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M S AMMALIAN PECIES No. 762, pp. 1–11, 3 figs. Panthera leo. BySarah K. Haas, Virginia Hayssen, and Paul R. Krausman Published 15 July 2005 by the American Society of Mammalogists Panthera leo (Linnaeus, 1758) [Felis leo] var. somaliensis Noack, 1891:120. Type locality ‘‘So- mali-Halbinsel.’’ Lion Felis leo kamptzi Matschie, 1900:92. Type locality ‘‘Yoko am ob- erenSanaga,’’5Yoko,UpperSanagaRiver,Cameroon. Felis leo Linnaeus, 1758:41. Type locality ‘‘Africa,’’ restricted to Felis leo goojrattensis Matschie, 1900:94. Incorrect subsequent ‘‘Constantine, Algeria’’ by J. A. Allen (1924:222–223) and spellingofFelis leo goojratensisSmee. based on Cuvier’sLion deBarbarie (GeoffreySt.Hilaireand Felis leo massaicus Neumann, 1900:550–551. Type locality ‘‘Ki- Cuvier1819). baya,Massai-Land.’’ [Felis leo] barbaricus Meyer, 1826:6. Type locality‘‘Barbarische’’ 5Barbary,NorthAfrica,videG. M.Allen(1939:244). FelisleosabakiensisLo¨nnberg,1910:22.Typelocality‘‘Kilimand- [Felis leo] persicus Meyer, 1826:6. Type locality ‘‘Persische’’ 5 jaro:... Kibonotointhecultavatedzone,1,500m.abovethe sea’’ or ‘‘in the plains NW. ofKilimandjaro,Leitokitok;...’’ Persia,videEllermanandMorrison-Scott(1951:319). restrictedtoKibonoto,Kilimanjaro,byJ.A.Allen(1924:223). [Felis leo] senegalensis Meyer, 1826:6. Type locality ‘‘Senegalis- che’’ 5 Senegal, vide G. M. Allen (1939:244); not Felis se- FelisleorooseveltiHeller,1913:2.Typelocality‘‘highlandsofAb- yssinianearAddisAbaba.’’Typelocalityexplicitlyquestioned negalensisLesson,1839:plate10,orFelisjubatasenegalen- and stated more broadly as ‘‘Abyssinia’’ by Hollister (1918: sisBlainville,1843. 165). Leo asiaticusBrehm,1829:638. Notypelocalityspecified. FelisleonyanzaeHeller,1913:4.Typelocality‘‘Kampala,’’Ugan- Leo africanusBrehm,1829:638. Notypelocalityspecified. FelisleoabarbarusFischer,1829:197.Notypelocalityspecified; da. Felis leo bleyenberghi Lo¨nnberg, 1914:273–274. Type locality based on Cuvier’sLion deBarbarie (GeoffreySt.Hilaireand ‘‘Katanga,’’BelgiumCongo. Cuvier1819). Felis leobsenegalensisFischer,1829:197.Notypelocalityspec- Leo leo leo:J. A.Allen,1924:223. Namecombination. LeoleoazandicusJ.A.Allen,1924:224.Typelocality‘‘Vankerck- ified;basedonCuvierandGeoffroy’s(1819)LionduSenegal. hovenville,northeasternBelgianCongo.’’ Preoccupied by [Felis leo]senegalensisMeyer,1826:6(Hol- lister, 1910:123). Not Felis senegalensis Lesson, 1839:plate 10, orFelis jubata senegalensisBlainville,1843. FelisleobengalensisBennett,1829:1.Typelocality‘‘Bengal’’;cor- rected to ‘‘Hurriana,’’ 5 Harriana, northern India by Blyth (1863).PreoccupiedbyFelis bengalensisKerr,1792:151. Felis leo e capensisFischer, 1830:565 [page mislabeled365].No type locality specified; based on Griffith’s (1827) ‘‘South Af- rican Lion.’’ Preoccupied by Felis capensis Forster, 1781:4, plate1. Felis leo var. goojratensis Smee, 1833(1834):165, 174. Type locality‘‘Guzerat’’5Gujerat,India;statedas‘‘Ahmedabad’’by Pocock(1930:660). Leo asiaticus:Jardine,1834:121. Namecombination. [?] guzcratensis Wagner, 1841:461. Incorrect subsequent spelling ofgoojratensisSmee, videPocock(1930:660). Felis (Leo) gougeratensis H. Smith, 1842:178 and plate XI. In- correctsubsequentspellingofFelis leo goojratensisSmee. Felis(Leo)melanochaitusH.Smith,1842:177.Typelocality‘‘The Cape’’videShortridge(1934:77)or‘‘CapeofGoodHope’’vide Roberts (1951:190); correct original spelling, by selection (Hollister1910:123). Felis(Leo)melanochoetusH.Smith,1842:plateX.Incorrectorig- inalspellingofFelis melanochaitusH. Smith. Leo gambianus Gray, 1843:40. Type locality ‘‘W. Africa, interior ofGambia’’;renamingofFelisleosenegalensisFischer,1829: 197; nomen nudum. F[elis].leonubicusBlainville,1843:58.Notypelocalityspecified; statedas‘‘Nubia’’by G.M.Allen(1939:243). Leo indicus Blainville, 1843:196. No type locality specified; pre- sumably India (Ellerman and Morrison-Scott 1951:319; Po- cock1930). Felis leo nigra Loche, 1858:7. Type locality not known, given as ‘‘Constantine,Algerien’’videHemmer(1974:230). Tigris leo:Severtzoff,1858:388. Namecombination. L[eo].gozeratensisBrehm,1863:59.Incorrectsubsequentspelling ofFelis leo goojratensisSmee. LeonobilisGray,1867:263.RenamingofFelisleoLinnaeus,1758; nomen nudum. L[eo].goorgrattensisGray,1867:263.Incorrectsubsequentspelling ofFelis leo goojratensisSmee. Leo guzeratensis Fitzinger, 1868:443. Incorrect subsequent spell- FIG. 1. Photograph of an adult male Panthera leo, Kenya, ingofFelis leo goojratensisSmee. Africa.UsedwithpermissionofthephotographerS. P. Goyal. 