4). known, occasional, or possible lion contain fewer than 50 lions (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 Central Africa Africa (percent)

Total ...... 20 66 86. Estimated to contain: >500 lions ...... 2 12 14 (16 percent). 50–500 lions ...... 5 28 33 (38 percent).

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

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

Eastern & Number of LCUs Western & Southern All regions Central Africa Africa (percent)

<50 lions ...... 13 26 39 (45 percent). Considered: Viable ...... 4 19 23 (27 percent). Potentially Viable ...... 12 34 46 (53 percent). Doubtful Viability ...... 4 13 17 (20 percent). With Populations Considered to be: Increasing ...... 3 5 8 (9 percent). Stable ...... 5 21 26 (30 percent). Decreasing ...... 12 24 36 (42 percent). Unknown ...... 16 16 (19 percent).

Riggio et al. (2013, entire) provide the appears to be working. Potential population of at least 50 prides, but most recent, most comprehensive strongholds are described, broadly, as preferably 100 prides, with no limits to estimates to date of free-ranging lion areas where immediate interventions dispersal is required.’’ Bauer et al. 2008 populations in Africa. They compiled might create a viable population. (in Riggio et al. 2013, p. 32) indicate the all existing estimates of African lion Specifically defined, strongholds (1) average lion pride as containing populations since 2002, including data contain at least 500 lions, (2) are within approximately five adults. from Chardonnet (2002), Bauer and Van protected areas (including those that The results of Riggio et al. indicate the Der Merwe (2004), IUCN (2006a, 2006b), allow hunting), and (3) have stable or size of the African lion population to be over 40 mainly country-specific reports, increasing lion numbers as assessed by about 35,000, which falls within the and their own experiences. They then IUCN (2006a, 2006b) (Riggio et al. 2013, range of the other recent estimates combined these data with satellite p. 22). Potential strongholds contain at (Table 3). However, they state that imagery and information on habitat least 250 lions, but do not satisfy either ‘‘Although these numbers are similar to condition to estimate lion abundance requirement (2) or (3) above. The previous estimates, they are and identify lion areas that they remaining lion areas—those not meeting geographically more comprehensive. characterized as strongholds and the requirements of a stronghold or There is abundant evidence of potential strongholds. They conducted potential stronghold—are described as widespread declines and local this within the context of savannah extinctions’’ (Riggio et al. 2013, p. 18). Africa, which they defined as areas that areas ‘‘where present management receive between 300 and 1,500 clearly isn’t working’’ (Riggio et al. Riggio et al. identified lions as millimeters (mm) of rain annually, and 2013, p. 32). Riggio et al. (2013, p. 32) occurring in 67 areas (Table 5). While a within which most of the present range derived the thresholds of 500 and 250 small portion (22 percent) of lion areas of the African lion occurs. Also, they using information in Bjo¨rklund (2003) identified by Riggio et al. contain large used the LCUs identified in the 2005– on the number of prides needed to avoid populations, the majority are small and 2006 lion workshops as the general the risk of inbreeding in lion isolated (Riggio et al. 2013, p. 30; Table framework within which to identify lion populations, and information in Bauer 5). Most (69 percent) contain fewer than areas, strongholds, and potential et al. (2008) on the average size of lion 250 lions. A considerable portion (39 strongholds. prides. Bjo¨rklund (in Riggio et al. 2013, percent) contains very small Riggio et al. (2013, p. 32) describe lion p. 32) assessed the risk of inbreeding populations of fewer than 50 lions. strongholds as areas meeting the due to habitat loss and determined that, These include 63 percent of the lion necessary requirements for long-term ‘‘. . . to sustain a large out-bred areas in western and central Africa, and viability; broadly, where management population of lions, a continuous 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

All regions Number of lion areas Western Central Eastern Southern (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 approximately 24,000 lions, or about 70 4,000 lions. More than 6,000 lions are strongholds (viable populations) and 7 percent of the current African lion located in areas not considered potential strongholds (Table 6). population. Of those, most (about strongholds or potential strongholds and According to Riggio et al. (2013, p. 29), 19,000 lions) are in protected areas. have a very high risk of being extirpated the 10 strongholds contain Potential strongholds contain about (Riggio et al. 2013, p. 33).

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

Lion Population IUCN Lion area Country Area Stronghold population size in (2006a, b) (km2) protected size areas Trend

Western Africa

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

Central Africa

SE Chad ...... Chad ...... 133,408 Potential 5 ..... 400 140 Stable. E CAR ...... Central African Republic ...... 328,721 Potential 6 ..... 1,244 148 Stable.

Eastern Africa

Boma-Gambella ...... Ethiopia, South Sudan ...... 106,941 Potential ...... 500 ∼ 500 Unknown. Laikipia-Samburu ...... Kenya ...... 35,511 Potential ...... 271 46 Stable. Tarangire ...... Tanzania ...... 28,771 Potential ...... 731 208 Decreasing. Ruaha-Rungwa ...... Tanzania ...... 195,993 Stronghold .... 3,779 2,235 Stable. Selous ...... Tanzania ...... 138,035 Stronghold .... 7,644 4,953 Stable. Serengeti-Mara ...... Kenya, Tanzania ...... 35,852 Stronghold .... 3,673 3,516 Increasing. Tsavo-Mkomazi ...... Kenya, Tanzania ...... 39,216 Stronghold .... 880 820 Decreasing.

Southern Africa

Etosha-Kunene ...... Angola, Namibia ...... 123,800 Potential ...... 455 ∼ 315–595 Increasing. Kafue ...... Zambia ...... 58,898 Potential ...... 386 386 Stable. Great Limpopo ...... Mozambique, South Africa, 150,347 Stronghold .... 2,311 2,179 Increasing. Zimbabwe. Kgalagadi ...... Botswana, South Africa ...... 163,329 Stronghold .... 800 ∼ 800 Stable. 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.

protected areas. Packer et al. (2013a, increased or remained stable (see Most of the strongholds and potential entire; 2013b, entire) looked at the Distribution and Abundance), and some strongholds identified by Riggio et al. relationship between lion densities, have been restored to areas from which are trans-boundary areas. The vast population trends, management they were previously extirpated (Packer majority, including all 10 strongholds, practices, and several other variables et al. 2013, p. 636). Reports from the are located in southern and eastern (human population densities, IUCN Species Survival Commission Cat Africa. Of the 17 strongholds and governance, sport hunting, private Specialist Group (IUN 2006a, b) potential strongholds, only two management, and reserve size) from 42 characterize the population as potential strongholds are located in sites in 11 countries in Africa. Results increasing in 3 of the lion strongholds western and central Africa, one each in of modeling indicate that by 2050 about identified by Riggio et al. (Table 6), as western Africa and central Africa. Only 43 percent of lion populations in stable in 6 of the strongholds, and as a small portion of the lions in the unfenced reserves may decline to less decreasing in 1 stronghold. While four central Africa potential stronghold are than 10 percent of the carrying of the lion strongholds or potential within protected areas. The western capacities of the unfenced reserves, strongholds identified by Riggio et al. Africa potential stronghold has one of including those in Botswana, Kenya, (Table 6) are considered to be the smallest lion populations of the 17 Cameroon, Ghana, Tanzania, and increasing, several African lion strongholds/potential strongholds and, Uganda. According to the same populations, containing a total of more according to Herschel et al. (2014, p. 5), modeling results lion populations in than 6,000 individuals, have a very high contains 88–90 percent of all lions in fenced reserves are expected to remain risk of local extinction (Riggio et al. the western Africa region. at or above the carrying capacity of the 2013, p. 33). During the 2005–2006 By definition, all 10 strongholds fenced reserves for the next 100 years, African lion workshops, lion experts identified by Riggio et al. include although most are small protected areas characterized lion populations in 36 (42 with small lion populations (Creel et al. percent) of the 86 LCUs as decreasing. 5 Two lion areas in central Africa make up one 2013, entire). In extensive surveys recently conducted potential stronghold. within 15 of the 20 LCUs in western and 6 Riggio et al. make one exception to the Trends requirement that lion strongholds contain central Africa, Henschel et al. (2010, populations that are stable or increasing. IUCN 2006 Based on the best available entire) were able to confirm lion indicate lion numbers in the Tsavo/Mkomazi lion information, as discussed above, African presence in only four. The work of area are decreasing in numbers, but Riggio et al. lion range and numbers have clearly Packer et al. (2013) suggests future believe that, while lion numbers are declining outside of protected areas, lions within the parks declined over the past several decades. declines within a number of protected are usually well protected and in sufficient However, not all African lion areas. Craigie et al. (2010, entire) numbers to meet the criteria. populations have declined—some have provide evidence of declining large

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mammal populations in Africa’s Prides vary in size and structure, but Namibia, where Stander (1992) showed that protected areas, indicating that typically contain 5–9 adult females (range, 1– lions hunting in pairs met their minimum protected areas in Africa have generally 18), their dependent offspring, and a requirements hunting springboks which, at failed to mitigate threats to large coalition of 2–6 immigrant males (Heinsohn <50 kg, are the smallest preferred prey and Packer 1995; Packer et al. 1991). . . . species recorded. mammal populations, including African Pride sizes are smallest in arid environments lion. Although Craigie et al. (2010, p. Prey availability affects the with limited prey species (Elliott and Cowan reproduction, recruitment, and foraging 2,225) found large regional differences 1977; Hanby and Bygott 1979; Ruggiero 1991; (from large declines in western Africa to Schaller 1972; Stander 1992b; Wright 1960) behavior of lions and, as a result, positive rates of change in southern . . . Males reside in a pride for strongly influences lion movements, Africa), they found overall populations [approximately] 2 years before being replaced abundance, and population viability decreased steadily from 1970 to 2005. by another group of males (Packer et al. (Winterbach et al. 2012, p. 7, citing 1988). . . . In the absence of a pride takeover, several sources). Lion densities are Biology/Ecology males generally leave their natal pride when directly dependent on prey biomass 2–4 years old (Bertram 1975b; Pusey and Habitat (Van Orsdol et al. 1985, in Packer et al. Packer 1987). Most females are incorporated 2013a, p. 636; Hayward et al. 2007, Historically, the species occurred in into their natal prides (Pusey and Packer entire), and range from 0.08–0.13 adults all habitats in Africa, except rainforest 1987; Van Orsdol et al. 1985). . . . A small and subadults per 100 km2 in Selous and the hyper-arid interior of the Sahara proportion of lions is nomadic, including 2 young and adult males without a pride. Game Reserve up to 18 per km in (Ray et al. 2006, p. 66). Today they are protected areas of eastern Africa and found primarily in savannah, although Nomadic lions follow the migrations of prey and hunt and scavenge cooperatively South Africa (Creel and Creel 1997, there are some remnant populations in (Bertram 1975a; Bygott et al. 1979; Schaller Nowell and Jackson 1996, in Haas et al. other habitat types (Riggio et al. 2013, p. 1968, 1969; Van Orsdol et al. 1985). 2005, p. 4). Aside from human-related 19). According to Nowell and Jackson . . . Lion productivity (measured as mortality, prey availability is likely the (1996, p. 19), optimal habitat appears to number of surviving cubs) is limited by food. primary determinant of lion density be open woodlands and thick bush, . . . Cub mortality is high in lions and is (Fuller & Sievert 2001, in Winterbach et scrub, and grass complexes, where linked to periods of prey scarcity and al. 2012, p. 7). In areas of low natural sufficient cover is provided for hunting infanticide by male lions during pride prey density, or high human contact, takeovers (Packer and Pusey 1983b; Schaller and denning. The highest lion densities lions may prey on livestock (see are reached in savannah woodlands 1972; Van Orsdol et al. 1985; Whitman and Packer 1997). Human-Lion Conflict). plains mosaics of eastern and southern . . . Lions are mainly active at night . . . Movements/Home Range Africa (Ray et al. 2005, p. 66). The [They] usually hunt in groups; males hunt species is intolerant of anthropogenic less frequently than do females, but males are Availability of prey is perhaps the (human-caused) habitat conversion, stronger and can gain access to kills made by primary factor that determines the such as farming or overgrazing by females (Bertram 1975a; Scheel and Packer ranging behavior of large carnivores livestock (Ray et al. 2005, p. 66). 1991). Prey selection is related to seasonal (Gittleman & Harvey 1982, Van Orsdol weather patterns and the migration of large et al. 1985, Grant et al. 2005, Hayward General Biology herbivores in some parts of Africa (Hanby et et al. 2009, in Winterbach et al. 2012, Lions are well studied. Much al. 1995). . . . Lions exhibit individual p. 4). Home-range sizes of lion prides preferences in prey selection within and information exists on African lion correlate with lean-season prey biomass habits, behavior, and ecology. CITES between prides in the same area (Rudnai 1973b; Van Orsdol 1984). (Van Orsdol et al. 1985, in Haas et al. (2014a, p. 3) provides a general 2005, p. 4) and, therefore, vary widely overview as follows: Diet and Prey among habitats. Average range sizes of Lions are generalist, cooperative hunters, Lions are opportunistic hunters and African lion prides are 26–226 km2, but with foraging preferences changing with scavengers. As scavengers, lions are can be considerably larger (Stander season and with lion group size. Lions live dominant and can usually readily 1992b; Van Orsdol et al. 1985; Viljoen in groups called ‘‘prides’’, which are ‘‘fission- displace other predators from their kills 1993, in Haas et al. 2005, p. 4). In areas fusion’’ social units with a stable (Packer 1986, Schaller 1972, in Haas et of low or variable prey biomass, annual membership that sometimes divide into al. 2005, pp. 4–5). As hunters they are range requirements for a single lion small groups throughout the range. Lions 2 have no fixed breeding season. Females give known to take a variety of prey. pride can exceed 1,000 km (Packer et birth every 20 months if they raise their cubs However, they are also the largest al. 2013, p. 636). Funston (2011, p. 5) to maturity, but the interval can be as short carnivore in Africa and, as a result, found the home ranges of lion prides in as 4–6 weeks if their litter is lost. Gestation require large prey to survive. Ray et al. the dune-savannah habitat of Kgalagadi lasts 110 days, litter size ranges 1–4 cubs, (2005, pp. 66–67) summarizes lion prey Transfrontier Park to range from 1,762 and sex ratio at birth is 1:1. At about four as follows: to 4,532 km2. years of age, females will have their first litter Because lion home ranges can be very Lions are generalists and have been and males will become resident in a pride. large, many protected areas are not large Pride takeovers by male lions and subsequent recorded to consume virtually every mammal species larger than 1 kg in their range, as well enough to sustain them (Winterbach et infanticide of cubs sired by the ousted male al. 2014, p. 1; Funston 2011, p. 1, citing lions greatly influences reproductive success. as a wide variety of larger reptiles and birds Lionesses defending their cubs from the (Nowell & Jackson 1996; Sunquist & Sunquist several sources). Where lion ranges victorious males are sometimes killed during 2002). The constraints of large physical size approximate protected area size, lions the takeover. Infanticide accounts for 27 and extended social groups, however, bind roam near or beyond the protected area percent of cub mortality. Adult mortality is them to large-bodied prey, and their diet is boundary, increasing human-lion typically caused by humans, starvation, dominated by medium-large ungulates. In contact and human-caused lion disease or attacks from other lions. Injury and fact, only a few species of large ungulates mortality. In these situations, local or comprise a majority of their diet wherever death can also occur during hunting attempts regional extirpation probability is high on some of their larger prey. they occur (Schaller 1972; Stander 1992; Packer et al. 1995), and they are unable to due to the population sink created Haas et al. (2005, entire) provide a persist in areas without large-bodied prey. around the boundary of the protected summary of information on lion, The threshold of this requirement is perhaps area (Davidson et al. 2011, in including the following: represented at Etosha National Park, Winterbach et al. 2012, p. 5; Funston

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2011, p. 1, citing several sources; the amount of land required to meet the et al. (2013, p. 29) estimate that there Brashares et al. 2001, entire). This ‘‘edge needs of those populations is constantly were originally approximately 13.5 effect’’ is a major threat to carnivore increasing (Brink et al. 2014, entire; million km2 of savannah habitat in populations inside protected areas Brink and Eva 2009, entire; Eva et al. Africa. In 1960, 11.9 million km2 of throughout the world (Woodroffe 2001, 2006, p. 4), a problem accentuated by these habitats had fewer than 25 people in Winterbach et al. 2012, p. 5) (also see slow rates of technological progress in per km2, and in 2000 this number Human-Lion Conflict). food production and land degradation decreased to 9.7 million km2. Based on from both overuse and natural causes analysis of land-use conversion using Habitat Loss (United Nations Environment satellite imagery and human population Habitat loss and degradation is Programme (UNEP) 2012a, p. 3; densities, Riggio et al. (2013, p. 29) reported to be among the main threats Chardonnet et al. 2010a p. 19; found current savannah habitat that is to African lions (IUCN 2006a, p. 18; Ray International Assessment of Agricultural suitable for lions to be fragmented and et al. 2005, pp. 68–69). The main cause Knowledge, Science and Technology for to total about 3.4 million km2 (or 25 of lion habitat loss and degradation is Development (IAASTD) 2009, pp. 3–4, percent of African savannah habitat). expansion of human settlements and 8; United Nations Economic These data suggest a substantial activities, particularly agriculture and Commission for Africa 2008, pp. 3–5). decrease in lion habitat over the past 50 intensive livestock grazing in lion The result of this process is accelerated years. habitat (IUCN 2006a, p. 18; IUCN 2006b, transformation of natural landscapes at Projections of future human p. 23; Ray et al. 2005, pp. 68–69; the expense of wilderness that sustains population growth, area of conversion to Chardonnet 2002, pp. 103–106). species such as lions and their prey agriculture, and livestock numbers in Expansion of human settlements and (Chardonnet et al. 2010a p. 19). From Africa suggest suitable lion habitat will activities into lion habitat renders the 1970 to 2000, the human population in continue to decrease into the foreseeable habitat unsuitable for lions primarily sub-Saharan Africa increased by 126 future. Africa has the fastest population because it results in reduced availability percent (from 282 million to 639 growth rate in the world (UNEP 2012a, of the wild prey that lions depend on for million) (United Nations (UN) 2013, p. p. 2). Future population growth in sub- survival (see Loss of Prey Base) and 9), while at about the same time (1975 Saharan Africa is projected to be large increased human-lion conflict resulting to 2000), there was a 57 percent increase and rapid (UN 2013, p. 9). Although in lion mortality (see Human-Lion in agriculture area (from just over 200 urbanization is increasing in sub- Conflict)—two of the main factors that million ha to almost 340 million ha) and Saharan Africa (UN 2014, p. 20), the influence the distribution and 21 percent decrease in natural majority of the population is rural, and population viability of large carnivores vegetation in the region (Brink and Eva about 60–70 percent of the population such as lions (Winterbach et al. 2014, p. 2009, p. 507). In 2009, approximately relies on agriculture and livestock for 1). Ray et al. (2005, p. 69) note that, 1.2 billion ha, or 40 percent, of Africa’s their livelihood (UNEP 2006, pp. 82, although lions have a wide tolerance for land area was in permanent pasture or 100, 106; IAASTD 2009, p. 2). Much of habitats, they are generally incompatible crops, with the vast majority (31 the agriculture and livestock-raising is with humans and human-caused habitat percent) in pasture (UNEP 2012b, p. 68). at subsistence level (IAASTD 2009, pp. alteration and loss. Lions are sensitive Growing human populations have 8, 28). As a result, a large portion of the to loss of cover or prey. Riggio et al. been associated with declines in large growing population will depend (2013, p. 18) state that dense human carnivore populations all over the directly on expansion of agriculture and populations and widespread conversion world, and high human density is livestock grazing to survive. Between of land to human use preclude use by strongly associated with local 2010 and 2050 the population of sub- lions. extirpation of large carnivores (Linnell Saharan Africa is projected to more than Habitat destruction and degradation et al. 2001, Woodroffe 2001, in double to more than 2 billion (from 831 has been extensive throughout the range Woodroffe and Frank 2005, p. 91; million to 2.1 billion) (UN 2013, p. 9). of the African lion, resulting in local Woodroffe 2000, entire). Chardonnet et During about this same time period and regional lion population al. (2002, p. 103) indicate that the (2005 to 2050), Alexandratos and extirpations, reduced lion densities, a distribution maps of lion Bruinsma (2012, p. 107) project the area dramatically reduced subspecies range subpopulations tend to confirm a direct of cultivated land to increase by 51 (see Range), and small, fragmented, and inverse correlation of lion density and million ha (approximately 21 percent). isolated lion populations that are numbers with human activity and However, this figure does not include increasingly limited to protected areas presence. Further, Packer et al. (2013, range land, and the majority of (see Distribution and Abundance) (Ray entire) found that lions in unfenced agricultural land in Africa is devoted to et al. 2005, p. 69; Bauer and Van der reserves are highly sensitive to human grazing (UNEP 2012b, p. 68). The Merwe 2004, pp. 29–30; Nowell and population densities in surrounding number of livestock (cattle, sheep, and Jackson 1996, pp. 20–21). Lions appear communities. goats) in sub-Saharan Africa is projected to have one of the lowest levels of Based on a comparison of land-use to increase about 73 percent, from 688 ecological resilience to human-caused and human population data, Riggio et million to 1.2 billion, by 2050 habitat fragmentation; they are the least al. (2013, p. 23) determined that a (Alexandratos and Bruinsma 2012, p. successful large African carnivore density of 25 or more people per km2 133). outside conservation areas (Woodroffe served as a proxy for the extent of land- Expansion of human settlements, 2001, in Winterbach et al. 2012, p. 6). use conversion that would render agriculture, and/or livestock grazing are Large carnivores with low ecological habitat unsuitable for lions. Woodroffe reported as occurring in or on the resilience have a high risk of local (2000, p. 167) analyzed the impact of periphery of several of the areas extinction. In order to survive, they people on predators by relating local identified by Riggio et al. (2013, suppl. require larger contiguous habitats with carnivore extinctions to past and 1) as lion strongholds (viable lower negative human impacts than do projected human population densities populations) and potential strongholds more resilient species (Winterbach et al. and estimated 26 people per km2 as the (IUCN 2006a, p. 16; IUCN 2006b, pp. 2012, p. 5). As human populations mean human density at which lions 20–22), and are particularly a threat in continue to rise in sub-Saharan Africa, went locally or regionally extinct. Riggio western, central, and eastern Africa and