2 MAMMALIANSPECIES 762—Panthera leo Leo leo hollisteri J. A. Allen, 1924:228. Type locality ‘‘Lime Springs,Sotik,BritishEastAfrica.’’ Leo leo krugeri Roberts, 1929:91. Type locality ‘‘Sabi Game Re- serve(KrugerNationalPark),’’southernAfrica. Panthera leo: Pocock, 1930:660. First use of current name combination. LeoleovernayiRoberts,1945:65.Typelocality‘‘MatapaPan,cen- tralKalahari,’’southernAfrica. Felis leo blevenberghi Jobaert, 1954:479. Incorrect subsequent spellingofFelis leo bleyenberghiLo¨nnberg,1914. Leo maculatesHeuvelmans,1955:443. Nomennudem. Panthera leo webbiensis Zukowsky, 1964:270. Type locality ‘‘So- malilande.’’ CONTEXTANDCONTENT.Genericcontextgivenabove. Hemmer(1974),Pocock(1930),andMeesteretal.(1986)provided several synonyms without sufficient citation information for vali- dation. These are listed above and in the ‘‘Literature Cited’’ with as much information as we could ascertain. The lion in NorthAf- rica, Panthera leo leo (Linnaeus), is extinct; barbaricus (Meyer), barbarus (Fischer), leo (Allen), and nigra (Loche) are synonyms. P. leohas7 extantsubspecies(Hemmer1974). P. l. azandicus (Allen, 1924:224), see above; massaicus (Neu- mann)partimandsomaliensis(Noack)partimaresynonyms. P. l. bleyenberghi (Lo¨nnberg, 1914:273–274), see above; blevenberghi(Jobaert)isa synonym. P. l. krugeri(Roberts,1929:91),seeabove;vernayi(Roberts)isa synonym. P. l. melanochaitus(Smith, 1842:177), seeabove;melanochaitus (Smith)andmelanochætus(Smith)aresynonyms. P. l. nubicus (Blainville, 1843:58), see above; hollisteri (Allen), massaicus (Neumann) partim, nyanzæ (Heller), roosevelti (Heller),sabakiensis(Lo¨nnberg),somaliensis(Noack)partim, andwebbiensisZukowskyaresynonyms. P. l. persica (Meyer, 1826:6), see above; asiaticus (Brehm), ben- galensis (Bennett), goojratensis (Smee), goojrattensis (Mat- schie), goorgrattensis (Gray), gougeratensis(Smith),gozera- tensis (Brehm), guzcratensis (Wagner), guzeratensis (Fitzin- ger),andindicus(Blainville)aresynonyms. P. l. senegalensis (Meyer, 1826:6) see above; gambianus (Gray), kamptzi(Matschie),andnobilis(Gray)aresynonyms. DIAGNOSIS.The lion (Fig. 1)isthe 2nd largestspeciesof Felidae, slightly smaller than the tiger, Panthera tigris (Maza´k 1981). Skull of P. leo (Fig. 2) is similar in appearance to that of the leopard (Panthera pardus) but is nearly twice the size.P. leo isdigitigradewithsharpretractileclaws;broadface,roundedears, relatively short neck; and well-proportioned, muscular body more drawn inatbellycomparedwiththetiger(Rudnai1973a). GENERAL CHARACTERS. Body is unicolor, lacking ro- FIG.2. Dorsal,ventral,andlateralviewsofcraniumandlat- settes; color is light tawny, white on abdomen and inner side of eral view of mandible of adult male Panthera leo, Chicago Field legs,withblackonbackofears(Smithers1971).Insizeandgen- Museumno. 20757. Greatestlengthofcraniumis330 mm. eral appearance, Asian and African subspecies are similar; the most obvious physical difference between Asian and Africansub- species is a longitudinalfold ofskin running alongbelly ofAsian Smutsetal.1980).InKrugerNationalPark,SouthAfrica,4males subspecies (Pocock 1930). Variations between African subspecies .4 years old averaged 181 kg, 25 males 2–4 years old averaged have been noted between and within geographic regions on the 146 kg, and large male cubs averaged 77 kg (Smuts 1976). The basisofsize, coatcolor,anddevelopmentandcolorofmane,with largestmaleinKrugerNationalParkweighed225kg(Smuts1982), thosefromsouthernpartsofcontinentbeinglargeronaveragethan butamalenearMountKenyaweighed272kg(NowellandJackson thosefromeasternparts(TurnerandAntoń1997). 