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some parts of southern Africa. There are population densities relate the number human population in sub-Saharan only two potential strongholds in of humans to land area and, Africa is rural, and land supports the western and central Africa (one in each consequently, are indicative of the level livelihood of most of the population, region). Expansion of agriculture and of pressure that will exist to convert loss and degradation of lion habitat can livestock grazing are reported in or land to uses that will meet the needs of be expected to accompany the rapid around both (Heschel et al. 2014, pp. 5– the human population. This is growth in sub-Saharan Africa’s human 6; Houessou et al. 2013, entire; particularly the case given that much of population. Therefore, overall, because Chardonnet et al. 2010, pp. 24–26; IUCN sub-Saharan Africa is rural and locals (1) lion prides have vast ranges and the 2008, pp. 8, 28–29), and management of depend on agriculture for their subspecies requires large areas of protected areas in portions of both is livelihood. suitable habitat to survive, (2) the reported as weak (Heschel et al. 2014, In southern Africa, the extent of subspecies’ range has already declined pp. 5–6; IUCN 2008, p. 8). Eastern current habitat destruction and dramatically and is increasingly limited Africa contains over half of all the lions degradation appears to vary widely. For to protected areas, and (3) habitat loss in Africa (Table 3). Seven of the example, according to the Zambia and degradation is occurring in or seventeen African lion strongholds and Wildlife Authority (2009 pp. 4–5), around several of the remaining lion potential strongholds identified by unplanned human settlement and other strongholds (viable populations) and Riggio et al. occur in eastern Africa, and land-use activities in game management potential strongholds, we conclude six of those seven (all four strongholds areas are a major threat to the long-term based on the best available scientific and two of three potential strongholds) survival of the lion in Zambia. They and commercial information that the are located in Tanzania and Kenya note that conversion of natural habitat continued destruction, modification, (Table 6). in game management areas for cropping and curtailment of lion habitat is likely Between 1990 and 2010, Kenya’s and grazing of livestock has led to to become a significant threat to the human population grew from 23 million habitat destruction and indicate that African lion throughout its range. (40/km2) to 41 million (70/km2), elimination of tsetse flies and whereas Tanzania’s grew from 25 subsequent increase in pastoralist Human-Lion Conflict million (27/km2) to 45 million (48/km2) activities in game management areas Human-lion conflict and associated (UN 2013, pp. 421, 798). Not places the lion under renewed direct retaliatory killing of lions has played a unexpectedly, sources indicate that conflict with humans. On the other major role in the reduction of lion expansion of agriculture and livestock hand, according to Funston (2008, populations (Lion Guardians 2013, p. 1; grazing is occurring in these countries pp. 123–126), in several areas of Lion Guardians 2011, p. 2; Hazzah and (Brink et al. 2014, entire; UNEP 2009, p. southern Africa where lions were Dolrenry 2007, p. 21; Frank et al. 2006, 91; Mesochina et al. 2010, p. 74), recently extirpated, lions are p. 1; Patterson et al. 2004, p. 508) and including in or around lion strongholds reestablishing as a result of, among is the greatest threat to remaining lion and potential strongholds (Ogutu et al. other factors, adequate protection of populations (Hazzah et al. 2009, p. 2011, entire; Mesochina et al. 2010, pp. habitat and prey. Human population 2,428; Moghari 2009, p. 31; Kissui 2008, 71–74, 76; Packer et al. 2010, pp. 8–9; growth, and resulting pressures exerted p. 422; Frank et al. 2006, pp. 1, 3, 10; UNEP 2009, pp. 98–99; Newmark 2008, on habitat, are also expected to vary Ray et al. 2005 in Hazzah 2006, p. 2; pp. 322–324; IUCN 2006b, pp. 20–22; widely in the region. Population IUCN 2006b, p. 18). Conflict between Ogutu et al. 2005, entire). Mesochina et increases from 2010 to 2050 are humans and wildlife has been linked to al. (2010, p. 74) state that widespread projected to range from about 23 percent population declines, reduction in range, destruction of wildlife habitat and (South Africa) to well over 200 percent impacts to small population human encroachment in wildlife (Zambia), with 2050 densities in the demographics, and even species corridors are major threats to lion region ranging from 5 people per km 2 extinctions (Dickman 2013, p. 377; Begg conservation in Tanzania and consider (Botswana and Namibia) to 348 people and Begg 2010, p. 2; Hazzah et al. 2009, loss of suitable habitat as a top threat to per km 2 (Malawi) (Table 7). p. 2,428; Moghari 2009, p. 36; Kissui lion survival in the country. In Kenya, 2008, p. 422; Hazzah 2006, pp. 15, 23, Summary of Habitat Loss the Kenya Wildlife Service (2009, p. 21) 25). indicates that habitat loss due to land- In the past several decades the human Human-wildlife conflict stems from use changes and human encroachment population has been expanding with human population growth and the into previously wild areas is having a concomitant large decreases in lion resulting overlap of humans and major impact on lion range size. By habitat and lion populations, resulting wildlife habitat (Chardonnet et al. 2010, 2050 the UN projects the human in an extremely large reduction in the p. 6; Hazzah 2006, pp. 14, 15). Lion population of Tanzania to almost triple species’ range. Habitat for African lion populations are increasingly restricted its 2010 population, reaching a density continues to be threatened with to protected areas, due to human of 137 people per km2, whereas Kenya’s destruction, modification, and expansion and associated expansion of population is projected to more than curtailment. Human populations are livestock husbandry and agricultural double, reaching a density of 167 people projected to increase dramatically in activities. However, despite being per km2 (Table 7). sub-Saharan Africa in coming decades. within protected areas, lions continue to The human populations of most other As human populations continue to rise be impacted by people living on current and recent lion range countries in sub-Saharan Africa, the amount of adjacent land. Villages are established are also expected to have very high land required to meet the expanding on the borders of protected areas, cattle growth rates (Table 7). It is important to human population’s needs is constantly herders enter the protected areas, and note that the country-wide human increasing. In addition, as indicated lions move beyond the borders of population densities provided here (and above, lions are increasingly limited to protected areas in search of food, in Table 7) are not directly comparable protected areas, and human population increasing interactions between humans to the density thresholds determined by growth rates around protected areas in and lions and the risk of human-lion Riggio et al. (discussed above) due to the Africa tend to be higher than the average conflict (Hazzah et al. 2013, p. 1; differences in scale at which they were rural growth rate (Wittemyer et al. 2008, Republic of Namibia 2013, p. 13; made. However, country-wide entire). Considering the majority of the Chardonnet et al. 2010, pp. 11–12;

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Mesochina et al. 2010a, p. 39; range (Chardonnet et al. 2005, p. 9; can be expected, and some lions can Mesochina et al. 2010b, p. 33; Packer IUCN 2006b, p. 19). In addition, the sale become habitual livestock killers (Frank et al. 2010, pp. 2, 6; Gebresenbet et al. of bushmeat is an important livelihood et al. 2006, p. 9). 2009, p. 9; Moghari 2009, pp. 1, 14, 25, in Africa, (Chardonnet et al. 2010, p. 27; Studies have shown variation in rates 26, 78; Kissui 2008, p. 422; Hazzah Mesochina et al. 2010a, p. 38; Abwe and of livestock depredation with regional 2006, p. 2). The most significant cause Morgan 2008, p. 26; Bennett et al. 2007, rainfall that correlate with prey of human-lion conflict is livestock p. 885; Fa et al. 2006, p. 507). This availability, including changes in depredation. Poor husbandry practices growing demand and widely available herding strategies, movement of prey, and grazing of livestock within or modern weapons has led to increased and movement of lions (Lion Guardians adjacent to protected areas increase poaching of native wildlife (Chardonnet exposure of livestock to lions and et al. 2010, pp. 13–14, 27; Packer et al. 2011, p. 6; Moghari 2009, p. 32; Hazzah increase livestock loss (Uganda Wildlife 2010, p. 8). Because many wildlife 2006, pp. 17, 18; Patterson et al. 2004, Authority 2010, p. 27; Woodroffe and species are being hunted at p. 514). For example, in some parts of Frank 2005 in Moghari 2009, p. 35; unsustainable levels to meet this Zimbabwe, Kenya, and Tanzania, Hazzah and Dolrenry 2007, pp. 22–23). demand within the range of the lion, its livestock losses occur during the dry Although lions generally avoid people, prey base is becoming depleted in many season. During this time, herders travel they will occasionally prey on humans, areas, which has led lions to seek out further for forage and water, they use causing serious injury or death livestock (and in some cases, humans) temporary bomas (a livestock enclosure) (Dickman 2013, pp. 380, 384; for food (Hoppe-Dominik et al. 2011, p. that are typically weak, they are Chardonnet et al. 2010, pp. 11, 12, 13; 452; Chardonnet et al. 2010, pp. 6, 13– unfamiliar with carnivore movements in Moghari 2009, pp. 14, 49, 26, 88; Bauer 14; Frank et al. 2006, p. 12). these new areas, and livestock are weak et al. 2001 in Moghari 2009, pp. 31, 78, Further, the demand for agriculture to due to disease, which makes them more 84; Frank et al. 2006, p. 1; Hazzah 2006, meet the increasing needs of a growing vulnerable to predator attacks by lions pp. 14, 17; Patterson et al. 2004, p. 507). population has been met by intensified (Hazzah 2006, p. 17). Additionally, Attacks on humans appears to be more agricultural and livestock practices herders are dependent on resources frequent in southern and eastern Africa (Chardonnet et al. 2010, p. 19). As within protected areas, and livestock (Chardonnet et al. 2010, pp. 12, 13; natural habitats are converted to may be left to wander for days or weeks Mesochina et al. 2010a, pp. 29–30; agricultural or pastoral land, it removes during a prolonged drought to find Frank et al. 2006, pp. 1, 10). Lion attacks the food and cover needed by wildlife, forage, increasing opportunities for can have various impacts on those and the lion’s natural prey base is attacks on livestock by lions communities that coexist with conflict- reduced, causing them to prey on (Chardonnet et al. 2010, p. 24; Frank et causing animals, generating resentment domestic livestock (Chardonnet et al. al. 2006, p. 6). In other parts of Kenya, towards them. When lions cause or are 2010, p. 27; Gebresenbet et al. 2009, the Maasai Steppe region of Tanzania, perceived to cause damage to livestock, p. 9). and Queen Elizabeth National Park, property, or people, the response is In Tanzania, which is home to more Uganda, livestock losses were greater generally to kill them (Dickman 2013, than 40 percent of the African lion during or following the rainy season pp. 378–379; Moghari 2009, p. 25; Frank population, conversion of rangeland to (Moghari 2009, p. 88; Kissui 2008, pp. et al. 2006, p. 1). agricultural use has blocked several 427, 428; Frank et al. 2006, p. 6; migratory routes for wildebeest and Loss of Prey Base Patterson et al. 2004, pp. 510, 514). zebra populations, both lion prey Weakened prey and readily available The lion’s prey base has decreased in species, which likely forces lions to rely carcasses provide easy meals during many parts of its range for various more on livestock (Packer et al. 2010, p. times of drought, leading to fewer reasons, but a large factor is due to 9). Conditions worsen as livestock livestock attacks. However, when rains competition for meat by humans. numbers and area under cultivation return, the abundant grass makes wild Humans in Africa rely on protein increase, leading to overgrazing, further obtained from bushmeat, resulting in habitat destruction, and greater prey harder to catch and lions may turn direct competition for prey between depredation rates by lions (Gebresenbet to livestock. Migratory prey species, humans and lions, and commercial et al. 2009, p. 9; Hazzah 2006, p. 61; such as zebra and wildebeest, will move poaching of wildlife is becoming a Frank et al. 2005, Ntiati 2002, Mishra to other areas for forage and replenished significant threat to many species, 1997, Meriggi and Lovari 1996, Rao water sources, leaving lions to turn to including those that lions rely upon for 1996, Mech et al. 1988 in Hazzah 2006, livestock as an alternate food source. food. Historically, subsistence hunting p. 18). Additionally, the use of fences to Migratory prey may also move outside with spears was traditionally used to subdivide group ranches interferes with of protected areas. Opportunities for hunt wildlife, which had minimal traditional wet and dry season grazing livestock predation on communal land impact to wildlife populations. Spears schedules for livestock and wildlife increase when lions follow (Packer et al. have since been replaced by automatic (Hazzah 2006, pp. 58–59). Restricting 2010, p. 9; Kissui 2008, p. 427; Patterson weaponry (Chardonnet et al. 2010, wildlife movement reduces wild prey et al. 2004, p. 514; Frank et al. 2006, p. p. 27), allowing for poaching of large and, when combined with an increase 6). Similarly, environmental factors numbers of animals for the bushmeat in livestock numbers, increases the rate such as vegetative cover, habitat, trade. of human-lion conflict (Hazzah 2006, climate, seasonality, and prey The human population in a majority pp. 59, 61). Although well-built bomas availability may affect the rate of attacks of African countries within the range of can effectively constrain cattle and keep on humans. A certain amount of the lion has quadrupled since the 1960s predators out (Frank et al. 2006, p. 8), vegetative cover is crucial for hunting (Riggio et al. 2013, p. 29; IUCN 2009, they are traditionally built to keep success; however, in some cases, the p. 15), increasing the demand for livestock confined, but do not offer vegetative cover may make it more bushmeat. Bushmeat comprises between effective protection from predators difficult to catch prey, leading to more 6 percent (southern Africa) and 55 (Moghari 2009, p. 35). In the absence of attacks on humans. Additionally, dense percent (Central African Republic) of a reliable methods for protecting cover near settlements allows lions to human’s diet within the African lion’s livestock, some amount of depredation hide or stalk humans at a close distance

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(Mesochina et al. 2010a, p. 39; Moghari protecting crops (Frank et al. 2006, p. rates of lions killed per 100 km2, yet it 2009, p. 85; Frank et al. 2006, p. 12). 12). may be that just under half of the lions that are killed are reported (Republic of Attacks on Livestock Retaliatory Killing of Lions Namibia 2013, p. 14). Although most of Traditional livestock husbandry Competition with humans, habitat the information on human-lion conflict practices are effective at reducing changes, and regional climate variations comes from just a few areas of the lion’s depredation of livestock by lions can decrease availability of prey and range (e.g., Kenya, Tanzania, and (Chardonnet et al. 2010, p. 35; Moghari increase human-lion conflict. When Uganda), it is reasonable to conclude 2009, p. 35; Frank et al. 2006, p. 2; native prey are unavailable or difficult that lions are being killed due to conflict Hazzah 2006, p. 22). These practices to find and kill, lions will target in all major range countries, due to their include livestock being closely herded domestic livestock or humans depredation on livestock (Frank et al. by men and dogs during the day and (Chardonnet et al. 2010, p. 27; Moghari 2006, p. 4). being brought into bomas at night with 2009, pp. 78, 83; Hazzah 2006, pp. 17– In areas of high conflict, identifying people living in huts around them 18; Patterson et al. 2004, pp. 507, 514). the responsible animal is often difficult, (Frank et al. 2006, p. 4). However, these Lion attacks occur at the highest and a token animal may be killed traditional practices are being replaced frequency in areas where natural prey instead (Hazzah 2006, p. 25), leaving the by less diligent husbandry practices, abundance is lowest (Packer et al. 2010, problem lion to continue to attack and which are increasing conflict p. 9; Frank et al. 2006, pp. 9, 12; the potential for additional retaliatory (Woodroffe and Frank 2005 in Moghari Patterson et al. 2004, p. 507). Livestock killings. In Tanzania, game officers kill 2009, p. 35; Frank et al. 2006, pp. 2, 10; provide an economic value to humans, numerous lions each year in retaliation Hazzah and Dolrenry 2007, p. 23). In particularly those in extreme poverty for attacks (Frank et al. 2006, p. 12). Botswana, livestock are often left to who rely solely on livestock for their Whereas shooting or spearing target wander outside bomas at night (Frank et protein source and livelihood. When specific problem animals, poisoning is al. 2006, p. 5). In Kenya and Tanzania, lions have no economic value to local indiscriminate and is known to remove social changes are altering traditional communities, and they kill or are entire prides at once (Frank et al. 2006, Maasai pastoral livelihoods, reducing perceived to kill livestock that do have pp. 2, 10, Living with Lions no date, dependency on livestock, and reducing an economic value to people, they are unpaginated). In the absence of reliable traditional livestock care and subject to retaliatory killing. This greatly methods for protecting livestock, rural management, leaving livestock more impacts already-dwindling lion people often turn to indiscriminant vulnerable to predation (Chardonnet et populations (Chardonnet et al. 2010, pp. methods, like poisoning, to control al. 2010, p. 35; Hazzah and Dolrenry 12–14; Mesochina et al. 2010a, p. 38; livestock depredation. Poisoning is an 2007, pp. 22–23). Young Maasai boys Mesochina et al. 2010b, p. 32; easy method for lethal control since it traditionally guarded herds at night; Gebresenbet et al. 2009, p. 9; Moghari is readily available, and reinforcing however, increased access to schools 2009, pp. 4, 25, 49; Kissui 2008, pp. 423, bomas or more carefully tending has left herds unattended to wander into 429; Hazzah 2006, p. 24; IUCN 2006a, livestock requires time and effort. The predator areas at night (Chardonnet et pp. 23, 24; IUCN 2006b. pp. 18–19; use of Furadan, a widely available and al. 2010, p. 35). Frank et al. 2006, p. 3). The availability cheap agricultural pesticide, is of guns and poison makes killing Attacks on Humans particularly lethal to wildlife and is suspected predators cheaper and easier increasingly being used to kill predators Provoked attacks on humans are than other control methods, such as in small pastoralist areas of Kenya and usually associated with someone reinforcing bomas (Hazzah et al. 2009, Tanzania. Livestock carcasses are approaching a lion too closely or trying p. 2,429; Moghari 2009, p. 35; Frank et doused with the poison, killing to injure or kill it and stealing a lion’s al. 2006, p. 14; Hazzah 2006, p. 3). predators and scavengers that feed on prey for bushmeat (Chardonnet et al. Spearing, shooting, trapping, and them (Frank et al. 2006, pp. 2, 10, Living 2010, p. 14; Uganda Wildlife Authority poisoning of lions, as either a preventive with Lions no date, unpaginated). 2010, p. 27). Unprovoked attacks are measure or in retaliation for livestock Poisoning of bush pig carcasses to kill usually associated with old, sick, or and human attacks, occurs regularly lions is not uncommon after attacks on injured lions that turn to humans as (Government of Namibia 2013, pp. 12, humans. These practices have serious easy prey. Additionally, there are risks 13–14; Begg and Begg 2010, p. 15; negative impacts on lion populations of unprovoked attacks associated with Chardonnet et al. 2010, pp. 41–42; (Frank et al. 2006, p. 9). certain human activities. These Packer et al. 2010, pp. 9–10; Uganda activities include walking alone at Wildlife Authority 2010, pp. 13, 42; Factors That Drive Retaliation night, sleeping outside, and surprising a Gebrensenbet et al. 2009, p. 7; Hazzah Several anthropogenic factors drive lion, particularly if it has cubs (Begg and et al. 2009, p. 2,429; Moghari 2009, pp. the level of resentment towards lions Begg 2010, pp. 3, 21; Chardonnet et al. 52, 89, 91; Ikanda 2008, pp. 5–6; Hazzah and the extent of retaliatory killing 2010, pp. 14, 15; Mesochina et al. and Dolrenry 2007, p. 21; Frank et al. (Dickman 2013, pp. 379, 385), including 2010a, pp. 38, 39; Mesochina et al. 2006, pp. 2–4, 7; Hazzah 2006, p. 52; the extent of the loss caused by the 2010b, p. 32; Uganda Wildlife Authority IUCN 2006b, p. 15). Studies have shown lions, and the wealth and security of the 2010, p. 27; Moghari 2009, p. 85; Frank that lion populations are declining in people affected (Dickman 2013, p. 381; et al. 2006, pp. 11, 12). Inebriated areas where pastoralism persists Mesochina et al. 2010b, p. 54; Moghari people may walk in an altered manner (Hazzah et al. 2009, p. 2,428). Within 2009, pp. 14, 25; Hazzah 2006, p. 81). that resembles sick or injured prey, protected areas, human-wildlife conflict Depending on alternative assets or attracting the attention of lions (Moghari is likely under-reported because cattle incomes, the economic impact of lions 2009, p. 85). The most common context herders are within the protected areas killing livestock can be significant. for attacks on humans occurs during illegally and, therefore, unlikely to Domestic livestock can provide manure, harvest, due to prey dispersal during the report it (Chardonnet et al. 2010, p. 14; milk, and meat, and are the basis of wet season, bush pig attraction to crops, Mesochina et al. 2010b, p. 34). For many family incomes, savings, and and because humans are particularly example, Etosha National Park and social standing; losses can amount to a vulnerable in makeshift tents while Caprivi Game Park have the highest large proportion of a subsistence

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herder’s annual income. These losses Africa, locals believe in ‘‘spirit lions’’, a sustainable. Attacks on humans can are generally uncompensated, lion whose body is overtaken by evil to impact long-term viability for lions as reinforcing negative community kill rivals or their livestock (West 2001 people who fear for their lives or safety attitudes toward lions and causing in Dickman 2013, pp. 381–382). Because are unlikely to support conservation retaliation (Dickman 2013, pp. 380, 381; people believe spirit lions are created by actions and are more likely to retaliate Chardonnet et al. 2010, pp. 11, 12, 18, their enemies, the number of perceived by killing any lions found near 29; Hazzah et al. 2009, p. 2,428; spirit lions, and killing of these lions, settlements (Frank et al. 2006, p. 12). Moghari 2009, pp. 14, 25, 27, 36; Kissui increases during times of social tension Every year, human-lion conflicts 2008, pp. 422–423). Furthermore, a (Dickman 2013, p. 382. The prohibition intensify due to habitat loss, poor common perception among local of ritual lion hunts provides a greater livestock management, and decreased communities is that lions are conserved incentive for participating in retaliatory availability of wild prey, further at the cost of community safety and hunts (Packer et al. 2010, p. 10; Moghari increasing the likelihood that the uncompensated financial losses. When 2009, pp. 13–14, 28; Ikanda 2008, pp. 5, subspecies will be at risk of extinction the people who suffer significant costs 6; Kissui 2008, p. 423; Frank et al. 2006, within the foreseeable future (Lion from wildlife feel that the wildlife’s p. 10; Hazzah 2006, p. 99). Guardians 2013, p. 1). needs are being put before their own Social tensions within tribes and Human population growth within the needs, their frustration can lead to between local communities and other lion’s range is projected to be 2.1 billion retaliatory killings (Dickman 2013, p. communities, the government, park by 2050 (UN 2012, p. 2). The number of 382). This situation further contributes officials, or tourists can lead to conflict livestock within the lion’s range is to negative attitudes toward lion and retributive killing of lions (Dickman projected to increase by about 73 conservation programs (Moghari 2009, 2013, p. 382; Hazzah 2006, p. 75). percent by 2050 (Food and Agriculture p. 37). Locals often report that wildlife Organization of the United Nations Lions are particularly vulnerable to authorities do not react effectively when 2012, p. 133). Given this expected retributive killing because they are often chronic livestock raiders are reported increase in humans and livestock by driven by a perceived level of lion (Frank et al. 2006, p. 9). Significant 2050, we conclude the conditions predation on livestock rather than actual numbers of lions have been killed when described above will continue to worsen levels of conflict. In some locations, promised benefits were not received or to the point that African lions will likely other predators (e.g., baboons (Papio adequate compensation was not be at risk of extinction within the ursinus), spotted hyenas (Crocuta provided for livestock and human losses foreseeable future. As livestock numbers crocuta), and leopards (Panthera (Dickman 2013, p. 383; Hazzah 2006, p. increase, expansion of agricultural and pardus)) as well as disease are 45). pastoral practices continue, and the responsible for the majority of livestock Summary of Human-Lion Conflict lion’s prey base is hunted at losses and human casualties, yet it is unsustainable levels to meet a growing lions that are sought and killed more Human-lion conflict and associated demand for food, livestock depredation often. Negative perceptions of lions may retaliatory killing of lions has played a and retributive killing of lions will be based on an over-estimated number major role in the reduction of lion likely increase (Dickman 2013, p. 379; of lions in a community or protected populations and is the greatest threat to Hoppe-Dominik et al. 2011, p. 452; area and an over-estimated number of remaining lion populations. The most Chardonnet et al. 2010, p. 19; human-lion conflicts (Dickman 2013, p. significant cause of human-lion conflict 380; Begg and Begg 2010, p. 20; is livestock depredation and, to a lesser Gebresenbet et al. 2009, p. 9; Hazzah Chardonnet et al. 2010, pp. 12, 21–22; extent, attacks on humans. Expansion of and Dolrenry 2007, p. 3). Furthermore, Hazzah et al. 2009, p. 2,436; Maclennan human settlements and agricultural and as the need for grazing land becomes et al. 2009 in Hazzah et al. 2009, p. pastoral activities into lion habitat, and more critical, expansion of livestock 2,429; Moghari 2009, pp. 77–78, 107, even into protected areas, decreases numbers may be partially supported by 150; Holmern et al. 2007 in Moghari prey availability and increases exposure the network of protected areas, seen by 2009, p. 34; Butler 2001 in Moghari of livestock and humans to lions. herders as unused pastures (Chardonnet 2009, p. 34; Kissui 2008, pp. 426, 428, The most common solution to lion et al. 2010, p. 25). 429; Hazzah 2006, pp. 18–19, 83–85, 96, attacks is retaliatory killing. Spearing, Retaliatory killing of lions continue in 98, 107, 111; Patterson et al. 2004, pp. shooting, trapping, and poisoning of many areas and this practice impacts 514, 515). One cause for the lions occur regularly. Although a the viability of lion populations disproportionate blame put on lions is majority of information on human-lion throughout its range. The killing of lions that the lion is a highly visible species. conflict comes from a few areas of the due to human-lion conflict is enough to It is a large-bodied species that lives in lion’s range, we can reasonably result in the local extirpation of lion groups and has cultural significance. conclude that lions are being killed due populations, though at present does not Because of its physical presence, there to conflict in all major range countries, place the subspecies in danger of is often a ‘‘hyper-awareness’’ of the because of their depredation on extinction. Human-lion conflict is potential risk for lion attacks and lions livestock (Frank et al. 2006, p. 4). exacerbated by an increasing human may be blamed simply because they Impacts on victims of lion attacks population, the expansion of human have been seen in an area (Dickman create resentment towards lions and settlements, loss of prey base due to the 2013, pp. 380–381). lion conservation, and a greater bushmeat trade and expanding Cultural beliefs and traditions can likelihood of retaliation. Even when agriculture, as well as increasing have a negative impact on lions. lions are not the predators responsible pressures on natural resources to meet Because cattle are of great cultural for the majority of attacks, lions incite the needs of the growing human significance to Maasai, their loss can a greater response and are killed more population. We expect retaliatory impose social or cultural costs and often than other predators of livestock. killings due to human-lion conflict to incite greater resentment and higher In areas of high human density and continue to increase into the foreseeable levels of retributive killing (Dickman low lion density, mainly in smaller future. We conclude based on the best 2013, p. 384; Kissui 2008, p. 429; reserves and outside large protected available scientific and commercial Hazzah 2006, p. 99). In some areas of areas, lion populations may not be information that the continuation of this

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activity is a significant threat to the Cleaveland et al. 2005, p. 450). Because they experience another bTB infection, African lion throughout its range. the lion’s primary prey are infected with suffer from poor nutrition or advancing bTB, they are frequently exposed to age, or become super-infected with Disease large amounts of infected tissue and are other diseases that may exacerbate the Wild lions are known to be infected at risk of infection (Keet et al. 2009, pp. infection (Renwick et al. 2007, p. 533). with various pathogens (Hunter et al. 4, 6; Renwick et al. 2007, pp. 532, 536; The impact of bTB on lions is largely 2012, p. 2; Craft 2008, p. 6; Michel et al. Michel et al. 2006, p. 93; Cleaveland et unknown. Researchers suggest that bTB 2006, p. 92; Hofmann-Lehmann et al. al. 2005, pp. 450, 451). Furthermore, may lower breeding success, reduce 1996, pp. 559–561). The human predators prey on weak animals and resiliency, and may be a mortality factor population within the range of the lion scavenge on carcasses, increasing their based on data that indicate survival is is expanding into lion habitat, likelihood of being exposed to M. bovis shortened in infected lions, with death increasing the exposure of lions to (Renwick et al. 2007, p. 536; Michel et ranging between 2 and 5 years after diseases from domestic animals (IUCN al. 2006, p. 93). Transmission may also infection (Maas et al. 2012, p. 4,212; 2006b, p. 26). Because lions are a top occur among lions via scratching and Renwick et al. 2007, p. 536; Michel et predator, they are at a particularly high biting (Keet et al. 2009, p. 7; Renwick al. 2006, p. 93; Cleaveland et al. 2005, risk of exposure to pathogens (Keet et al. et al. 2007, pp. 532–533). M. bovis is a pp. 450, 451). Thirty percent of the 2009, p. 11). Some pathogens are pathogen that causes the infected inbred populations in Hluhluwe- endemic, meaning they are constantly animal to remain infectious and, iMfolozi Park died due to a combination present, but often do not cause disease. therefore, a source of infection, until it of bTB and malnutrition (Hunter et al. Others are epidemic and cause a sudden dies (Renwick et al. 2007, p. 531). 2012, p. 3). A study from Kruger severe outbreak with the potential to The social behavior of buffalo and National Park indicated that bTB cause high mortality (Craft 2008, pp. 5, lions allows M. bovis to spread to larger spreads quickly through lion 6). Although lions are known to be areas and facilitates the transmission populations; in an area with high herd infected with certain pathogens, within and between prides. Drought prevalence of M. bovis, 90 percent of information on the extent of the conditions may also encourage the lions became infected (Cleaveland et al. subspecies’ infections and impacts of spread of this pathogen as herds must 2005, p. 451). However, despite bTB these diseases on lion populations is move into new areas in search of forage, infection and a high prevalence in prey limited, because few long-term studies potentially putting them in contact with species, the lion population in Kruger have been conducted; for example, new, uninfected herds (Keet et al. 2009, National Park has remained stable those lion populations found in pp. 4, 6; Renwick et al. 2007, p. 533; (Ferreira and Funston 2010, p. 201). Serengeti National Park, Ngorongoro Michel et al. 2006, p. 93). In Kruger Epidemics of canine distemper virus Crater, and Kruger National Park. National Park, bTB was introduced in (CDV) are known to have occurred in Feline calicivirus, feline herpesvirus, the southeastern corner of the park the Serengeti-Mara Ecosystem, an area feline parvovirus, feline coronavirus, between 1950 and 1960. It gradually that encompasses the Serengeti National and feline leukemia virus are endemic made a northern progress and reached Park, Ngorongoro Conservation Area, viruses known to occur in lions of the park’s northern boundary in 2006. In and Maasai Mara National Reserve Serengeti National Park, Ngorongoro 2009, the disease was found in buffalo (Craft 2008, pp. 13–14; Cleaveland et al. Crater, Lake Manyara National Park, across the river boundary in Zimbabwe 2007, pp. 613, 616, 618). CDV is a Kruger National Park, and Etosha (Keet et al. 2009, pp. 6, 11; Renwick et common pathogen in the large National Park (but not all viruses are al. 2007, pp. 532, 533; Michel et al. population of domestic dogs around the known in all parks). However, these 2006, pp. 92, 96, 98). In time it will Serengeti-Mara Ecosystem, which are diseases are not known to affect lion likely spread to Mozambique (Keet et al. believed to be the source of CDV survival (Hunter et al. 2012, p. 2; Craft 2009, p. 6). In Serengeti National Park, (Cleaveland et al. 2007, pp. 613, 617). 2008, p. 6; Hofmann-Lehmann 1996, pp. infection may be widespread due to the CDV is assumed to be transferred to 559, 561). large, migratory wildebeest population lions by the sharing of food sources with Lions within Kruger National Park that ranges throughout the Serengeti spotted hyenas (Crocuta crocuta) or and Hluhluwe-iMfolozi Park, South ecosystem, including Maasai Mara jackals (Canis spp.) that become Africa, and Serengeti National Park, National Reserve (Cleaveland et al. infected by consuming the infected Tanzania, are known to be infected with 2005, p. 450). Although an eradication carcasses of domestic dogs (Canis lupus Mycobacterium bovis, a pathogen that program has been implemented for familiaris). Lions may also transmit causes bovine tuberculosis (bTB). This cattle in South Africa, once an infection CDV among themselves via sharing pathogen is not endemic to African is established in a free-ranging food, fights, and mating (Craft et al. wildlife and was likely introduced from maintenance host, like buffalo, it is 2009, pp. 1,778, 1,783; Craft 2008, pp. cattle imported from Europe. M. bovis is unlikely to be eradicated (Keet et al. 13, 18, 71). transmitted to ungulates, such as 2009, p. 11; Renwick et al. 2007, pp. CDV generally lacks clinical signs or African buffalo (Syncerus caffer) and 537, 538; Michel et al. 2006, p. 96). In measurable mortality in lions, and most wildebeest (Connochaetes taurinus) fact, modeling has predicted that CDV events have been harmless. from domestic cattle located on the prevalence could reach as high as 90 However, in 1994 and 2001, CDV periphery of the parks (Maas et al. 2012, percent over the next 25 years, with epidemics in the Serengeti National p. 4,206; Keet et al. 2009, pp. 4, 11; similar consequences for predators Park/Maasai Mara National Reserve and Renwick et al. 2007, p. 532; Michel et (Renwick et al. 2007, p. 535). Ngorongoro Crater, respectively, al. 2006, pp. 92, 93; Cleaveland et al. Clinical signs of bTB in lions include: resulted in unusually high mortality 2005, pp. 446, 449, 450). Spillover of the emaciation, respiratory complications, rates (Hunter et al. 2012, p. 2; Craft disease from buffalo to other lion prey swollen lymph nodes, draining sinuses, 2008, p. 14; Munson et al. 2008, pp. 1, species, such as kudu (Tragelaphus ataxia, and lameness (Keet et al. 2009, 2; Cleaveland et al. 2007, pp. 613, 618; strepsiceros) and warthog p. 13; Renwick et al. 2007, pp. 533, 534; Roelke-Parker et al. 1996, pp. 441, 443). (Phacochoerus africanus), have also Cleaveland et al. 2005, p. 450), although These outbreaks coincided with climate been documented (Keet et al. 2009, pp. some lions may be subclinically extremes that resulted in a higher 4, 11; Renwick et al. 2007, p. 535; infected but remain asymptomatic until number of Babesia, a tick-borne