1996). Males weigh 20–27% more than females. For females, Pelage varies across populations and between and within lengthofheadandbodyis1,400–1,750mm,lengthoftailis700– neighboringprides(Smutsetal.1978).Colorvariesfromlightbuff 1,000 mm, height at shoulder averages 1,070 mm, and mass is andsilverygraytoyellowishredanddarkbrown;whitelionshave 120–182kg(NowakandParadiso1983).InKrugerNationalPark, been reported from KrugerNationalParkandUmfoloziGameRe- SouthAfrica,25females.4yearsoldaveraged126kg,11females servein SouthAfrica(McBride1977;Smuts1982).Coatcolorre- 2–4yearsoldaveraged103kg,andlargefemalecubsaveraged60 sultsfromacombinationofshortersandy-yellowhairs,mixedwith kg(Smuts1976).Thelargestfemaleweighed152kg(Smuts1982). and overlain by longer, black guard hairs (Rudnai 1973b). Juve- Asian subspeciesisslightlysmallerthanAfricansubspecies; nilesareheavilyspotted(Rosevear1974). adult males weigh 160–190 kg and females weigh 110–120 kg Manesofadultmalesvaryincolor(fromtawnytoblack)and (Nowelland Jackson 1996).Recordtotallengthsformalesareca. size varies among populations and age classes (Rosevear 1974). 3,300and2,920mmforAfricanandAsiansubspecies,respectively Manegrowthisinfluencedbytestosterone,andmanecolorisusu- (Nowelland Jackson1996). allylightinyounganimalsand darkenswithage(Schaller1972). Panthera leo has a muscular, deep-chested body with a Head is rounded and bears prominent whiskers. For males, rounded and shortened head and reduced dentition (Rudnai lengthofheadandbodyrangesfrom1,700to2,500mm,lengthof 1973b). Skull is wide and high compared with its length. Orbitis tailis900–1,050mm,heightatshoulderisca.1,230mm(Nowak large,anditsposteriorborderisslightlyanteriortomiddleofskull. andParadiso1983),andmassis150–225kg(average5188kg— Postorbital processes are broad, blunt, and ligulate. Upper profile 762—Panthera leo MAMMALIANSPECIES 3 parks are a refuge for free-ranging lions and within these areas their numbers increase (Orford et al. 1988). Countries in which lions are stillrelatively widespreadincludeBotswana,CentralAf- ricanRepublic,Ethiopia,Kenya,Tanzania,Zaire,andZambia.Sta- tus in Angola, Mozambique, Somalia, and Sudan is uncertain but groups may be patchy and widespread (Fagotto 1985). Lions sur- vivedinthedesertontheedgeofNiger’sAirMountainsuptoca. 1910 (Rosevear 1974). Lions may range into snowy mountainous regions of eastern Africa up to 3,600 mon Mt. Elgon, Mt.Kenya, andMt.Ruwenzoriandover4,200minEthiopia’sBaleMountains (Nowelland Jackson1996). FOSSILRECORD.Theearliestfossilrecordofthelionap- pears during the Late Pliocene at Laetoli in Tanzania ca. 3.5 3 106yearsago(TurnerandAntoń1997).The1stappearanceofthe lion in Europe occurred ca. 0.9 3 106 years ago at the French Mediterranean site of Vallonnet, and those European lions were larger than extant lions (Turner and Antoń 1997). Fossil remains of the cave lion (P. leo cf. spelaea) were recovered from northern Israel dating to the Middle Pleistocene (Tchernov and Tsoukala 1997).OtherareasintheOldWorldwherefossilremainshavebeen FIG. 3. Geographic distribution of Panthera leo in Africa found include the upper beds of Olduvai Gorge in easternAfrica, and India (modified from Hemmer 1974; Nowell and Jackson SwartkransandKromdraaiinSouthAfrica,andtheMosbachgrav- 1996): 1, P. l. azandicus; 2, P. l. bleyenberghi; 3, P. l. krugeri elsinGermany(Hemmer1979a).NumerousfindsinFrance,Eng- andP. l. melanochaitus; 4,P. l. nubicus;5,P. l. persica;6,P.l. land, and Germany indicate that the lion was common during the senegalensis. HolsteinandIlfordinterglacials.DuringtheLatePleistocene,lions spread over North America into Central America (Hemmer 1978; Seymour1983). of skull in lateral view is convex; muzzle is steeply sloped with VariouspopulationsknownfromfossilsbynamesP.atrox,P. nasal orifice set low. Temporal ridges meet on summit of sagittal fossilis,P.spelaea,andP.vereshchaginiarenowregardedascon- crest, are short, and are restricted to posterior portion of crown specific withP. leo(Burger et al.2004;Harington1977;Hemmer (Pocock 1951). Sagittal crest, mastoid, and paroccipitalprocesses 1976, 1979a). In Europe and