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parasite, infections (Munson et al. 2008, Slattery et al. 2008, p. 4; O’Brien et al. impacts on lion health, but climatic pp. 2, 5). Babesia is common in lions, 2006, p. 262). The social nature of lions conditions increased exposure of lions but typically at low levels with no allows for viral transmission within and to Babesia infections, which were measurable impacts on their health between prides through saliva when exacerbated by the immunosuppressive (Craft 2008, p. 14; Munson et al. 2008, biting (Maas et al. 2012, p. 4210; Pecon- effects of CDV and led to unusually high p. 3). However, droughts in 1993 and Slattery et al. 2008, p. 5; Brown et al. mortality rates. Furthermore, species 2000 in Serengeti National Park/Maasai 1994, p. 5,953). Prevalence of FIV in with reduced genetic variation may be Mara National Reserve and Ngorongoro infected lion populations is high, often less able to mount an effective immune Crater, respectively, led to large-scale approaching 100 percent of adults response against an emerging pathogen starvation and widespread die-offs of (O’Brien et al. 2012, p. 243; Troyer et al. (O’Brien et al. 2006, p. 255). Some lions buffalo. This situation combined with 2011, p. 2; Roelke et al. 2009, p. 3; infected with bTB may remain resumption of rains and fire suppression O’Brien et al. 2006, p. 262; Hofmann- asymptomatic until conditions change in Ngorongoro Crater favored Lehmann et al. 1996, p. 559). and they suffer from poor nutrition due propagation of ticks, vectors of Babesia, FIV causes immune deficiencies that to low prey density, advancing age, or leading to unusually high tick burdens. allow for opportunistic infections in the become super-infected with other The compromised health of buffalo host (Brown et al. 1994, p. 5,953). diseases that may exacerbate the allowed lions to feed on an inordinate Chronic effects of FIV are important to infection (Renwick et al. 2007, p. 533). number of tick-infested prey (Craft 2008, long-term survival and differ according Impacts of coinfections of FIV with p. 14; Munson et al. 2008, pp. 2, 4, 5). to subtype (Troyer et al. 2011, p. 6). FCV, FPV, FHV, and FCoV on Exposure to either CDV or Babesia Studies have indicated that lions may individual lions are negligible and do singly is not typically associated with a exhibit signs of opportunistic infection not endanger the lion population, at compromise in health or an increase in associated with AIDS, such as swollen least in the absence of other aggravating mortality (Craft 2008, p. 14; Munson et lymph nodes, gingivitis, tongue cofactors (Hofmann-Lehmann et al. al. 2008, pp. 1, 2, 3). However, the papillomas, dehydration, poor coat 1996, p. 561). Pathogen–pathogen Babesia infections were exacerbated by condition, and abnormal red blood cell interactions may become more the immunosuppressive effects of CDV parameters, and in some cases death important when lions are under and led to the unusually high mortality (Troyer et al. 2011, p. 2; Roelke et al. additional stress (e.g., increased parasite rates (Craft 2008, p. 14; Munson et al. 2009, pp. 2, 3–6). Lions in Botswana load or low prey density) (Maas et al. 2008, p. 5). The Serengeti National Park/ and Tanzania have demonstrated 2012, p. 4,212). Maasai Mara National Reserve lion multiple clinical features of chronic Although disease is known in several population lost 30 percent of its immune depletion similar to HIV and populations, the impacts are known in population (approximately 1,000 lions), domestic cat AIDS (Troyer et al. 2011, only a couple of populations where but has recovered to its pre-epidemic pp. 2–3). However, there is no evidence disease has been frequently studied. population levels (Craft 2008, pp. v, 14, that it poses a threat to wild populations Disease can be a factor in the decline of 41; Munson et al. 2008, p. 1; Cleaveland (Frank et al. 2006, p. 1); FIV does not lions when combined with other factors, et al. 2007, pp. 613, 617; Roelke-Parker appear to be impacting lions in Kruger including environmental changes, et al. 1996, p. 444). Thirty-four percent National Park (Maas et al. 2012, p. reduced prey density, and inbreeding of the Ngorongoro Crater lion 4,212), and no evidence of AIDS-like depression. However, this type of population was killed, but frequent illnesses or decreased lifespan has been impact has been observed in some small outbreaks of disease have prevented this found in FIV lion populations in the populations that are at a higher risk, but population from recovering back to its Serengeti (O’Brien et al. 2006, p. 263). has not been observed at the species carrying capacity (Craft 2008, p. 14; Infection with a single disease does population level. Therefore, we Munson et al. 2008, pp. 1, 2; Cleaveland not appear to have detrimental impacts conclude, based on the best scientific et al. 2007, p. 617). The difference in on lions, although general body and commercial information available, recovery is likely due to the highly condition, health, and lifespan may be that disease is not a significant threat to inbred nature of the Ngorongoro Crater compromised. Co-infections, however, the species. lion population, compared to the could have synergistic effects that lead Deleterious Effects Due to Small Serengeti population, and its greater to greater impacts on lions than a single Population Sizes susceptibility to parasitic and viral infection. Lions impacted by the 1994 infections (Hunter et al. 2012, p. 2; CDV outbreak in Serengeti National The risk of extinction is related to the Munson et al. 2008, p. 5; Brown et al. Park/Maasai Mara National Reserve may moment when a declining population 1994, pp. 5,953–5,954). have been more susceptible to CDV due becomes a small population and is often Feline immunodeficiency virus (FIV) to depleted immunity caused by FIV estimated using minimum viable is an endemic pathogen in many lion (O’Brien et al. 2006, p. 263). Troyer et population (MVP) sizes (Traill et al. populations of southern and eastern al. (2011, pp. 5–6) found that survival 2010, p. 28). The viability of a lion Africa (Maas et al. 2012, p. 4,206; during the CDV/Babesia outbreak in population is complex, but it partly Adams et al. 2011, p. 173; Pecon- Serengeti National Park/Maasai Mara depends on the number of prides and Slattery et al. 2008, p. 2; Hofmann- National Reserve was significantly less ability of males to disperse and interact Lehmann et al. 1996, pp. 555, 558; for lions infected with FIV A and/or C with other prides, which affects Brown et al. 1994, p. 5,966). FIV is than FIV B. This finding suggests that exchange of genetic material (Bjorklund believed to have been present in lions FIV A and C may predispose carriers to 2003, p. 518). Without genetic exchange, since the late Pliocene (O’Brien et al. CDV pathogenesis and may increase the or variation, individual fitness is 2012, p. 243; Troyer et al. 2011, p. 2; risk of mortality (O’Brien et al. 2012, p. reduced and species are less able to Roelke et al. 2009, p. 3; Pecon-Slattery 243). Additionally, certain adapt to environmental changes and et al. 2008, p. 8). There are 6 subtypes environmental conditions may stress, increasing the risk of extinction of FIV, A through F, each with a distinct exacerbate the effects of an otherwise (Bijlsma and Loeschcke 2012, pp. 117, geographic area of endemnicity (Adams innocuous infection. For example, as 119; Segelbacher et al. 2010, p. 2; Traill et al. 2011, p. 174; Troyer et al. 2011, discussed above, CDV and Babesia et al. 2010, p. 31; Bjorklund 2003, p. p. 2; Roelke et al. 2009, p. 3; Pecon- infections generally have no measurable 515).

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Some scientists believe that the the human population grows and lion hunting quota, the country has halted minimum viable population size (MVP) populations decline, as discussed above, authorization of trophy hunting to maintain genetic viability is between more lion populations could reach pursuant to that quota until some later 500 and 5,000 individuals, although this levels below the suggested minimum of date or until some further action is estimate is not specific to lion (Bijlsma 10 prides to maintain genetic diversity, taken, as prescribed by that country. and Loeschcke 2012, p. 122; Traill et al. putting more populations at risk of Therefore, you will see us refer to 2010, p. 30; Willi et al. 2006, p. 449). inbreeding and extirpation. Therefore, countries like Zambia and Botswana, The MVP for the African lion has not we conclude, based on the best each of which has hunting quotas and been formally established and agreed scientific and commercial information bans in place. Trophy hunting is upon by species experts (Riggio et al. available, that small population sizes currently banned in 12 countries: 2011, p. 5; CITES 2004, p. 2; Bjorkland currently pose a threat to the species. Angola, Botswana, Cameroon,7 Congo, Gabon, Ghana, Kenya, Malawi, 2003, p. 521); however, it has been Trophy Hunting suggested that, to conserve genetic Mauritania, Niger, Nigeria, and Rwanda diversity populations of 50 to 100 prides Trophy hunting (also known as sport (CITES 2014a, p.14; Lindsey et al. (250 to 500 individuals), with no limits hunting) has been identified by the 2013a, entire; Lindsey 2013, pers. to dispersal, are necessary because petitioners as one of the factors comm.; Jackson 2013, pp. 7–8). inbreeding increases significantly when contributing to the decline of African Botswana banned lion hunting between populations fall below 10 prides. If lions (Petition 2011, p. 24). Lions are a 2001 and 2004, and then again from there are less than 10 prides, inbreeding key species in sport hunting as they are 2008 to the present (Davison et al. 2011, will increase from an F-value of 0.0 in considered one of the ‘‘big five’’ (lion, p. 114). Kenya banned all sport hunting the initial state to an F-value 0.26–0.45 leopard, elephant, rhino, and cape in 1977 (African Wildlife Foundation after 30 generations, while if the number buffalo), touted to be the most 1998, p. 3). Trophy hunting is restricted of prides is 100 this F-value is only challenging species to hunt, due to their to problem or dangerous animals in around 0.05 assuming no migration into nimbleness, speed, and behavioral Ethiopia and Uganda (Lindsey 2008, p. the population (Bjorkland 2003, p. 515). unpredictability (Lindsey et al. 2012a, 42). Zambia banned all sport hunting in F is the probability that the two alleles p. 2). However, with the documented January of 2013; while restrictions were of a gene in an individual are identical decline in lion population numbers lifted from other trophy species in by descent. Therefore, the Service throughout Africa, the sport hunting of August 2014, the ban on lions and considers the MVP to be 50 prides. lions for trophies has become a highly leopards remains in place (ABC News Because the number of prides and male complex issue that has raised 2014, unpaginated; Flocken 2013, dispersal are the most important factors considerable controversy among unpaginated). In 2011, researchers in for maintaining viability, sufficient stakeholders. Cameroon suggested that there should areas are needed to support 50 or more Range Countries be an immediate moratorium of at least 5 years on the hunting of lions in prides and allow unrestricted male As of May 2014, approximately 18 dispersal. Unfortunately, few lion Cameroon, during which lions are countries in Africa permit lions to be allowed to recover and a management populations meet these criteria, and few hunted for trophies: Benin, Burkina protected areas are large enough to plan for lion hunting is established Faso, Central African Republic (CAR), (Croes et al. 2011). support viable populations (Bauer et al. Democratic Republic of Congo (DRC), 2008, unpaginated; Riggio 2011, p. 5; Ethiopia, Ivory Coast, Mali, Quotas Hazzah 2006, p. 2; Bauer and Van Der Mozambique, Namibia, Senegal, A scientifically based ‘‘quota’’ is the Merwe 2004, pp. 28–30; Bjorklund Somalia, South Africa (RSA), Sudan, maximum number of a given species 2003, p. 521). Even within large areas, Tanzania, Togo, Uganda, Zambia, and that can be removed from a specific inbreeding will increase if dispersal is Zimbabwe. However, in 2013 lion population without damaging the limited, (Bjorklund 2003, pp. 521–522). trophy hunting was only documented to biological integrity and sustainability of More than 6,000 lions are in occur in nine countries, specifically that population (World Wildlife Fund populations where their probability of Benin, Burkina Faso, CAR, (WWF) 1997, p. 9). For a quota to be survival is likely to be at risk of Mozambique, Namibia, RSA, Tanzania, scientifically based, it must be based extinction within the foreseeable future Zambia, and Zimbabwe (Lindsey 2013, upon available monitoring data of the (Riggio et al. 2013, p. 33). Furthermore, personal communication). Four species. Although varying by country research indicates that there is a general countries, Burundi, Guinea Bissau, and by economic resources, monitoring lack of gene flow in most lion Lesotho, and Swaziland, provide no data used to determine quotas have conservation units (Dubach et al. 2013, legal protection for lions (CITES 2014a, included, but are not limited to, past pp. 749, 750; Bertola et al. 2011, p. p. 14). hunting off-take records, trophy quality 1364; Chardonnet et al. 2009, p. 54). data, ground transect surveys, wildlife Small populations (e.g. fewer than 50 Hunting Moratoriums ranger and safari operator input, the lions) can persist in the wild for some In response to growing international species’ reproductive biology, and aerial time; however, the lack of dispersal and recognition of reduced population population census data, although genetic variation can negatively impact numbers, many countries began usually aerial data is limited to species the reproductive fitness of lions in these implementing moratoriums banning the that can be easily observed from the air, populations and local extirpation is sport hunting of lions. In this document such as elephants and buffalo (Barnett & likely (Traill et al. 2010, p. 30; O’Brien we use the terms moratorium and ban Patterson 2005, p. 102). Generally, the 1994, p. 5,748). interchangeably. A ban or moratorium conservation principle behind Increasing human population growth can be permanent, long term, or scientifically based quotas is to limit between now and 2050 will continue to temporary, and can occur in countries decrease and fragment large areas of that have hunting quotas in place. 7 We found conflicting data on Cameroon, which habitat needed to support viable lion Having both a moratorium and a quota was reported to prohibit trophy hunting (CITES 2014, p. 14), although other information provided populations and disrupt dispersal routes in place at the same time means that, by Lindsey (2013, pers. comm.) and Jackson (2013, for genetic exchange. Additionally, as although the country may have a p. 8) state that trophy hunting is legal in Cameroon.

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offtake of the species to either equal or (Lindsey et al. 2013a, p. 9; Packer et al. According to Packer et al. (2006, pp. 2– slightly lower than the growth rate of 2006, pp. 5, 9). 3), regardless of population estimates, the target specimens (e.g., males vs. Two primary concerns have been countries are allowing for only a small female), provided the offtake does not raised by the scientific and international proportion of their lion populations to damage the integrity and sustainability community with regards to current lion be hunted, with most countries ranging of that population. quotas. Specifically, that existing quotas from 2–4 percent annually (excluding In order for scientifically based quotas are set above sustainable levels and the offtake from South Africa, where offtake to result in offtake less than the growth data used for setting quotas is has been increasing from the trophy rate of target specimens, many factors inconsistent and not scientifically based hunting of primarily captive-born lions, are evaluated including the species’ (Hunter et al. 2013, unpaginated; and Zimbabwe, where offtake was 2–3 biological factors (reproductive rate, Lindsey et al. 2006, p. 284). For percent higher than other countries from gender, age, and behavior), as well as example, recent quotas appear rarely to 1998–2004. community and client objectives (WWF address safeguards for sustainability or Regardless of these reductions, many 1997, pp. 14–19). Each quota should be establish a systematic approach to stakeholders consider the quota system then assigned to a geographical area setting lion quotas (Hunter et al. 2013, to be outdated and ineffective because it and/or population based on this p. 2; Lindsey et al. 2013b, p. 8). does not address the biological and information. Thus, for lions, a Additionally, it has been noted that social impacts of trophy hunting on lion scientifically based quota defines the previous quotas in Namibia, prides. Opponents also state that trophy specific number of lions that can be Mozambique, and Zimbabwe may have hunting affects the social structure of removed from a specific geographical been influenced by human-lion conflict, the pride and results in increased area and population, for any purpose, with higher quotas being allocated to infanticide of lion cubs. This within a particular year. Scientifically locations with reportedly higher human- supposition is inconclusive and not based quotas do not apply solely to lion conflict levels (Lindsey et al. well supported (CITES 2014a, p. 14; sport hunting, but set the limits for all 2013b, p. 4). Apparently, in recognition Dagg 2000, pp. 831–835) (See offtake for a particular year; other of these inconsistencies, range countries Infanticide and Age-based Hunting potential offtake includes problem- and conservationists have been working Strategies). Regardless, since 2006, animal control (to reduce human- to establish a set of best practices in researchers have recommended the wildlife conflict), translocation (to order to create a more consistent, implementation of age-based hunting expand conservation), culling (reducing scientifically based approach to strategies; these are discussed below population pressures), and local hunting determining quotas. The recommended (Packer et al. 2006, pp. 6–8). (for protein/meat or employment) (WWF best practices include: (1) Establishing Five countries maintain quotas to 1997, pp. 8–10). processes and procedures that are allow for approximately 6–15 lion While each of these uses offers clearly outlined, transparent, and trophies to be taken per year: Benin, advantages and disadvantages, quotas accountable; (2) establishing processes Burkina Faso, Cameroon,7 Mozambique, are typically utilized only for sport and procedures that are CITES and Namibia. Tanzania allows the take hunting, as it may provide the highest compliant;(3) demonstrating of approximately 50 lions annually, and all-around benefits to local management capacity; (4) standardizing Zimbabwe allows approximately 70 communities. For example, a portion of information sources; (5) establishing animals annually to be taken (Jackson a quota could be used to kill a problem monitoring systems for critical data; (6) 2013 pp. 7–8, CITES WCMC–UNEP animal; the benefits to the community recording and analyzing trophy hunting trade database, accessed December would then include the use of the data; (7) conducting data collection and 2013). In Ethiopia and Uganda, trophy animal parts for meat or trade and it analysis for each hunting block and hunting is restricted to problem or would theoretically reduce the conflict. concession; and (8) establishing a dangerous animals only (Lindsey 2008, However, this provides a more limited primary body who will approve quotas p. 42), and Botswana and Zambia economic benefit to the community than (Burnett and Patterson 2005, p. 103). We currently ban all trophy hunting (CITES would selling the same quota for trophy have no information on whether these 2014a, p.14). South Africa has not set a hunting, which could potentially best practices have been implemented quota for the take of wild lions since 99 eliminate the problem animal, provide by the lion range states. However, most percent of the trophy-hunted lions are meat and parts for trade, and provide countries that allow trophy hunting of reportedly not of wild origin, but revenue for the community (WWF 1997, lions appear to be reviewing their captive-born (Hunter et al. 2013, p. 2; pp. 31–33). trophy hunting practices (Jackson 2013, There are two primary types of RSA 2013, pp. 5, 7). pp. 2–3; White 2013, pp. 12–13). Benin Below is a summary of estimated quotas, ‘‘fixed’’ and ‘‘optional.’’ Trophy halved their quotas in 2002 after the annual hunting quotas for the African fees for ‘‘optional’’ quotas are paid only first population census of lions was lion: when the lion is shot, whereas, ‘‘fixed’’ conducted and resulted in the current quotas require the payment of a portion quota of six lions every 2 years in TABLE 7—ANNUAL TROPHY QUOTAS (40–100 percent) of the lion trophy fee, Pendjari and four lions every 2 years in (APPROXIMATE) AS OF 2013 regardless of whether the hunt is western Benin or one lion annually in successful. Until 1999, male lions were each of the five hunting zones. This was Annual lion typically on ‘‘fixed’’ quotas, whereas largely due to impacts to lions from trophy quotas female lions were under ‘‘optional’’ habitat degradation and fragmentation Country (Jackson 2013, quotas. Due to this approach, trophies (particularly exacerbated by the increase pp. 7–8) collected in the 1990’s were often of of human population), loss of prey by lower quality, younger, less desirable Benin ...... 6 poaching, trade (both legal and illegal), Botswana (moratorium) ...... 30 male lions, as operators and hunters had and human-lion conflict. (CITES 2014a, Burkina Faso ...... 6 no incentive to be selective (e.g. the p. 5; Sogbohossou 2014, p. 1). Cameroon 7 ...... 6 hunter had already paid for it). Throughout the countries in Africa, Mozambique ...... 15 Therefore, current recommendation for most appear to have reduced their Namibia ...... 10 all quotas is to be the ‘‘optional’’ type offtake considerably since the 1990’s. Tanzania (as of 2012) ...... 50