northern Asia, lion populationsmay areprominent(Smithers1971).Ca.50%ofAsiansubspecieshave havebecomeisolatedduetowidespreadicesheets.Americanand bifurcatedinfraorbitalforamina,whereasAfricanlionshaveonly1 cavelionsbecameextinctca.10,000 yearsago(Harington1996). foramenoneitherside(O’Brienetal.1987a;Pocock1939).Base- lineoflowerjawisslightlyconvex.Carnassialsarehighlysectorial, FORMANDFUNCTION.Skullhasshortjaws.Backward- andcanineshaveafinelongitudinalgroove(Ewer1973).Averages curved horny papillae cover surface of tongue (Pocock 1951). Al- orrangesforcranialmeasurements(inmm)areasfollows:greatest thoughthesenseofsmelliswelldeveloped,visionandhearingare length of skull, 250–460; condylobasal length, 316–345; basal ofgreaterimportanceinlocatingprey(Macdonald1984).Themane length, 284; zygomatic breadth, 229; interorbital breadth, 69.5; servesasprotectionduringintraspecificfighting,asignpostofgen- braincase breadth, 106 (Rosevear 1974; Van Valkenburgh 1996). derdistinguishableatadistance,anindicatorofindividualfitness, Meanlengthof48skullswas277.1(SD522.9—VanValkenburge and insulation for the neck (Caputo 2002), but may have evolved and Ruff 1987). Ranges of cranial measurements for 5 males and recently(320,000–190,000yearsago—Yamaguchietal.2004). 6 females, respectively, from eastern Africa are: greatest length, Lions have an incompletely ossified hyoid apparatus with an 353–401,293–315;condylobasallength,316–345,264–280;man- elasticligamentmeasuring15cmwhenrelaxedand22.5cmwhen dibular length, 235–260, 196–212; width of mastoid, 133–144, stretched,allowinglionstomovethelarynxfrompalateandlength- 112–120 (Hollister 1918). Similar cranial measurements are pre- en pharyngeal passage (Hast 1989). This elastic ligament makes sentedby Roberts(1945). roaringpossible(Owen1834; Pocock1916). Panthera leo is unique in having a horny spur that is sepa- Vertebral formula is 7 C, 13 T, 7 L, 3 S; number of caudal rated from last caudal vertebra and that is covered by a tuft of vertebrae not reported. Dental formula is i 3/3, c 1/1, p 3/2, m blackhairattipoftail(Rudnai1973b).Peculiaritiesofhairgrowth 1/1, total 30 (Groves 1982; Rosevear 1974; Smuts et al. 1978). alsodistinguishlionsfromotherfelids.Pelageiscomposedofrel- Maxillary canines are ca. 43–52 mm tall, elongated, posteriorly ativelysparse,fineunderfur6–8mmlongandofflattish-sectioned recurved, conical, and somewhat longer than lower canines. P4 subbristles 10–14 mm long (Rosevear 1974). Lions have a large carnassials measure 38 mm long (Parker 1982; Rosevear 1974; patchofreversehairgrowingforward,startingwithavortexabove Smutsetal.1978).Lowercarnassialhasadeep,narrowcarnassial lumbar vertebrae and continuing forward up to middorsal region notch and upper carnassial has a large protocone. Incisors are where it forms a small transverse crest at interface withopposite- small, spatulate, and arranged inacompacttransverserow;upper growing hair and extends laterally about one-third down side of lateral incisors are larger than more mesial incisors (Rosevear body (Rudnai1973b). 1974). Occiput is inclined posteriorly to upper toothrow and par- amastoidsandparaoccipitalsaresmallandseparate(Martin1989). DISTRIBUTION. Ancestors of modern lions occurred Lionshave sebaceousglands around chin,lips,cheeks,mys- throughout Europe and the Americas until Late Pleistocene(Har- tacial and genal whiskers, above tail, and between toes (Schaller ington1996;KurteńandAnderson1980;Vereshchagin1971).Ca. 1972). Females have 2 pairs of mammae, rarely 3 (Ewer 1973). 