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TABLE 7—ANNUAL TROPHY QUOTAS using the CITES Trade Database have been hunted over the past two (APPROXIMATE) AS OF 2013—Con- indicates that the outputs produced by decades (IUCN 2009, pp. 36–37; Bauer tinued the CITES Trade Database can be easily and Nowell 2004, p. 36), although their misinterpreted if one is not familiar current annual quota is 6 animals. In Annual lion with it (CITES 2004b, p. 5). The number Botswana, a quota of 30 lions per year Country trophy quotas of ‘‘trophies’’ reported does not was authorized for nearly two decades; (Jackson 2013, necessarily equate to the number of however, Botswana has recently pp. 7–8) lions hunted. Additionally, the number implemented a hunting moratorium Zambia (moratorium) ...... 50 of trophies reported for a given year in (Jackson 2013, p. 8). (CITES lion gross Zimbabwe ...... 70 the trade report does not equate directly exports, http://trade.cites.org, accessed to the number of animals hunted in that April 23, 2014; CITES UNEP–WCMC Import/Export of Lion Trophies given year (CITES export permits may database, accessed January 8, 2014, and be valid for 6 months, and a trophy August 16, 2013). Although each country has its own could in theory be exported the year method of regulating trophy hunting, after it was hunted). The second Potential Impacts of Trophy Hunting international trade of lion trophies must limitation to interpreting this Infanticide and Age-Based Hunting adhere to CITES (see Conservation information is, although many permits Strategies Status). International trade of lion parts may indicate that an animal is of wild and products (including trophies) are Tourist safari hunting of males has origin (source code ‘‘W’’), these permits been suggested by the petitioners to reported by both the exporting and may be incorrectly coded. This is true importing countries and tracked by the increase infanticide rates (when males for South Africa, where during the kill young lion cubs sired by other United Nations Environment period of 2000 to 2009, animals that males) (Petition 2011, p. 24; Whitman et Programme World Conservation were captive-born and released into al. 2004, p. 175), due in part to trophy Monitoring Centre (UNEP–WCMC). The private reserve systems were assigned hunters taking males under a certain international trade data on the African an incorrect source code of ‘‘wild.’’ age. Removing a younger male lion is lion that has been compiled in the South Africa has since requested their purported to allow another male to take CITES UNEP–WCMC Trade Database is provincial authorities to use the correct over the pride, and kill the former extensive. Therefore, it is likely that the source code for ‘‘captive bred’’ in order patriarch’s cubs. This supposition is actual numbers of African lion parts and to correctly reflect the source of sport- inconclusive and not well supported products in international trade is hunted lion trophies; however, some (CITES 2014a, p. 14; Dagg 2000, pp. slightly smaller than what we have provinces are still not complying (RSA reported using the UNEP–WCMC ‘‘gross 2013, pp. 8–9). However, based on 831–835). Infanticide is a common exports’’ report (CITES lion gross South African trade data, the bulk of the practice among many species, including exports, http://trade.cites.org, accessed exports of lions and their parts and lions (Hausfater et al. 1984, pp. 31, 145, April 23, 2014). products (including trophies) from 173, 487). When an adult male lion in In 2012, the most recent year for South Africa were from captive-born a pride is killed, surviving males who which CITES trade data are available, lions (RSA 2013, p. 7). form the pride’s coalition become U.S. CITES Annual Report trade data Tanzania, with the highest lion vulnerable to takeover by other male indicated that the United States allowed populations (Hamunyela et al. 2013, pp. coalitions, and this often results in the direct import of African lion 29, 283; Riggio et al. 2013, p. 32; Ikanda injury or death of the defeated males trophies from eight African countries, as 2008, p. 4; Baldus 2004, pp. 5, 6), was (Davidson et al. 2011, p. 115). In some follows: the largest exporter of wild-origin lion cases, replacement males who take over trophies, but their exports have the pride will kill all cubs less than 9 Central African Republic = 1 trophy months of age in the pride (Whitman et Ethiopia = 1 trophy decreased significantly since 2006. In Mozambique = 5 trophies 2008, approximately 138 lions had been al. 2004, p. 175). One range country Namibia = 5 trophies estimated to be killed in Tanzania as specifically addressed this issue; the South Africa = 413 trophies (the majority of trophies. In 2010, Tanzania’s numbers Republic of Namibia indicates that lion which are reported to be of captive-born declined to 128 exports, 55 in 2011, and populations reproduce at similar rates origin) 42 in 2012 (CITES lion gross exports, in both harvested and non-harvested Tanzania = 42 trophies http://trade.cites.org/, accessed April populations, but it is unclear whether Zambia = 32 trophies 25, 2014). In 2012, Tanzania established cub survival is consistent in harvested Zimbabwe = 49 trophies an annual quota to limit trophy hunting vs. non-harvested lion populations. According to the CITES UNEP– to no more than 50 animals (Jackson While utilizing individual-based WCMC database, between 2005 and 2013, p. 7). Again, it should be noted simulation models, Whitman et al. 2012, exports of lion trophies have that there may be discrepancies between (2004, pp. 175–177) found that if offtake demonstrated a decreasing trend when the annual quota and the actual number is restricted to males older than 6 years exports of captive-born lions from South of trophies exported in a given year (see of age, then trophy hunting will likely Africa are excluded (CITES lion gross http://www.cites.org/common/ have minimal impact on the pride’s exports, http://trade.cites.org, accessed resources/TradeDatabaseGuide.pdf for social structure and young (Packer et al. April 23, 2014). For example, in 2005 additional information). Regardless, the 2006, p. 6). This 6-year age restriction there were 874 lion trophy exports numbers of lion trophies exported by approach for lion trophies is in the reported in UNEP–WCMC, 521 if South Tanzania according to the UNEP– process of being self-implemented, Africa were excluded; whereas in 2012, WCMC database suggest a decreasing along with other best practices, by there were 1,237 lion trophy exports trend. professional hunting guides, and is reported in UNEP–WCMC, 336 if South In other areas within the range of the being adopted by certain range states Africa is excluded. African lion, the number of lions hunted (White 2013, p. 14; Davidson et al. 2011, Here it should be noted that there are or authorized to be hunted annually has p. 114; Whitman et al. 2004, p. 176). It limitations to interpreting the above remained fairly consistent. In Burkina involves conducting an age assessment reported information. The 2004 guide to Faso, approximately 12 lions per year of male lions using identification

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techniques, such as mane development, viable lion populations. Developing and malpractice among the hunting facial markings, nose pigmentation and implementing best management companies and various government tooth-aging, to establish the relative age practices, while not categorically departments. Some game management of male lions. Tooth wear on incisors, establishing a direct correlation with areas and privately owned game ranches yellowing and chipping of teeth, increased population numbers and were not included in the ban, but lion coupled with scars, head size, mane health, do appear to have practical hunting appears to be currently length and color, and thinning hair on impacts on lion populations. Based on prohibited throughout the country the face, as well as other factors can be the best available scientific and (Michler 2013, pp. 1–3). According to an indicator of advanced age in lions commercial information, infanticide, as some authors (Martin 2012, pp. 4, 104; (Whitman and Packer 2006, entire). a result of the removal of lions through Kimati 2012, p. 1; Kideghesho 2008, pp. Although these characteristics may be hunting, is not a threat to African lions. 16–17), corruption in the wildlife sector subjective, as regional differences may Further, it is not likely to become a has often been one of the most occur between lion populations, there threat in the foreseeable future since the discussed topics in Tanzania’s National are clear attempts by the trophy hunting science is not well supported as to Assembly, which presumably would community to establish and implement whether infanticide resulting from indicate the awareness of and best practices. Promoting the removal of offtake due to trophy hunting is a willingness to address the corrupting males 6 years of age or older, significant threat to the subspecies factors in the wildlife sector. theoretically allows younger males the (Whitman et al. 2004, pp. 175–176; Provided that countries continue to opportunity to remain resident long CITES 2014a, p. 14). address corruption within the wildlife enough to rear a cohort of cubs Corruption sector, we conclude, based on the best (allowing their genes to enter the gene scientific and commercial information Corruption is common in some areas pool; increasing the overall genetic available, that corruption, in and of within the range of the African lion, diversity). By removing males in a itself, does not currently pose a threat to manner that promotes healthy particularly in areas with extreme poverty (Michler 2013, pp. 1–3; Kimati the species. However, if efforts to population growth, the lion population address corruption do not continue, it could yield more males in the long term 2012, p. 1; Garnett et al. 2011, p. 1; IUCN 2009, p. 89; Leader-Williams et al. could become a threat to African lions (Davidson et al. 2011, p. 114; Whitman in the future. et al. 2004, p. 176). The governments of 2009, p. 296–298; Kideghesho 2008, pp. 16–17; http://www.transparency.org). Tanzania, western Zimbabwe, Revenue From Trophy Hunting Several of the range countries of African Mozambique in the Niassa National lion have experienced political The high value of lions makes them Reserve, Zambia, and most recently instability for many years, which one of the most expensive large game Benin have instituted or are in the appears to be a contributing factor in species to hunt. The revenue derived process of instituting reforms such as 6- intensifying levels of corruption. from lion hunting is substantial. Lions year age restrictions on lion trophies to Political instability results in war and are reported to generate the highest increase the likelihood that trophy famine, which essentially halt daily rate of any mammal hunted (USD hunting of lion is sustainable in those conservation efforts and the $2,650 per day), the longest number of countries (Van der Merwe 2013, p. 2; enforcement of existing wildlife days that must be booked, and the Jackson 2013, p. 3; White 2013, p. 14; protection laws (Barnett & Patterson highest trophy fee ($24,500) (Jackson Dallas Safari Club 2013, pp. 1–2; Hunter 2005, p. 82). Corruption manifests itself 2013, p. 6; Lindsey et al. 2012a, p. 5). et al. 2013, p. 2). in several ways, including According to Groom (2013, p. 4), a 21- In addition to quota-setting, embezzlement of funds and acceptance day lion hunt in Zimbabwe may be sold moratoriums, and the 6 year age limit, of bribes to overlook illegal activities or for approximately $2,500 per day, with it has been reported that more protective for political influence (Garnett et al. an additional trophy fee of $10,000. standards and guidelines are 2011, p. 1). Given the financial aspects Depending on the country in which a implemented, such as the best practices of sport hunting, it is reasonable to hunter visits, there may be several listed below (Jackson 2013, pp. 3, 8–10, assume that corruption and the inability different fees required, including game Dallas Safari Club 2013, pp. 1–2). to control it could have a negative fees, observer fees, conservation fees, • Minimum trophy quality, sizes, and impact on decisions made in lion permit fees, trophy handling fees, and standards; government payments in terms of taxes, • Wildlife hunting regulations management by overriding biological rationales with financial concerns. as well as safari operator fees (Barnett & enacted and enforced; Patterson 2005, p. 71). In the late 1990’s, • Professional hunting associations Corruption has complex roots and will not end immediately, but from all Tanzania reported annual revenue of formed; • Professional hunting training appearances, it is being addressed in $29.9 million from all trophy hunting, courses; many of the African lion range countries South Africa reported $28.4 million, • Professional hunter standards where it has occurred in the past. Zimbabwe reported $23.9 million from established; Countries throughout the range of the all trophy hunting, Botswana reported • Quota-setting procedures; African lion are putting tools in place to $12.6 million, and Namibia reported • Compliance with CITES combat corruption and create awareness $11.5 million; the revenue generated demonstrated; (http://www.transparency.org/cpi2012/ solely from lion hunting was not broken • Monitoring; and results, accessed June 20, 2013). In out (Barnett & Patterson 2005, p. iv). In • Information and data collection and recent years, in several African lion the past, government and private land analysis. range countries, leadership has taken owners were the primary beneficiaries While the supposition of increased steps to address corruption, or activities of the revenue gained; however, a infanticide due to the remove of that facilitate corruption, associated portion of the revenue derived from established males from a pride is with wildlife management. For example, hunting, in some countries, is now inconclusive and not well supported, it in 2013, the Tourism Minister of Zambia being distributed to local communities is clear that improved management banned hunting in 19 game management as well, which benefits the livelihoods practices are beneficial to maintaining areas for 1 year due to corruption and of local people as well as contributes to

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national economies of African range In Botswana, despite the current ban al. 2012a, p. 9). According to Barnett states (Barnett & Patterson 2005, p. vi). on lion hunting, the country currently and Patterson (2005, p. 82), in has over 128,000 km2 of gazetted Zimbabwe: Trophy Hunting as a Wildlife wildlife management areas and Management Tool Land invasions, resettlement and political controlled hunting areas set aside for instability has had dire consequences for The concept of using trophy hunting hunting purposes, which equates to 22.1 wildlife occurring in the commercial sector. to support lion conservation is complex percent of the country’s total area. This Land invasions have affected all wildlife and counterintuitive to many. Many is in addition to 111,000 km2 (or 19.1 management activities, and resulted in severe range countries rely heavily on tourism percent) that has been set aside as habitat destruction, increased poaching and (predominantly ecotourism and safari habitat in the form of National Parks, infrastructure damage with thousands of hunting) to provide funding for wildlife Game Reserves, and Forest Reserves kilometers of fences being destroyed to make management (IUCN 2006a, p. 24). The wire snares . . . A typical questionnaire (Barnett & Patterson 2005, p. 7). response from an invaded 50,000 acre farm countries that rely most on lion hunting Tanzania has land set aside for sport are proportionally the highest in in Masvingo Province . . . indicates hunting in the form of safari areas, substantial poaching losses of up to Mozambique, Tanzania, and Zambia communal land, and privately owned $1,819,040, with over 3,400 snares recovered (Lindsey et al. 2012a, pp. 7–8). The properties that make up 23.9 percent of and 134 poachers arrested in just two revenue generated from these industries the total land base (Barnett & Patterson months. provides jobs for locals, such as game 2005, pp. 76–77). In 2000, five countries Niassa National Reserve, guards, cooks, drivers, and security in southern Africa (Botswana, Namibia, personnel, and often brings in revenue Mozambique, incurs annual costs of South Africa, Tanzania, and Zimbabwe) approximately $1.9–2 million to for local microbusinesses that sell art, 2 had set aside a combined 420,000 km maintain a 42,000-km2 area (Lindsey et jewelry, and other native crafts. 2 of communal land, 188,000 km of al. 2012a, p. 9). As a single source of Revenue generated from scientifically 2 commercial land, and 420,089 km of revenue, the trophy hunting of lions based management program is used to 2 state land totaling over 1,028,000 km provides a substantial source of funds to build and maintain fences, provide for sport hunting purposes (Barnett & security personnel with weapons and pay for the management of lion habitat. Patterson 2005, p. iii). As a species with According to Lindsey et al. (2012a, p. 5), vehicles, provide resources for anti- a considerable range (up to 1,000 km2) poaching activities, and provides with the exception of rhinoceros and (Packer et al. 2013 p. 636; Haas et al. exceptional elephant trophies, ‘‘lions resources for habitat acquisition and 2005, p. 4), suitable habitat is important management (Chardonnet et al. 2010, generate the highest revenue per hunt of to the survival of the species, and the pp. 33–34; Newmark 2008, p. 321). any species in Africa.’’ In Niassa marked decline in suitable habitat is a Revenue from trophy hunting increases National Reserve, lion trophy hunting significant threat to the species (see the ability of many African countries to has generated $380,000–400,000 Habitat Loss). The land currently manage wildlife populations both annually (Lindsey et al. 2012a, p. 9). In designated for use in sport hunting has within and adjacent to reserves; many of the Save´ Valley Conservancy, between helped to reduce, but not eliminate, the these hunting areas are geographically 2005 and 2011, lion hunting in impact of habitat loss for the African linked to national parks and reserves, Zimbabwe provided an estimated net lion. providing wildlife corridors and buffer income (based on 26 lions) of zones (Chardonnet et al. 2010, p. 34; Cost estimates for maintaining lion approximately $1,365,000 in per-night Newmark 2008, p. 321). populations range, from an annual charges and roughly $260,000 in trophy Proponents and most species experts budget of $500 per km2 in smaller fees (Groom 2013, p. 4). support trophy hunting as a fenced reserves to $2,000 per km2 Trophy hunting of lions, if part of a conservation tool for the African lion annually for unfenced populations scientifically based management (Hunter 2011, entire; van der Merwe (Packer et al. 2013, p. 640; Lindsey et al. program, can provide direct benefits to 2013, entire; Hunter et al. 2013, entire) 2012a, p. 9). This includes but is not the species and its habitat, both at the because it provides: (1) Incentives for limited to costs associated with national and local level (See: Role of the conservation of large tracts of prime permanent and temporary staff, fencing Local Communities in Lion habitat, and (2) funding for park and installation and maintenance (fences Conservation). Trophy hunting and the reserve management, anti-poaching, and can cost $3,000 per km to install), revenue generated from trophy hunting security activities. As habitat loss has infrastructure maintenance, anti- are tools that range countries can use to been identified as one of the primary poaching activities such as surveillance facilitate maintaining habitat to sustain threats to lion populations, it is notable and snare/trap removal, wildlife large ungulates and other lion prey, that the total amount of land set aside restocking fees (both for lions killed by protecting habitat for lions, supporting for hunting throughout Africa, although illegal poaching/snares as well as other the management of lion habitat, and not ameliorating the concerns about trophy species killed by lions on the protecting both lions and their prey base habitat loss, exceeds the total area of the reserves), community outreach, and through anti-poaching efforts. While national parks, accounting for compensation for loss of livestock in hunting alone will not address all of the approximately half of the amount of surrounding communities (Packer et al. issues that are contributing to the viable habitat currently available to 2013, p. 640; Groom 2013, pp. 4–5; declined status of the species, it can lions (Chardonnet et al. 2010, p. 34; Lindsey et al. 2012a, p. 9; Barnett & provide benefits to the species. Packer et al. 2006, pp. 9–10). In Patterson 2005, p. 82). For example, in Tanzania, 25–33 percent of the total the past, the Save´ Valley Conservancy in Role of Local Communities in Lion area, encompassing 190 hunting units Zimbabwe invested $546,000 annually Conservation and over 247,000 km2, has been set on anti-poaching activities and Over the last few decades, aside for sport hunting purposes; this employed 186 permanent scouts, while conservationists and range countries has resulted in an area 5.1 times greater operators in Coutada 16, Mozambique, have realized the integral role local than Tanzania’s fully protected and spent $60,000 annually on anti- communities play in the conservation of gazetted parks (Jackson 2013, p. 6; poaching (such as the removal of 5,000 lions and their habitat; when Barnett & Patterson 2005, p. 61). gin traps) (Groom 2013, p. 5; Lindsey et communities benefit from a species,