2,000 years ago, lions became extinct in eastern Europe with the Anal glands of lions are ca. 25 mm long with ducts that open on development of dense forests (Guggisberg 1961, 1975). Until ca. either side of and close to anus (Ewer 1973). Penis is relatively 1850, lions were distributed across the Indian subcontinent and shortand baculumis6.0–9.5 mm long(Didier1949). werefoundinpresent-daystatesofGujarat,Haryana,MadhyaPra- Lions drink after every meal when water is available, and desh, Punjab, Rajasthan, and Uttar Pradesh (Chellam and John- drinkingcontributessignificantlytowaterinfluxratesinhotweath- singh1993).Byearly1900s,theAsiansubspecieswasdrastically er(Eloff1973;Greenetal.1984).Drinkingcomprisesca.40%of reducedandtodayisconfinedtotheGirSanctuaryofGujuratState thetotalwaterinfluxinlionsinEtoshaNationalPark(Clarkeand in western India, where it was initially protected by theNawabof Berry 1992). Lions can obtain water from prey and plants to a Junagadhin hisprivatehuntinggrounds(Kinnear1920). limited extent and can survive in very arid environments (Eloff AfricansubspeciesoccuralongthesouthernfringeoftheSa- 1973; Hanby et al. 1995). Asian lions frequently hunt within riv- harafromwesternAfricatoSudanandSomaliathensouththrough erineforestsneara watersource(Chellamand Johnsingh1993). most of Africa excluding the Congo forest region (Fig. 3). Lions havedisappearedinnorthernAfricaandtheCaperegion.Southof ONTOGENY AND REPRODUCTION. Lions have no theSahara,lionsarerareinwesternAfricabutsurviveinprotected fixed breeding season although birth peakshavebeenrecordedin areas of eastern and southern Africa (Rosevear 1974). African KrugerandSerengetiNationalParks(February–AprilandMarch– 4 MAMMALIANSPECIES 762—Panthera leo July,respectively)inrelationtoseasonalweatherpatternsandprey Largermammals,suchasyoungAfricanelephants(Loxodontaaf- availability (Packer et al. 1990; Smuts et al. 1978). Lion produc- ricana), buffalo (Syncerus caffer), eland (Tragelaphus), giraffe tivity(measuredinthenumberofsurvivingcubs)islimitedbyfood. (Giraffacamelopardalis),andkudu(Tragelaphusstrepsiceros)also Four-cub litters survive best where prey is most abundant, and maybecaptured(Pienaar1969;Stander1997).InSerengeti,7prey survival declines as the prey base declines (Packer and Pusey species that account for more than 90% of total meat intake are 1995). Whenthepreybaseisreduced,littersizesaresmaller. buffalo, kongoni (Alcelaphus buselaphus), Thomson’sgazelle(Ga- Femalescome into estrusinresponsetowithin-pridemecha- zella thomsonii), topi (Damaliscus lunatus), warthog (Phacocho- nisms such as estrus of pridemates as well as loss of cubs from erusaethiopicus),wildebeest,andzebra(Equusburchellii—Scheel male infanticide at pride takeovers (Bertram 1975b; Packer and and Packer 1995). In Kafue and Manyara nationalparks of South Pusey1983a;Schaller1972).Estruslasts4–7days,withintervals Africa, buffalo make up 62% of lion kills (Ewer 1973; Schaller betweenperiodslastingfromafewdaystomorethanayear(Rud- 1972). In Kalahari Gemsbok Park, lions survive on smaller prey nai 1973a). During estrus, vaginal smears display flat, nonnucle- suchasgemsbok(Oryxgazella)andporcupines(Hystrix)because ated, cornified cells with a smaller number of cells whose nuclei oftherelativeabsenceoflargeantelope(Eloff1973,1984;Turner are pycnotic (Asdell 1946). Ovulation is induced by copulatory and Antoń 1997). In Kruger NationalPark, male lionspreferbuf- stimuli or interaction with conspecifics during estrus (Schrammet falo,whereasfemalespreyuponsmaller,moreabundantungulates al.1994).Conceptionoccurson4thdayofestruswithaconception suchaswildebeestandzebra(Funstonetal.1998).Basedonnum- rateperestrouscycleof38%(Cooper1942;Rudnai1973b).Cop- berskilledinNairobiNationalPark,kongoni,warthog,wildebeest, ulationoccursupto100timesperdaywithmultiplemales(Davies and zebra provide most of the prey (Rudnai 1974). Asian lions andBoersma1984;Packer1986).Intervalsbetweenmatingsrange mainlypreyuponsmallerspeciesthantheAfricanlions.Themost between4and148minwithanaverageof17min(Rudnai1973a). common prey is chital (Axis axis), weighing ca. 50 kg, although Mean interbirth interval is 20 months if the previous litter sambar deer (Cervus unicolor), weighing ca. 166 kg, may be pre- survives to maturity and 4–6 months if the previous litter is lost ferred(Chellamand Johnsingh1993; Khan1990). (PuseyandPacker1987).Gestationlasts110days.Littersizeav- Lionskillagreaterproportionofmalesandyoungindividuals erages 1–4 (maximum of 6), and postpartum litter sizeisvirtually of several prey species (Viljoen 1997). In SerengetiNationalPark identical to litter size in utero (Cooper 1942; Packer and Pusey in Tanzania, lions prey on cheetah (Acinonyx jubatus) cubs and 1987;Smutsetal.1978).Sexratioatbirthis1:1(Rudnai1973a). may reduce cheetah density in that area (Durant1998). Lionsoc- Femalesgivebirthindensorareasofdensebrushandkeeplitters casionallytakesmallpreysuchasrodents,tortoises,fishinshallow hiddenforatleastamonthwhilecubsarenotverymobileandare ponds,termites,grass,andfruitssuchasthetsamamelon(Citrul- vulnerable to predators (Hanby et al. 1995; Packer and Pusey lus lanatus). They also dig warthogs out of burrows (Ewer 1973; 1997). Van Orsdol 1984). Chimpanzee (Pan troglodytes) remains have Eyes are open at birth or shortly thereafter. Mean weight at been found in the feces of lions from Mahale Mountains National birth is 1,650 g, and cubs gain ca. 106 g per day in the first 4 Park, Tanzania (Tsukahara 1993). Lions are opportunisticscaven- weeksoflife(Hemmer1979b;OftedalandGittleman1989).New- gers that readily displace other predators from their kills (Packer borns are marked with spots that persist on belly and legs until 1986; Schaller 1972). In Serengeti, ca. 40% of food items were adulthoodandmayremainthroughoutlife.Cubsmaybeginwalking scavenged,althoughscavengingislesscommonwherepreyanimals within 2 weeks of birth; eruption of milk teeth begins at 3 weeks arescarce(Packeretal.1990; Stander1992a). ofage,allowingtheyoungtotakesolidfoodsatca.8weeks(Oftedal Each pride has a territory of 20–500 km2 (Van Orsdol et al. andGittleman1989).Milkishighinfat(BenShaul1962;DeWaal 1985). Average range sizes of prides in Africa were 26–226 km2 et al. 2004). Cubs are weaned at ca. 8 months and eruption of but can be considerably larger (Stander 1992b; Van Orsdol et al. permanent teeth begins between 9th and 12th months (Grzimek 1985; Viljoen 1993). Use of space within a territory is related to 1975). the availability of prey (Spong 2002). Variation in vibrissa spot Within a pride, cubs born ,1 year apart constitute a cohort pattern enables study of individual members of prides and group- and are raised communally after 4–6 weeks of age (Hanby et al. level behaviors, and allows estimates of population size (Jhala et 1995; Packer 1986; Packer and Pusey 1983b; Van Orsdol et al. al.1999; Pennycuickand Rudnai1970). 1985). Once cubs can move, mothers bring them out in the open In Serengeti, pride ranges may overlap but each pride main- tojointheprideinanurserycre`chethatwillpersistuntiltheyare tains acore areawheremostactivitiesareconcentratedwithlittle ca.2yearsoldandthemotherresumesmating(Packeretal.1990; interactionwithothergroups(Schaller1972).Serengetilionsmain- Packerand Pusey1997). tain stable territories except during periods of extreme hardship At2 yearsofage,femalelionsareslightlyshorterthanadult (Packer et al. 1990). Studies on the impact of lion removalreveal females, whereas males may be slightly taller than adult females that the opening of a territory (by removal of the current pride) is withinitialmanegrowthevidentaroundneckandonchest(Smuts followed by an influx of competing new lions(Smuts1978b). New et al. 1978). By 3 years of age, young females are difficultto dis- pridesareformedbythedivisionofpreexistingkingroups(Hanby tinguish from adults, and by 4 years, females cannot be distin- and Bygott 1987; Pusey and Packer 1987). Although coalescence guished fromolder femalesonthebasisofbodysize(Smutsetal. ofunrelatedfemalesinaprideisrare,itoccurredduringprolonged 1978). For females, reproduction declines at 11 years of age and drought in Botswana’s Central Kalahari Game Reserve when orig- virtuallystopsat15years;although16-year-oldmalescanproduce inalrangesexpanded(OwensandOwens1984). viable sperm, few old males have access to a pride (Packer et al. Populationdensities(adultsandsubadultsper100km2)range 1988; Smutsetal.1978). from0.08–0.13inSelousGameReserve,0.17inBotswana’sChobe Sexualmaturityoccursatca.24monthsofageforbothsexes, National Park, 1.5–2.0 in Kalahari Gemsbok, to 3–10 and up to although malesmaynotbeginspermatogenesisuntil30monthsof 18inprotectedareasofeasternAfricaandSouthAfrica(Creeland age(Rudnai1973b;Smutsetal.1978).Malesinthewildmaynot