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they have incentive to protect it. even more vulnerable to expanding (including Cameroon), seven have Therefore, utilizing the wildlife sector human settlement (Van der Merwe developed National Poverty Reduction as a land-use option and source of 2013, p. 2). Strategies in partnership with the income for rural populations has Community conservancies that benefit International Monetary Fund (for a increasingly been employed throughout from trophy hunting have specifically complete list, see http://www.imf.org/ the range countries of the African lion. been formed as a way to protect wildlife external/np/prsp/prsp.aspx); each of Many of these countries are classified as and habitat. As an example, in Namibia, these has incorporated sustainable ‘developing’ nations; specifically, seven 160,000 km2 (61,776 mi2) of community natural resource development as a main of the ten countries (we include conservancies were established in part priority, and emphasized benefit Cameroon here) where trophy hunting is due to revenue from trophy hunting. distribution and management to rural permitted have 27–64 percent of their These conservancies benefit the local communities (Benin 2000, unpaginated; populations living in severe poverty communities, which in turn protect lion Burkina Faso 2000, unpaginated; (United Nations Development habitat. For example, in 2012, the Save´ unpaginated; CAR 2000, p. 45; Programme’s (UNDP) Human Valley Conservancy (Zimbabwe) Mozambique 2000, unpaginated; Development Report, http:// ‘‘provided over US$100,000 worth of Tanzania 2000, pp. 13, 21; Zambia 2000, hdr.undp.org/en/data, accessed July 7, support to adjacent villages or farmers unpaginated). As a result, an increase in 2014; Barnett & Patterson 2005, p. iii). in the resettled areas. Assistance participation by local communities in These countries often have high included drilling boreholes, maintaining managing natural resources that are population growth, high boreholes, dredging of dams, building adjacent to reserves is occurring in unemployment, limited industry, and a clinics and schools, assisting with several areas. repairs, maintenance and materials for Gross Domestic Product (GDP) per Captive Lions capita lower than the poverty level schools, education initiatives, school (Barnett & Patterson 2005, p. iii). These field trips, provision of computer In analyzing threats to a species, the combined challenges highlight the need equipment in schools, and craft Service focuses its analysis on threats for innovative solutions. programs’’ (Groom 2013, p. 5) acting upon wild specimens within the Conservationists and range countries Connecting conservation to community native range of the species, because the recognize the value of the wildlife benefits can provide a value for wildlife, goal of the Act is survival and recovery sector; if managed sustainably, there is including lions, where there was of the species within its native high potential to contribute to rural previously resentment or indifference, ecosystem. We do not separately analyze ‘‘threats’’ to captive-held economic development while helping to instill a sense of importance specimens because the statutory five simultaneously protecting the unique for lion conservation Additionally, an factors under section 4 (16 U.S.C. 1533) ecological habitats and species estimated 125,000 kg of game meat is are not well-suited to consideration of contained therein (Chardonnet et al. provided annually to rural communities specimens in captivity and captive-held 2010, p. 33; Kiss [editor] 1990, pp. 1, 5– by trophy hunters at an estimated value specimens are not eligible for separate 15). of $250,000 per year, which is considerable for rural locations where consideration for listing. However, we Studies have indicated that, in order severe poverty and malnutrition exists do consider the extent to which for species such as the African lion to (White 2013, p. 21), further providing a specimens held in captivity create, persist, the local communities must value for wildlife, including lions. contribute to, reduce, or remove threats benefit from or receive a percentage of Lastly, local communities benefit from to the species. funds generated from tourism such as the trophy hunting industry by gaining Captive-held African lions, including wildlife viewing, photography, or employment as cooks, drivers, game those that are managed for trophy trophy hunting (White 2013, p. 21; guards, security, and anti-poaching hunting in South Africa and lions held Martin 2012, p. 57; Kiss [editor] 1990, personnel, and they also obtain revenue in captivity in zoos, are believed to pp. 1, 5–15). The economic value of a for items purchased by trophy hunters number between a few thousand and species, such as lion, can encourage such as jewelry, art, and native 5,000 worldwide (Republic of South range countries to develop management handicrafts. Africa 2013, p. 5; Barnett et al. 2006a, and conservation programs that involve Trophy hunting as part of a p. 513). Captive lions in general are not local communities which would scientifically based management suitable for reintroduction due to their ultimately discourage indiscriminate program may provide direct economic uncertain origins (Barnett et al. 2006a, killings by local communities (Groom benefits to the local communities and p. 513; Hunter et al. 2012, p. 3), 2013, pp. 3, 5; Hazzah et al. 2013, p. 1; can create incentives for local potential maladaptive behaviors, and White 2013, p. 21; Martin 2012, p. 49). communities to conserve lions, reduce higher failure risk compared to If local communities see no beneficial the pressure on lion habitat, and end translocated individuals (Hunter et al. value of lions being present in their retaliatory killing, primarily because 2012, pp. 2–3). There may be cases communal areas, sustainable utilization lions are viewed as having value. where captive specimens provide a of lions as a land-use becomes less Conversely, lack of incentives could benefit to the species under certain competitive with other land-use cause declines in lion populations circumstances. For example, the display options, such as grazing and livestock because lions are viewed as lacking of Giant pandas in U.S. zoos has management, and local communities value and are perceived to kill livestock, generated considerable revenue that is become unwilling and unable to manage which do have value to communities used for in-situ conservation of the their wildlife heritage (Barnett & (see Human-lion Conflict). species in China. It may be possible that Patterson 2005, p. iii). When the value Many range countries have realized captive lions could also serve a purpose of lions in areas outside of national local communities must benefit from the of generating revenue for in-situ parks is diminished, those areas are conservation of the species because conservation. likely to be converted to forms of land [why?] and have revised their land use less suitable for lions, such as management and ownership policies to Summary of Trophy Hunting agriculture, livestock pastures, or areas reflect this. Of the ten countries where Although there is some indication of resource extraction, making them lion trophy hunting currently occurs that trophy hunting could contribute to

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local declines in lion populations needed revenue, jobs, and a much- anecdotal in nature, based on the best through unsustainable quotas, needed protein source to local people, available scientific and commercial corruption, and possible disruption of demonstrating the value of lions to local information, we find that the sale of pride structure through infanticide and communities (Groom 2013, pp. 1–3; these byproducts does not currently take of males that are too young, we do Lindsey et al. 2006, pp. 283, 289). In pose a threat to the species. Further, not find that any of these activities rises addition, the amount of habitat that has without a significant shift in the market, to the level of a threat to the African been set aside by range countries it is not likely to become a threat in the lion subspecies at this time. It appears specifically for trophy hunting has foreseeable future. that most range countries that allow greatly increased the range and habitat Conservation Measures in Place To trophy hunting of African lions restrict of lions and their prey base, which is Protect Lions offtake to approximately 2–4 percent of imperative given the current ongoing their lion populations for trophy rate of habitat destruction occurring in There has been awareness for several hunting annually, excluding South Africa. The total amount of land set years that conservation strategies need Africa, where offtake is from aside for trophy hunting throughout to be implemented for the African lion predominantly captive-born animals, Africa exceeds the total area of the due to the apparent decrease in its and Zimbabwe, where offtake is 2–3 national parks, providing half the population numbers (Hamunyela et al. percent higher than in other countries amount of viable lion habitat 2013, p. 1; Henschel et al. 2010, p. 34; (Packer et al. (2006, pp. 2–3). Exports of (Chardonnet et al. 2010, p. 34; Packer et Gebresenbet et al. 2009, p. 5; IUCN lion trophies have demonstrated a al. 2006, pp. 9–10). However, expanding 2006a, b, entire). Prior to 2006, decreasing trend when exports of likely protected areas without taking the institutional inconsistencies throughout captive-born lions from South Africa are human population into consideration the African lion’s range resulted in poor excluded (CITES lion gross exports, could lead to more resentment and lion conservation policies and little to http://trade.cites.org, accessed April 23, retaliatory killing of lions (Nelson et al. no enforcement of existing laws (IUCN 2014), and lions from South Africa are 2009, p. 315). 2006b, p. 18). As mentioned, in 2005 likely captive-born (RSA 2013, p. 5). Therefore, we conclude, based on the and 2006, nongovernmental Most of the range countries that allow best scientific and commercial organizations (NGOs) and several trophy hunting have quotas in place to information available, that trophy governments at various levels organized limit take. Tanzania, with a population hunting is not a significant threat to the two regional lion conservation of approximately 16,000 lions, has a species. workshops. Species specialists, wildlife quota of 50 animals per year. Many managers, and government officials Traditional Use of Lion Parts and attended these regional workshops in other range countries have laws in effect Products that address trophy hunting, and several order to provide range country have moratoriums in place. The hunting CITES (2014, p. 8) reports that many governments with frameworks for community is taking the lead in African countries, including Somalia, developing their own national action developing best management practices Nigeria, Burkina Faso, Kenya, and plans for the conservation of lions. Over to address take of males that are under Cameroon, maintain local markets in 50 lion specialists, representing all lion 6 years of age, and they are guiding the lion products, which include teeth, range countries, participated in these development of scientifically based claws, fat, whiskers, bone, bile, testicles, workshops (Henschel et al. 2010, p. 34). tools for minimizing the impact of meat, and tails for use as talismans, During the workshops lion experts trophy hunting on the social structure of decorations, and in traditional African collectively assessed what they believed lion populations. This 6-year age medicine. In Ghana, lion parts and to be the then-current status of African restriction on lion trophies is in the products are used for ceremonial, lions based on a variety of information, process of being self-implemented by medicinal, and nutritional purposes and subsequently identified 86 African professional hunting guides, and is (Burton et al. 2010, p. 4). Skins and LCUs. This information was then used being adopted by certain range states, claws of lions were observed for sale in as a framework to identify lion areas, such as Tanzania (White 2013, p. 14; a market in Tamale, Ghana. Lions in and strongholds, and potential strongholds Whitman et al. 2004, p. 176). around Mole National Park in Ghana by Riggio et al. (2013, p. 32). Currently, most countries that allow have been killed for traditional Many countries with very small lion trophy hunting of lions appear to be consumptive purposes (Burton et al. populations have developed or updated reviewing their trophy hunting practices 2010, p. 4). In some cases, lions (either their conservation plans for the African (Jackson 2013, pp. 2–3; White 2013, pp. alive or dead) have been ‘‘laundered’’ lion. Some of these include Benin, 12–13). Range countries have through other countries so that their Cameroon, Uganda, and Malawi. Some recognized the need to incorporate best country of origin is unknown. As an range countries participate in management practices, and have been example, lions have been found to be transboundary conservation projects progressively updating the policies and shot in Zimbabwe and Mozambique and and are collaborating on transboundary management systems in order to declared as South African trophies (Lion lion conservation initiatives for shared implement them (Lindsey et al. 2013a, Aid 2011, p. 20). In other cases, there lion populations. Most range countries pp. 4–10). have been reports of captive-born lions have a national lion action plan or Finally, we found that, if trophy being smuggled between Botswana and strategies in place, particularly if there hunting of lions is part of a scientifically South Africa and described as wild are economic incentives for them to based management program, it could (Mouton 2013, pp. 1–2). Lion products, have viable lion populations (Groom provide considerable benefits to the such as the trade in lion bone, seem to 2013; Nghidinwa et al. 2013, pp. 11–12; species, by reducing or removing be primarily byproducts of trophy Zambia Wildlife Authority 2012; Lion incentives by locals to kill lions in hunting; hunters are primarily Aid 2011, pp. 1–2; Mesochina et al. retaliation for livestock losses, and by interested in the trophy and skin and, 2010; Government of Tanzania 2010; reducing the conversion of lion habitat therefore, the bones and other parts are Begg and Begg 2010). Range states have to agriculture. Trophy hunting, if sold separately (CITES 2014a, p. 10). also implemented a number of managed well and with local However, since the reports of these conservation strategies designed to communities in mind, can bring in types of activities are primarily conserve habitat, reduce human-lion

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conflict, and preserve the lion’s prey- human populations, declining km corridor through an area that had base. purchasing power of external funds, and been heavily impacted by human- corruption (Garnett et al. 2011, pp. 1–2; wildlife conflict. The purpose of the Conservation Measures To Stem Habitat Wittemyer et al. 2008, pp. 123, 125). corridor was primarily to reduce Loss Another mechanism for protecting human-elephant conflict and appears to Habitat loss represents one of the habitat is to reconnect fragmented have been successful (Mount Kenya main threats facing the African lion habitat across national boundaries. Trust 2011, p. 1). The corridor also (Bauer et al. 2008, unpaginated). Corridors are being restored, fences are allows other wildlife such as lions to Attempts by range countries to address being removed, and protected areas are disperse through habitat that otherwise this decline in habitat are manifested in being connected. Restoration of these would have been unfavorable for a number of ways, such as the creation corridors allows wildlife to travel wildlife to travel through (Mount Kenya of protected areas and the establishment between areas of suitable habitat (Jones Trust 2011, p. 1). It was an expensive of wildlife corridors to connect et al. 2012, pp. 469–470). In some areas, project, but recent reports indicate that fragmented habitats. fences have been constructed to protect the effort has served its purpose: Two conservation tools utilized by grazing resources for domestic livestock Elephants are using the corridor on a range countries for African lions include as well as to provide barriers to disease regular basis (particularly an underpass the establishment of protected areas and (Gadd 2012, pp. 153, 176). One aspect under a highway), and humans are the enforcement of protections in these of these fences is that they separate reporting less human-wildlife conflict areas (Mesochina et al. 2010a and b; lions from their prey. In southern (Mount Kenya Trust 2011, p. 1). Treves et al. 2009, pp. 60, 64). Over the Africa, the trend now is to take down However, connectivity alone does not past few decades, the effectiveness of fences to increase the size of connected ensure the dispersal of animals (Roever protected areas in protecting habitat has habitat and link it to reserves and et al. 2013, pp. 19–21). The Tanzania been studied, particularly in Africa national parks (IUCN 2009, p. 101; Wildlife Research Institute (TAWIRI) is (Pfeifer et al. 2012, p. 1; Craigie et al. IUCN 2008, various). The Limpopo a parastatal organization under 2010, pp. 2,221–2,222). A study Transfrontier Park is another example of Tanzania’s Ministry of Natural conducted by the Wildlife Conservation where this is being implemented Resources and Tourism, and is Society in 2005 found that most lion (Newmark 2008, p. 327). Boundary responsible for conducting and populations in protected areas of fences along national borders that coordinating wildlife research activities southern and eastern Africa have been separate many reserves are being in Tanzania (http://tawiri.or.tz/). In this essentially stable over the previous removed to form a 35,000-km2 park. role, TAWIRI has been actively involved three decades (Ray et al. 2005, p. 67). Limpopo National Park (formerly in promoting the development of and However, several problems have known as Coutada 16) in Mozambique; monitoring the use of wildlife corridors emerged. For example, certain land- Kruger National Park in South Africa; in Tanzania (http:// tenure systems do not recognize Gonarezhou National Park, Manjinji Pan www.tzwildlifecorridors.org). Surveys community ownership of land and Sanctuary, and Malipati Safari Area in conducted in 2009 and 2010 suggest wildlife and undermine the extent to Zimbabwe will all be connected, as will that the Nyanganje Corridor in Tanzania which benefits are converted into be the area between Kruger and is no longer being used by elephants incentives for conservation. Protected- Gonarezhou, and the Sengwe communal and other wildlife. This corridor is at a area ‘‘boundaries’’ are not always land in Zimbabwe and the Makuleke narrow passage in the Kilombero Valley visible. Additionally, law enforcement region in South Africa (Newmark 2008, and is the shortest distance for animals in protected areas can be sporadic, and p. 327). However, in some locations, to cross between the Udzungwa and parks are often understaffed (Pfeifer et areas that have previously been Selous ecosystems. Despite efforts in al. 2012, pp. 1, 7). Lastly, despite the designated as corridors have been place, much of the corridor is being Wildlife Conservation Society’s encroached upon by human settlements encroached upon by conversion of land findings, more recent evidence suggests and agriculture (Estes et al. 2012, pp. to rice farming and cattle grazing (Jones that some protected areas are being 258–261; Jones et al. 2012, p. 469). et al. 2012, p. 469). Because these more commonly encroached upon as Tanzania is an example of a country activities often deter wildlife from human populations expand and search attempting to reconnect habitat. As of passing through, the corridor is for resources. 2002, the Tanzanian Government, with ineffective (Jones et al. 2012, p. 469). Despite encroachment, protected donor and NGO support, was TAWIRI reminds wildlife managers that areas are somewhat effective at reconnecting the nine largest blocks of they need to continue to implement protecting wildlife and habitat as rates forest in the East Usambara Mountains steps to ensure that corridors are of habitat loss tend to be lower in using wildlife corridors (Newmark 2002, functioning properly. protected areas than outside them various). Additionally, the 2009 (Green et al. 2013, p. 70; Pfeifer et al. Wildlife Act of Tanzania allows the Conservation Measures in Place To 2012, p. 2). African countries are Minister, in consultation with relevant Stem the Loss of Prey Base realizing the benefits of managing their local authorities, to designate wildlife Lions, like most large carnivores, prey wildlife populations and parks for corridors, dispersal areas, buffer zones, upon a variety of species including tourism; however, conservation of vast and migratory routes. The 2010–2015 buffalo, plains zebra, wildebeest, giraffe, areas of land for megafauna such as the National Elephant Management Plan of gemsbok, kob, and warthog (Kenya African lion is not only complex, but Tanzania indicates that corridors are the Wildlife Service 2013, p. 13; Niassa also expensive. As an example, the 28- primary objective of the plan, and National Reserve Technical Report km (17-mi) elephant corridor, although primarily designed for 2011, p. 4; Nowell and Jackson 1996, p. completed in 2011 in Kenya, cost $1 elephants, these corridors allow for 18). Depletion of these prey species due million (The Nature Conservancy 2013, continuity of populations of other large to competition with humans represents unpaginated). Additionally, the overall mammal species such as lions (Jones et a threat to the lion (Chardonnet et al. costs of anti-poaching and al. 2012, p. 470). 2005, pp. 8–9). As noted, the increase in compensation is expected to increase in In 2011, Kenya (which neighbors the human population in Africa is a range states concurrently with growing Tanzania to the North), completed a 28- major contributor to the increase in the