Creel 1997; Nowell and Jackson 1996). Density of Asian lions is haveanopportunitytoreproducebeforetheageof5years(Smuts estimated at 1 lion/7 km2, with mean home ranges for males and 1982). Four years is the median age when females have their 1st femalesof110km2and50km2,respectively(Chellam1993).Var- surviving litter and when males 1st become resident in a pride iationinpopulationdensitiesistheresultofdifferencesinresource (Packeretal.1988; Puseyand Packer1987). availabilityandcompetition(Stander1997).AdultsexratioforAf- rican and Asian subspecies (1 male to 2.1 females) is skewed ECOLOGY.Lionspreferopenwoodlands,thickbush,scrub, heavily in favor of females because of high male mortality, espe- and grass complexes (Kingdon 1977; Yalden et al. 1980) but will cially during subadulthood (Chellam and Johnsingh 1993; Smuts occurinsemideserts,forests,andinmountainsupto5,000m.The 1978b; VanOrsdoletal.1985). GirForestsofIndiaaredry,mixeddeciduousforestsdominatedby Home-range size of a pride correlates with lean-season prey teak (Tectona grandis) in the upper story with acacia thorn(Aca- biomass,butnotwithgood-seasonpreybiomass(VanOrsdoletal. cia)savannaintheeasternportion(Berwick1976). 1985).Inthedryseason,lionshuntcooperativelymorethanduring Prey selection of African lions is well studied (Elliott et al. other seasons. During the wet season, groups of lionesses obtain 1976; Eloff 1984; Mills and Shenk 1992; Rudnai 1974; Ruggiero morefoodthandosolitaryfemales(Stander1991). 1991;Scheel1993a,1993b;Smuts1978a;Stander1992b).Ineast- Prides allow greater protection for the individual lionessand ern Africa and parts of South Africa, wildebeest (Connochaetes her offspring and greater success in hunting effort (Packer 1986; taurinus) comprised the majority of the lion diet (Schaller 1972). PackerandRuttan1988).Coordinatedgrouphuntsallowforgreater 762—Panthera leo MAMMALIANSPECIES 5 capture and kill of prey (Packer 1986; Packer and Ruttan 1988; vival or subsequent reproduction (Packer et al. 1998). Disappear- Stander1992a,1992b). ance of cubs 18 months or older is often due to death, but some Lionsandspottedhyenas(Crocutacrocuta)havesimilarprey young may disperse with the mother to a new area (Packer et al. preferencesandoftencompeteatkillsites(Cooper1991;Schaller 1988). In Serengeti, solitary females are less able to successfully 1972). Lions are dominant to hyenas except when substantially rearlargelittersofcubsascomparedwithfemalesinapride(Pack- outnumbered (1 or 2lionsper20–40hyenas);lionsgenerallysur- er and Pusey 1995). Survival of young remains constant with ma- render the remnantsofakilltohyenasonlyafterconsumingmost ternalagebutlittersizedeclinesat14years(Packeretal.1998). ofthemeat(Packer1986).Lionsareoccasionallyinjuredorkilled Conflict with people on reserve borders is a major cause of by prey such as buffalo, rhinoceros (Rhinoceros), warthog, wilde- mortalityandmayaccelerateextinctioninisolatedareas(Woodroffe beest, orzebraduring failed hunting attempts(Bertram1979;Ro- and Ginsberg 1998). Hunting is restricted to problem animals in sevear1974). Kenya and Uganda, with trophy hunting allowed in South Africa, TuberculosishasbeendiagnosedinlionsfromKrugerNation- Tanzania, and Zimbabwe. Lions are an importantgame animalfor alParkandmaybecontracteddirectlyorindirectlyfrombuffaloes tourist hunting and generate ca. 12% of the hunting revenue in (Keet et al. 1996). An epidemic caused by a morbillivirusclosely somepartsofAfrica(Creeland Creel1997). related to canine distemper virus emerged in a lion population of Lions are easily maintained in captivity, breed successfully, Serengeti National Park in 1994 and spread north in the Maasai and survive for more than 20 years (Shoemaker and Pfaff 1997). Mara reserve. The fatal neurological disease was characterizedby In1992,theInternationalSpeciesInformationSystemreportedca. grand mal seizures and myoclonus (Roelke-Parker et al. 1996). 