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demand for bushmeat, which in turn area as a whole. This open approach to npn; NASCO 2010, p. 4). The majority increases human encroachment into protected area management reflects a of the incomes from these conservancies wildlife lands (Lindsey et al. 2012b, p. trend in recent years to bring in local come from ecotourism, followed by 36). In addition to the increase in the communities to assist in the trophy hunting (NASCO 2010, p. 22). human population, lack of an management of protected areas (Lindsey These incomes are then used to support alternative livelihood, lack of alternate et al. 2012b, p. 53). Wildlife infrastructure improvement in the food sources, and lack of clear rights management programs run by local community. In addition, legal bushmeat over land or wildlife are contributing communities are defined by two goals: acquired within conservancy lands is factors toward the increase in demand Conserving wildlife and providing distributed to local families (NASCO for bushmeat (Lindsey et al. 2012b, pp. economic aids to the community 2010, p. 25). The success of the program 36–41). The advent of automatic (Bandyopadhyay et al. 2010, p. 5). With in Namibia has been attributed to weapons in the bushmeat trade impacts regards to discouraging the Namibia’s unique characteristics, the lion’s prey base, which is being consumption of bushmeat, this new including low population density and hunted at unsustainable levels. approach is seen in the creation of favorable seasonal rain, which helps Reconnecting fragmented habitat has community-based wildlife management prey species recover (van Vliet 2011, p. the additive effects of not only programs (van Villet 2011, p. 26). The 30). Despite the successes in Namibia, conserving the biodiversity of the purpose of these programs is to give the the country’s unique characteristics African lion’s habitat, but also that of its local community a direct stake in the mean that adapting Namibia’s success to prey base (Lindsey et al. 2012b, p. 43). management of wildlife areas. One use other, more densely populated countries These types of restoration practices for these areas is to turn them into game will be difficult. enhance the health of species by ranches. These areas are used both for allowing genetic interchange to occur legal bushmeat production as well as Conservation Measures To Stem and, thus, conserve the genetic diversity trophy hunting and ecotourism. Human-Lion Conflict of all wildlife. Wildlife management One such program is the Chivaraidze As the human population expands, entities are linking many of the major Game Ranch in Zimbabwe (van Villet the potential for conflict with wildlife protected areas by removing boundary 2011, pp. 28–29). The Chivaraidze Game increases. In Africa, conflict between fences along national borders that Ranch started in 1996 with the stated villagers and lions, who prey upon separate many reserves in addition to goal of reducing poaching through livestock, represent a threat to the creating or improving corridors to link providing bushmeat at a reduced price. species (Chardonnet et al. 2010, p. 12; good-quality habitat for wildlife (Gadd However, internal infighting in the Moghari 2009, p. 14; IUCN 2006a, p. 2012, p. 179; Newmark 2008, pp. 323– organization over the devolution of 23). In addition, habitat loss due to 324). To address the increasing power to local communities, between conversion of land increases the chance consumption of bushmeat, host those in favor of devolution and a of villagers coming into direct contact countries have employed a variety of powerful local interest group, limited with lions (Chardonnet et al. 2010, p. different strategies, including the the effectiveness of the organization. In 24). In an attempt to address these development of alternative industries the span of 8 years (between 2001 and problems, range countries have for communities. Helping local 2009), the Chivaraidze Game Ranch has employed a variety of different communities develop alternate had six different boards of directors strategies to help the lion. Such industries represents one of the ways (Mombeshora and Le Bel 2010, p. 5). strategies involve education, an effective range countries can reduce their Furthermore, a power shake-up in local conservation plan, and interacting with dependence on bushmeat. Throughout communities along party lines and the local community. Africa, several ideas have been kinship affiliation limited the abilities attempted with varying levels of for communities to cooperate with each Historically, range countries seek to success. For example, the Anne Kent other (van Villet 2011, pp. 28–29; mitigate human-lion conflict through Taylor Fund (AKTF) helps local Maasai Mombeshora and Le Bel 2010, p. 7). The controlling rather than conserving the women to buy beads and other supplies result was that the cost of maintaining predator population. In countries such to produce traditional items for the local the program exceeded the benefits to the as Malawi, for example, the Department tourist industry (AKTF 2012, p. 7; local community. The decline in of Game, Fish and Tsetse Control would Lindsey et al. 2012b, p. 45; van Villet economic benefits to the local shoot large carnivores that prey upon 2011, p. 17). In addition, AKTF helps community coincided with a resurgence livestock. The result of this policy was organize local men into anti-poaching in poaching within areas of the park that, between 1948 and 1961, over 560 and de-snaring teams (AKTF 2012, p. 5; (Mombeshora and Le Bel 2010, p. 3). predators (which include lions and van Villet 2011, p. 17). By creating The result of the Chivaraidze Game leopards) were killed in the country programs targeting both men and Ranch project reflects the difficulty in (Mesochina et al. 2010b, p. 35). While women, AKTF creates an environment shifting wildlife management from a this department was disbanded in 1963 that provides communities with centralized national government and jurisdiction shifted to the new financial stability as well as direct approach towards a more decentralized, Department of Forestry, crop and community interest in protecting local community-based approach. livestock protection still remains an wildlife. With 13 years assisting local Unlike the difficulties encountered in important part of its function. Despite communities, the AKTF represents one Zimbabwe, Namibia has had greater the department focusing on protecting of the more successful attempts to success in setting up community-run crops and livestock, the number of lions encourage locals to shift away from conservancies. After gaining killed in the country has declined. relying on bushmeat. independence in 1990, Namibia began Between 1977 and 1982, eight lions Studies compiled by Huzzah 2013 to turn over ownership of wildlife areas were killed, whereas six lions were (pp. 1, 8) have shown that local to local communities (van Vliet 2011, p. killed between 1998 and 2007 communities who lived near protected 29; Bandyopadhyay et al. 2010, p. 6). By (Mesochina et al. 2010b, p. 35). While areas with more lenient policies have a 2011, Namibia had 64 communities that fewer lions are being killed than in the more positive attitude and relationship covered 17 percent of the country total previous decades, problems remain, with both the manager and the protected area (van Vliet 2011, p. 29; Connif 2011, including lack of resources, lack of

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manpower, and corruption within the al. 2014, p. 858; Lion Guardians 2013, conservation areas, and to conserve and range countries. p. 20). Since its establishment in 2007, regulate the take, hunting, and trade of Current governmental management of only five lions had been killed in wildlife, including parts and products, lions in countries such as Malawi, territories where Lion Guardians but could find no legislation specific to Tanzania, and Zambia are managed by operates, in contrast to more than 100 lions, nor to the main threats affecting the Problem Animal Control units lions killed in adjacent areas (Lion lions: habitat loss, human-lion conflict, (Mesochina et al. 2010a, p. 41; Guardians 2013, p. 5). Furthermore, and loss of prey base (See: Appendix A, Mesochina et al. 2010b, p. 36). When reduced lion mortality was sustained Ecolex information was accessed July 7– lion attack incidents occur, Problem across multiple years, resulting in the 10, 2014, at http://www.ecolex.org.8). Animal Control dispatches officials to reserve having one of the highest lion Our status review did not reveal investigate the problems. If the problem densities in Africa (Hazzah et al. 2014, regulatory mechanisms in place that lion is located, it is either removed or p. 857; Schuette et l. 2013, p. 149). specifically address the main threats eliminated. When properly funded, this Despite the success of this program, affecting lions. We are requesting program has helped in reducing not retaliatory as well as ceremonial killings comments or information from lion only conflicts between lions and of lions outside the program areas range states, other concerned humans but also has driven down the remain a threat to the species. governmental agencies, the scientific numbers of lions killed. Between 2005 We found that many of the lion range community, or any other interested and 2009, there were 116 reported cases states are trying to address lion parties concerning regulatory of lions killed, with the number of lions conservation through the establishment mechanisms that address the three main killed being less than 50 per year in of protected areas, wildlife management threats to lions: habitat loss, human-lion Tanzania (Mesochina et al. 2010a, p. areas, wildlife corridors, and conflict, and loss of prey base. 41). However, limitations of resources reconnecting habitat. In some areas, Finding (including both manpower and funds) creating incentives for lion conservation have hampered the effectiveness of is occurring through community Section 4 of the Act (16 U.S.C. 1533) these officials in responding to these conservation programs in range and its implementing regulations at 50 incidents. In addition, many Problem countries. In other cases, participatory CFR 424 set forth the procedures for Animal Control interventions resulted strategies have been implemented to adding a species to, and/or removing a in the death of the lion (Mesochina et enhance local tolerance for large species from, the Federal Lists of al. 2010a, p. 41; Chardonnet et al. 2009, carnivores in Africa. An increasing Endangered and Threatened Wildlife p. 36). Even in cases of translocation, number of programs encourage local and Plants. As noted in the Information the lions that were being transported communities to solve problems that Requested section, a species may be often end up injured or continue to pose arise from human-lion conflict without determined to be an endangered or problems to the community (Bauer et al. killing lions. However, the effectiveness threatened species due to one or more 2007, p. 91). of these measures still ranges from of the five factors set forth in section NGOs are also assisting in protecting successful to unsuccessful, due in part 4(a)(1) of the Act: lions. Intervention by NGOs often takes to lack of resources, political will, and (A) The present or threatened the form of interacting with the local infighting. It is imperative that range destruction, modification, or community (Winterbach et al. 2010, p. countries continue to recognize and curtailment of its habitat or range; 98). Lion Guardians, which operate in support the role that local communities (B) Overutilization for commercial, Kenya, recruits and educates local play in lion conservation. Greater recreational, scientific, or educational young men. These men then monitor support by countries to address the purposes; and track lion movement and warn needs of local communities, and thereby (C) Disease or predation; (D) The inadequacy of existing herders of lion presence in the area, address the needs of lions, may be the regulatory mechanisms; thereby mitigating or preventing single-most important role these (E) Other natural or manmade factors possible lion-human conflict (Hazzah et countries can play in changing the affecting its continued existence. al. 2014, p. 853; Lion Guardians 2013, trajectory of lion declines. In assessing whether the African lion p. 7; Lion Guardians 2012, p. 3). In meets the definition of an endangered or addition, Lion Guardians work with Regulatory Mechanisms threatened species, we considered the tribal elders to dissuade young men Regulatory mechanisms in place to five factors in section 4(a)(1) of the Act. from killing lions for ceremonial provide protections to African lions A species is ‘‘endangered’’ for purposes purposes. Historically, the killing of vary substantially throughout Africa. As of the Act if it is in danger of extinction lions through ritualized lion hunts mentioned in the Conservation Status of throughout all or a significant portion of called ilmurran is rewarded with gifting African Lions CITES section, lions are its range and is ‘‘threatened’’ if it is of cows and other rewards (Lion listed in Appendix II under CITES, and likely to become endangered within the Guardians 2012, p. 5; Goldman et al. with the exception of South Sudan, all foreseeable future throughout all or a 2010, p. 334). After introducing village of the lion range states are parties to significant portion of its range. The elders to the Lion Guardians program CITES. According to the draft CITES first hand, many return home to their Periodic Review of the Status of African ‘‘foreseeable future’’ is the period of village and give their blessings to the Lions (CITES 2014a, pp. 14–15) outside time over which events or effects project. This education led to significant of CITES, lions have no legal protections reasonably can or should be anticipated, results; on August 11, 2013, two Lion in four countries: Burundi, Guinea or trends extrapolated. Guardians stopped a group of hunters Bissau, Lesotho, and Swaziland. 8 ECOLEX is a comprehensive database on who were planning to hunt a lion in However, CITES 2014a (p. 15), states environmental law, maintained by the International retaliation for the lion preying on their that most of the southern and eastern Union for Conservation of Nature (IUCN), the livestock. The local village elders fined lion range states have regulatory United Nations Environment Programme (UNEP), the potential hunters two cattle each for mechanisms in place to protect lions. and the Food and Agriculture Organization of the United Nations (FAO). Our search terms used with going on a lion hunt, marking a gradual We found that most of the range states respect to wildlife laws were ‘‘African lion’’ and but significant shift in the cultural have national environmental legislation ‘‘country’’, e.g., ‘‘Angola’’, ‘‘Benin’’, etc. See attitudes regarding the lion (Hazzah et to establish national parks and Appendix A.

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When considering what factors might Saharan Africa’s human population into automatic weapons is a significant constitute threats to a species, we must the foreseeable future. threat to lion prey (Chardonnet et al. look beyond the mere exposure of the Africa has the fastest population 2010, p. 27). Because many wildlife species to a factor to evaluate whether growth rate in the world (UNEP 2012a, species are being hunted at the species may respond to the factor in p. 2). The majority of the population is unsustainable levels to meet this a way that causes actual impacts to the rural, and about 60–70 percent of the demand within the range of the lion, its species. If there is exposure to a factor population relies on agriculture and prey base is becoming depleted in many and the species responds negatively, the livestock for their livelihood (UNEP areas and has led to lion attacks on factor may be a threat and we attempt 2006, pp. 82, 100, 106; IAASTD 2009, p. livestock and humans (Hoppe-Dominik to determine how significant a threat it 2). As a result, a large portion of the et al. 2011, p. 452; Chardonnet et al. is. The threat is significant if it drives, growing population will depend 2010, pp. 6, 13–14; Frank et al. 2006, p. or contributes to, the risk of extinction directly on expansion of agriculture and 12). Given the rapid increase in humans of the species such that the species may livestock grazing to survive in the and livestock by 2050, we can warrant listing as endangered or future. Between 2010 and 2050, the reasonably expect the conditions threatened as those terms are defined in population of sub-Saharan Africa is described above to worsen. Also, as the Act. We conducted a review of the projected to more than double to more livestock numbers increase and as best scientific and commercial data than 2 billion (from 831 million to 2.1 expansion of agricultural and pastoral available regarding the status of the billion) (UN 2013, p. 9). During about practices continue to deplete and African lion and assessed whether the this same time period (2005 to 2050), degrade the habitat that lion’s prey rely African lion is endangered or threatened the area of cultivated land is projected on, the lion’s prey base is expected to throughout all of its range. to increase by 51 million ha further decline. As the lion’s prey base There is consensus within the (approximately 21 percent) is hunted at unsustainable levels to research community as well as lion (Alexandratos and Bruinsma 2012, p. meet a growing demand for food, range states that the African lion is 107). However, this figure does not livestock depredation and retributive impacted by a number of factors actively include rangeland, and the majority of killing of lions through spearing, contributing to its population decline agricultural land in Africa is devoted to shooting, trapping, and poisoning will throughout Africa: habitat loss grazing (UNEP 2012b, p. 68), thus that continue to occur, and will likely (fragmentation and degradation) (Factor figure may be much larger. The number increase (Dickman 2013, p. 379; Hoppe- of livestock (cattle, sheep, and goats) in A); decreased access to food prey Dominik et al. 2011, p. 452; Chardonnet sub-Saharan Africa is projected to sources (aka loss of prey base) (Factor et al. 2010, p. 19; Gebresenbet et al. increase about 73 percent, from 688 B); retaliatory killing, snaring, and 2009, p. 9; Hazzah and Dolrenry 2007, million to 1.2 billion, by 2050 poaching (both intentional and p. 3). (Alexandratos and Bruinsma 2012, p. Lion range countries are aware of the unintentional), and deleterious effects 133). Therefore, in the case of African threats affecting lions, and many are in its viability due to small populations lion, the best available scientific and working to address them. NGOs and in some areas within its range (Factor E) commercial data that we rely upon in several governments at various levels (Nyanganji et al. 2012, p. 12; Seguya et projecting future conditions for the have organized regional lion al. 2010, p. 26). purpose of this listing determination conservation workshops, which have We find three main threats, habitat establish the foreseeable future to be helped them to identify Lion loss, loss of prey base, and human-lion 2050. Conservation Units. Most range conflict, are impacting lions, alone and Human settlements and agricultural countries have a national lion action in combination, such that the and pastoral activities have expanded plan or strategy in place (Groom 2013; subspecies is likely to become into lion habitat and protected areas, Nghidinwa et al. 2013, pp. 11–12; endangered within the foreseeable decreasing prey availability and Zambia Wildlife Authority 2012; Lion future throughout all of its range. In the increasing exposure of livestock and Aid 2011, pp. 1–2; Mesochina et al. past several decades, the human humans to lions. Human-lion conflict 2010; Government of Tanzania 2010; population has been expanding with and associated retaliatory killing of Begg and Begg 2010). Some range concomitant large decreases in lion lions will continue to play a major role countries participate in transboundary habitat and likely lion numbers, in the reduction of lion populations and conservation projects to create wildlife resulting in an extremely large is the greatest current threat to corridors and reconnect habitat, and are reduction in the species’ range. As remaining lion populations. The lion’s collaborating on transboundary lion human populations continue to rise in prey base has decreased in many parts conservation initiatives for shared lion sub-Saharan Africa, the amount of land of its range in large part due to the populations. Reconnecting fragmented required to meet the expanding human bushmeat trade habitat has the additive effects of not population’s needs is constantly Bushmeat is the primary source of only strengthening the biodiversity of increasing. Lions are increasingly protein for humans in much of the lion’s the African lion but also that of its prey limited to protected areas, and human range (Chardonnet et al. 2010, p. 27; species (Lindsey et al. 2012b, p. 43). population growth rates around Mesochina et al. 2010a, p. 38; Abwe and Wildlife management entities are protected areas in Africa tend to be Morgan 2008, p. 26; Bennett et al. 2007, linking many of the major protected higher than the average rural growth p. 885; Fa et al. 2006, p. 507), areas by removing boundary fences rate (Wittemyer et al. 2008, entire). comprising between 6 percent (southern along national borders that separate Considering the majority of the human Africa) and 55 percent (Central African many reserves, in addition to creating or population in sub-Saharan Africa is Republic) of a human’s diet (Chardonnet improving corridors to link good-quality rural, and land supports the livelihood et al. 2005, p. 9; IUCN 2006b, p. 19). habitat for wildlife (Gadd 2012, p. 179; of most of the population, loss and This reliance by humans on protein Newmark 2008, pp. 323–324). degradation of lion habitat, loss of prey obtained from bushmeat results in direct Range states have also implemented a base, and increased human-lion conflict competition for prey species between number of conservation strategies can reasonably be expected to humans and lions, and commercial designed to conserve habitat, reduce accompany the rapid growth in sub- poaching of wildlife through the use of human-lion conflict, and preserve lion

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prey-base. In order to address the ceremonial purposes. The result of such In Tanzania, which is home to 40 increasing consumption of bushmeat, programs has been a gradual change in percent of all lions, land set aside for host countries have employed a variety cultural attitudes towards lions (Hazzah sport hunting purposes has resulted in of different strategies, including the et al. 2014, p. 858; Lion Guardians 2013, an area 5.1 times greater than Tanzania’s development of alternative industries p. 20). fully protected and gazetted parks for communities, which can reduce Finally, many range countries rely (Jackson 2013, p. 6; Barnett & Patterson their dependence on bushmeat. For heavily on tourism (predominantly 2005, p. 61). In Botswana, despite the example, the Anne Kent Taylor Fund ecotourism and safari hunting) to current ban on lion hunting, the country (AKTF) helps local Maasai women to provide funding for wildlife currently has more than 128,000 km2 of buy beads and other supplies to produce management (IUCN 2006a, p. 24). The gazetted wildlife management areas and traditional items for the local tourist revenue generated from these industries controlled hunting areas set aside for industry (AKTF 2012, p. 7; Lindsey et can be critical to fund wildlife hunting purposes, which equates to 22.1 al. 2012b, p. 45; van Villet 2011, p. 17) management programs in range states. percent of the country’s total area; this and has organized local men to Tourism, through ecotourism and is in addition to 111,000 km2 (or 19.1 participate in anti-poaching and de- trophy hunting, can provide jobs to percent) that has been set aside as snaring teams (AKTF 2012, p. 5; van locals (such as game guards, cooks, habitat in the form of National Parks, Villet 2011, p. 17). By targeting both drivers, security personnel) and often Game Reserves, and Forest Reserves men and women in the community, brings in revenue for local (Barnett & Patterson 2005, p. 7). In 2000, such programs provide communities microbusinesses that sell art, jewelry, five countries in southern Africa with financial stability as well as direct and other native crafts. Lions can (Botswana, Namibia, South Africa, community interest in protecting local generate the highest daily rate of any Tanzania, and Zimbabwe) had set aside wildlife. African countries are realizing mammal hunted (USD $2,650 per day), a combined 420,000 km2 of communal the benefits of managing their wildlife the longest number of days that must be land, 188,000 km2 of commercial land, populations and parks for tourism; booked, and the highest trophy fee and 420,089 km2 of state land totaling however, conservation of vast areas of ($24,500) (Jackson 2013, p. 6; Lindsey et more than 1,028,000 km2 for sport land for megafauna such as the African al. 2012a, p. 5), thus generating hunting purposes (Barnett & Patterson lion is expensive. The costs of anti- significant revenue for range countries. 2005, p. iii). As a species with a poaching and compensation is expected Creating community-based incentives to considerable range (up to 1,000 km2) to increase in range states concurrently conserve lions from revenue derived (Packer et al. 2013 p. 636; Haas et al. with growing human populations, from trophy hunting may ameliorate the 2005, p. 4), suitable habitat is important declining purchasing power of external human-lion conflict that arises from to the survival of the species, and the funds, and corruption (Garnett et al. lions and humans coexisting in the marked decline in suitable habitat is a 2011, pp. 1–2; Wittemyer et al. 2008, same area. significant threat to the species. The Revenue from scientifically based pp. 123, 125). habitat currently preserved for use in management programs that include sport hunting has helped to reduce the Studies have shown that local trophy hunting can increase the ability impact of habitat loss for the African communities who live near protected of many African countries to manage lion, but as discussed previously, areas (PAs) with community-based wildlife populations both within and habitat loss remains a significant threat conservation policies have more adjacent to reserves; many of these to the species. positive attitudes and relationships with hunting areas are geographically linked both the park manager and the PA as a to national parks and reserves, Within its current range, the African whole (Huzzah 2013, pp. 1, 8). This providing wildlife corridors and buffer lion exists in 10 stronghold populations open approach to PA management zones (Chardonnet et al. 2010, p. 34; containing approximately 24,000 lions reflects a trend in recent years to bring Newmark 2008, p. 321). In the past, (70 percent of the current African lion in local communities to assist in the government and private land owners population), 19,000 of which are in management of PAs (Lindsey et al. were the primary beneficiaries of the protected areas, and in 7 potential 2012b, p. 53). Wildlife management revenue gained; however, a portion of stronghold populations containing programs run by local communities are the revenue derived from hunting is another 4,000 lions. Reports from the defined by two goals: conserving reportedly now being distributed to IUCN Species Survival Commission Cat wildlife and providing economic aids to local communities, creating a value for Specialist Group (IUN 2006a, b) the community (Bandyopadhyay et al. lions that encourages their conservation characterize the population as 2010, p. 5). NGOs are also assisting in (Barnett & Patterson 2005, p. iv). increasing in 3 of those strongholds, as protecting lions. Intervention by NGOs Revenue from trophy hunting is stable in 6 of the strongholds, and as often takes the form of interacting with purported to create: (1) Incentives for decreasing in 1 stronghold. Most lion the local community (Winterbach et al. countries to conserve large tracts of populations in protected areas of 2010, p. 98). For example, Lion prime habitat; and (2) funding for park southern and eastern Africa have been Guardians, which operates in Kenya, and reserve management, anti-poaching, essentially stable over the last three has shown great success with its Lion and security activities. Because habitat decades (Ray et al. 2005, p. 67). In Guard program. Lion Guardians loss has been identified as one of the contrast to the stronghold or potential educates local young men who monitor primary threats to lion populations, it is stronghold populations, other African and track lion movement and warn notable that trophy hunting has lion populations, containing a total of herders of lion presence in the area, provided lion range states incentives to more than 6,000 individuals, have a thereby mitigating or preventing set land aside for hunting throughout very high risk of local extinction (Reggio possible lion-yhuman conflict (Hazzah Africa, and the land set aside exceeds et al. 2013, p. 33. During the 2005–2006 et al. 2014, p. 853; Lion Guardians 2013, the total area of the national parks, African lion workshops, lion experts p. 7; Lion Guardians 2012, p. 3). accounting for approximately half of the characterized lion populations in 36 (42 Outreach to tribal elders has amount of viable lion habitat percent) of the 86 LCUs as decreasing. successfully helped elders to dissuade (Chardonnet et al. 2010, p. 34; Packer et In extensive surveys recently conducted young men from killing lions for al. 2006, pp. 9–10). within 15 of the 20 LCUs in western and

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central Africa, Henschel et al. (2010, DPS, we will list the DPS rather than the We propose to add a 4(d) (special) entire) were able to confirm lion entire taxonomic species or subspecies. rule for the African lion (Panthera leo presence in only four. The work of We found the African lion to be in leo) at 50 CFR 17.40(n). This 4(d) rule Packer et al. (2013) suggests future danger of extinction within the would maintain all of the prohibitions declines within a number of protected foreseeable future throughout all of its and exceptions codified in 50 CFR 17.31 areas. Craigie et al. (2010, entire) range. Therefore, no portions of the and 17.32 and would supersede with provide evidence of declining large species’ range are ‘‘significant’’ as regard to African lion the import mammal populations in Africa’s defined in our SPR policy and no exemption found in 50 CFR 17.8 for protected areas, indicating that additional SPR analysis is required. threatened wildlife listed in Appendix II protected areas in Africa have generally Proposed 4(d) Rule of CITES, such that a threatened species failed to mitigate threats to large import permit under 50 CFR 17.32 mammal populations, including African The purposes of the ESA are to would be required for the importation of lion. Although Craigie et al. (2010, p. provide a means whereby the all African lion specimens. Through the 2,225) found large regional differences ecosystems upon which endangered promulgation of the proposed 4(d) rule, (from large declines in western Africa to species and threatened species depend the presumption of legality provided positive rates of change in southern may be conserved, to provide a program under Section 9(c)(2) of the Act for the Africa), they found overall populations for the conservation of such endangered otherwise lawful importation of wildlife decreased steadily from 1970 to 2005. species and threatened species, and to listed in Appendix II of CITES that is take such steps as may be appropriate to The best available scientific and not an endangered species listed achieve the purposes of the treaties and pursuant to section 4 of the Act would commercial information leads us to conventions set forth in the ESA. When conclude that the African lion is in not apply to this subspecies. Thus, a species is listed as endangered, certain under the proposed 4(d) rule, all danger of extinction within the actions are prohibited under section 9 of otherwise prohibited activities, foreseeable future throughout all of its the ESA, as specified in 50 CFR 17.21. including all imports of African lion range. Accordingly, we find that listing These include, among others, specimens, would require prior is warranted and we propose to list it as prohibitions on take within the United authorization or permits under the Act. a threatened species throughout its States, within the territorial seas of the Under our regulations, permits or range, wherever found. United States, or upon the high seas; authorization to carry out an otherwise import; export; and shipment in Significant Portion of Its Range prohibited activity could be issued for interstate or foreign commerce in the scientific purposes, the enhancement of Under the Act and our implementing course of a commercial activity. regulations, a species may warrant The ESA does not specify particular propagation or survival of the species, listing if it is endangered or threatened prohibitions and exceptions to those economic hardship, zoological throughout all or a significant portion of prohibitions for threatened species. exhibitions, educational purposes, or its range. The term ‘‘species’’ includes Instead, under section 4(d) of the ESA, special purposes consistent with the ‘‘any subspecies of fish or wildlife or the Secretary, as well as the Secretary of purposes of the Act. Applications for plants, and any distinct population Commerce depending on the species, these activities are available from segment [DPS] of any species of was given the discretion to issue such http://www.fws.gov/forms/3-200-37.pdf. vertebrate fish or wildlife which regulations as deemed necessary and The intent of this proposed 4(d) rule interbreeds when mature.’’ We advisable to provide for the is to provide for the conservation of the published a final policy interpreting the conservation of such species. The African lion consistent with the phrase ‘‘Significant Portion of its Secretary also has the discretion to purposes of the Act. Under the proposed Range’’ (SPR) (79 FR 37578, July 1, prohibit by regulation with respect to 4(d) rule, the prohibitions would, in 2014). The final policy states that (1) if any threatened species any act part, make it illegal for any person a species is found to be endangered or prohibited under section 9(a)(1) of the subject to the jurisdiction of the United threatened throughout a significant ESA. Exercising this discretion, the States to ‘‘take’’ (includes harass, harm, portion of its range, the entire species is Service has developed general pursue, hunt, shoot, wound, kill, trap, listed as endangered or threatened, prohibitions (50 CFR 17.31) and capture, or to attempt any of these) respectively, and the Act’s protections exceptions to those prohibitions (50 within the United States or upon the apply to all individuals of the species CFR 17.32) under the ESA that apply to high seas; import or export; deliver, wherever found; (2) a portion of the most threatened species. Under 50 CFR receive, carry, transport, or ship in range of a species is ‘‘significant’’ if the 17.32, permits may be issued to allow interstate or foreign commerce, by any species is not currently endangered or persons to engage in otherwise means whatsoever, in the course of threatened throughout all of its range, prohibited acts for certain purposes. commercial activity; or sell or offer for but the portion’s contribution to the Under section 4(d) of the ESA, the sale in interstate or foreign commerce viability of the species is so important Secretary, who has delegated this any lion specimens. It would also be that, without the members in that authority to the Service, may also illegal to possess, sell, deliver, carry, portion, the species would be in danger develop specific prohibitions and transport, or ship any such wildlife that of extinction, or likely to become so in exceptions tailored to the particular has been taken in violation of the Act. the foreseeable future, throughout all of conservation needs of a threatened We believe that these protections, its range; (3) the range of a species is species. In such cases, the Service issues including the requirement for an import considered to be the general a 4 (d) rule that may include some or all permit for all African lion specimens, geographical area within which that of the prohibitions and authorizations will support and encourage species can be found at the time FWS set out in 50 CFR 17.31 and 17.32 but conservation actions for the African lion or NMFS makes any particular status which also may be more or less and require that permitted activities determination; and (4) if a vertebrate restrictive than the general provisions at involving lions are carried out in a species is endangered or threatened 50 CFR 17.31 and 17.32. For the African manner that is consistent with the throughout an SPR, and the population lion, the Service has determined that a purposes of the Act and our in that significant portion is a valid 4(d) rule is appropriate. implementing regulations.

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In connection with this proposed 4(d) on scientifically sound data and are Threatened Wildlife. This rule, if rule, the Service notes that the African being implemented to address the adopted, would also establish a 4(d) rule lion is listed in Appendix II of CITES, threats that are facing lions within that for the African lion, which implements and thus can be imported into the U.S. country. all of the prohibitions and exceptions pursuant to Section 9(c)(2) of the Act Such management plans would be under 50 CFR 17.31 and 17.32 and and upon presentation of a proper expected to address, but are not limited requires a threatened species import CITES export permit from the country of to, evaluating population levels and permit under 50 CFR 17.32 for the origin. Section 9(c)(2) of the Act trends; the biological needs of the importation of all African lion provides that the otherwise lawful species; quotas; management practices; specimens. Under the proposed 4(d) importation of wildlife that is not an legal protection; local community rule, the import exemption found in 50 endangered species listed pursuant to involvement; and use of hunting fees for CFR 17.8 for threatened wildlife listed section 4 of the Act, but that is listed in conservation. In evaluating these in Appendix II of CITES would not Appendix II of CITES, shall be factors, we will work closely with the apply to this subspecies. Through the presumed to be in compliance with range countries and interested parties to promulgation of the proposed 4(d) rule, provisions of the Act and implementing obtain the best available scientific and the presumption of legality provided regulations. While there has been commercial data. By allowing entry into under Section 9(c)(2) of the Act for the question as to whether this provision of the United States of African lion otherwise lawful importation of wildlife the Act might automatically require trophies from range countries that have listed in Appendix II of CITES that is allowing the importation of a species scientifically based management plans, not an endangered species listed that is both listed as threatened and in the range countries would be pursuant to section 4 of the Act would Appendix II, and preclude the issuance encouraged to adopt and financially not apply to this subspecies. (See: of more restrictive 4 (d) rules covering support the sustainable management of Proposed Special Rule section). lions that benefits both the species and importation, the Service has concluded Available Conservation Measures that such 4 (d) rules may be issued to local communities. In addition to provide for the conservation of the addressing the biological needs of the Conservation measures provided to involved species. Section 9(c)(2) does subspecies, a scientifically based species listed as endangered or not expressly refer to threatened species management plan would provide threatened under the Act include or prevent the issuance of appropriate 4 economic incentives for local recognition of conservation status, requirements for Federal protection, and (d) rules and could not logically have communities to protect and expand prohibitions against certain practices. been intended to allow the addition of African lion habitat. Recognition through listing encourages a species to an appendix of an As stated, anyone wishing to conduct any otherwise prohibited activity, such and results in public awareness and international convention to override the as interstate commerce or imports, must conservation actions by Federal and needs of U.S. law, where there is first obtain a permit under the current State governments in the United States, reliable evidence to affect the permitting regulations found at 50 CFR foreign governments, private agencies presumption of validity. Finally, the 13 and 50 CFR 17. As will all permits, and groups, and individuals. term ‘‘presumed’’ implies that the the individual requesting authorization Section 7(a) of the Act, as amended, established presumption is rebuttable to carry out an otherwise prohibited and as implemented by regulations at 50 under certain circumstances, including activity under the Act must submit a CFR part 402, requires Federal agencies through the promulgation of a protective permit application to the Service with to evaluate their actions that are to be regulation pursuant to section 4(d) of specific information concerning the conducted within the United States or the Act. proposed activity and the benefits/ upon the high seas, with respect to any In the case of the African lion, there impacts of the activity on the species. In species that is proposed to be listed or are substantive grounds on which to some cases, such as imports of sport- is listed as endangered or threatened. challenge the presumption. For the hunted trophies, it is not always Because the African lion is not native to import of sport-hunted trophies, while possible for the applicant to provide all the United States, no critical habitat is there is evidence that many of the range of the necessary information needed by being proposed for designation with this countries are implementing lion the Service to make a positive rule. Regulations implementing the management plans, we want to determination under the Act to interagency cooperation provision of the encourage and support efforts by these authorize the activity. For the import of Act are codified at 50 CFR part 402. countries to develop plans that are sport-hunted trophies, it is typical for Section 7(a)(2) of the Act requires based on sound scientific information. the Service to consult with the range Federal agencies to ensure that activities As noted, the proposed 4(d) rule for country and other interested parties to they authorize, fund, or carry out are not African lion would provide for the obtain the necessary information. To likely to jeopardize the continued importation into the United States of date, the Service typically has made the existence of a listed species or to trophies taken legally in range countries required findings on sport-hunted destroy or adversely modify its critical upon the issuance of a threatened trophy imports on a country-wide basis, habitat. If a proposed Federal action species import permit. While the although individual import permits are may adversely affect a listed species, the Service cannot control hunting of issued for each applicant. While the responsible Federal agency must enter foreign species such as African lion, we Service encourages the submission of into formal consultation with the can regulate their importation and information from individual applicants, Service. Currently, with respect to the thereby require that U.S. imports of we would primarily rely on information African lion, no Federal activities are sport-hunted African lion trophy from other sources when making a known that would require consultation. specimens are obtained in a manner that permitting decision. Section 8(a) of the Act authorizes the is consistent with the purposes of the provision of limited financial assistance Act and the conservation of the Effects of This Rule for the development and management of subspecies in the wild, by allowing This rule, if made final, would revise programs that the Secretary of the importation from range countries that 50 CFR 17.11(h) to add the African lion Interior determines to be necessary or have management plans that are based to the List of Endangered and useful for the conservation of

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endangered or threatened species in possess, sell, deliver, carry, transport, or Branch of Foreign Species (see FOR foreign countries. Sections 8(b) and 8(c) ship any such wildlife that has been FURTHER INFORMATION CONTACT). of the Act authorize the Secretary to taken in violation of the Act. Permits Authors encourage conservation programs for may be issued to carry out otherwise foreign listed species, and to provide prohibited activities involving The primary authors of this proposed assistance for such programs, in the threatened wildlife species under rule are staff of the Branch of Foreign form of personnel and the training of certain circumstances. Certain Species, Ecological Services, U.S. Fish personnel. exceptions apply to agents of the and Wildlife Service. Section 9 of the Act and its Service and State conservation agencies. implementing regulations at 50 CFR Proposed Regulation Promulgation Required Determinations 17.31 set forth a series of general For the reasons described in the prohibitions that apply to all threatened National Environmental Policy Act (42 preamble, we propose to amend part 17, wildlife, except where a 4(d) rule U.S.C. 4321 et seq.) subchapter B of chapter I, title 50 of the applies, in which case the 4(d) rule will We have determined that we do not Code of Federal Regulations, as follows: contain all the applicable prohibitions need to prepare an environmental and exceptions. If the 4(d) rule is assessment, as defined under the PART 17—[AMENDED] adopted as proposed, these prohibitions authority of the National Environmental ■ 1. The authority citation for part 17 would apply to the African lion. These Policy Act of 1969, in connection with continues to read as follows: prohibitions, in part, make it illegal for regulations adopted under section 4(a) any person subject to the jurisdiction of of the Act. We published a notice Authority: 16 U.S.C. 1361–1407; 1531– the United States to ‘‘take’’ (includes outlining our reasons for this 1544; and 4201–4245; unless otherwise harass, harm, pursue, hunt, shoot, determination in the Federal Register noted. wound, kill, trap, capture, or to attempt on October 25, 1983 (48 FR 49244). ■ 2. In § 17.11(h), the List of Endangered any of these) within the United States or and Threatened Wildlife, add an entry References Cited upon the high seas; import or export; for ‘‘Lion, African’’ under Mammals to deliver, receive, carry, transport, or ship A list of all references cited in this read as follows: in interstate or foreign commerce, by document is available at http:// any means whatsoever, in the course of www.regulations.gov at Docket No. § 17.11 Endangered and threatened commercial activity; or sell or offer for FWS–R9–ES–2012–0025, or upon wildlife. sale in interstate or foreign commerce request from the U.S. Fish and Wildlife * * * * * any lion specimens. It also is illegal to Service, Endangered Species Program, (h) * * *

Species Vertebrate population where When Critical Special Historic range endangered or Status listed habitat rules Common name Scientific name threatened

MAMMALS

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

*******

* * * * * (2) The import exemption found in (3) All applicable provisions of 50 ■ 3. Amend § 17.40 by adding paragraph § 17.8 of this part for threatened wildlife CFR parts 13, 14, 17, and 23 must be (n) to read as follows: listed in Appendix II of the Convention met. on International Trade in Endangered * * * * * § 17.40 Special rules—mammals. Species of Wild Fauna and Flora Dated: October 20, 2014. * * * * * (CITES) does not apply to this (n) African lion (Panthera leo leo). subspecies. A threatened species import Stephen Guertin, (1) General requirements. All permit under § 17.32 of this part is Director, Fish and Wildlife Service. prohibitions and provisions of §§ 17.31 required for the importation of all [FR Doc. 2014–25731 Filed 10–28–14; 8:45 am] and 17.32 of this part apply to this African lion specimens. BILLING CODE 4310–55–P subspecies.

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