715 lions in captivity throughout the world (Nowell and Jackson Canine distemper virus has been documented in Serengeti lions 1996).Theanestheticspropofol,ketamine,andxylazinehavebeen andcausedencephalitis,nervousdisease,andpneumonia(Carpen- usedsuccessfullyonlions(Epsteinetal.2002). ter et al. 1998). Feline herpesvirus and feline immunodeficiency RelativeamountofwearonP3,widthofthepulpchamberof virusareendemicinSerengetiandNgorongoroCraterpopulations canineteeth,spotmarkingsofcubs,prominenceofnipplesduring (Packer et al. 1999). Trichinella nelsoni occurred in 3 lion car- lactation, number of incremental cementum lines, and certainbe- casses from Serengeti (Pozio et al. 1997). Antibodiestothegram- havioralcharacteristicscanbeusedtodetermineage(David1962; negative bacteriaBartonella werefoundin5%of58free-ranging Rudnai 1973b; Schaller 1972; Smuts et al. 1978, 1980; Spinage lions(Moliaetal.2004).Incaptivity,felineimmunodeficiencyvirus 1976). Skull measurements provide the only means of assessing and canine distemper require individual isolation and treatment ageofyounglions,3yearsofagemorepreciselythantootherup- (ShoemakerandPfaff1997). tion and replacement (Smuts et al. 1978). Shortly after 3 years of Lionsareharassedbystable(Stomoxyscalcitrans)andtsetse age,skullsizeincreasesandskulldimensionsstarttooverlapthose (Glossina morsitans)fliesandotherbitingpestssuchasticks(or- ofolderlions(Smutsetal.1978). der Acarina) and fleas (order Siphonaptera). Lions generally have Lionsareindividuallyrecognizedfromscars,eardamage,and tapewormsfromswallowingcystswithmeatoftheirherbivoreprey. natural markings (e.g., whisker spot patterns—Pennycuick and Sarcoptes scabieihasbeen foundonlioncubsinKrugerNational Rudnai1970).Lionscanbeimmobilizedbydartingfromavehicle Park and may result in mortality in severe cases (Young 1975). (Mills 1996). Death caused by capture is rare in lions, although Variousparasites,includingAncylostomaparaduodenale,Cylicos- severe stress due to human hunting has been linked with fatal pirura, Dirofilaria sudanensis, Lagochilascaris major, Linguat- capturemyopathy(Joubertand Stander1990). ula, Schistosoma mattheei, and Trichinella spiralis, have been found in fecal samples of lions throughout Africa (Bjork et al. BEHAVIOR. The pride is the primary fission–fusion social 2000). Twenty-eight parasites associated with lions include Ancy- unit of lions (Gittleman 1996; Packer et al. 1991; Schaller1972). lostomaparaduodenale,A.tubaeforme,Cylicospirurasubaequal- Prides vary in size and structure, but typically contain 5–9 adult is, Dipylidium, Diphyllobothrium theileri, Dirofilaria repens, D. females(range,1–18),theirdependentoffspring,andacoalitionof sudanensis, Echinococcus granulosus, Filaria leonis, F. martis, 2–6 immigrant males (Heinsohn and Packer 1995; Packer et al. Galoncus perniciosus, Gnathostoma, G. spinigerum,Lagochilas- 1991). Observed groups may be much smaller (1.2–1.9 adults— carismajor,Mesocestoides,Ollulanustricuspis,Pharyngostomum Bauer et al. 2003). Pride sizes are smallest in arid environments cordatum, Physaloptera praeputialis, Taenia, T. bubesi, T. gon- with limited prey species (Elliott and Cowan 1977; Hanby and gyamia,T.hydatigena,T.regis,T.taeniaeformis,Toxocaracanis, Bygott1979;Ruggiero1991;Schaller1972;Stander1992b;Wright T. cati, T. leonina, and Trichinella spiralis (Round 1968). Other 1960). On average, lionessesinaprideshareone-seventhoftheir fecal samples of wild lions had 15 parasite taxa, includingAncy- geneswithotherpridemembers.Numberoflionsinapridevaries lostoma, Coccidia, Giardia, helminth larvae (Ollulanus tricuspis acrosssuccessive monthsbecauseofsynchronyofbirthsandhigh and Aeluronstrongylus), Physaloptera, Sarcocystis, Spirometra, mortalityofcubs(VanOrsdoletal.1985).Malesresideinapride Spirurida, Taeniidae(EchinococcusandTaenia),Toxocara,Trem- for ca. 2 years before being replaced by another group of males atoda,Trichostrongylidae,andTrichuris(MullerGraf1995).Ofthe (Packer et al. 1988). Pride membership is stable but pridemates hostssampled,97.3%wereinfectedwithatleast1parasite.Twelve are often scattered in subgroups throughout the range and each previously undocumented parasites (Aelurostrongylus, Acantho- individual spends time alone (Pusey and Packer 1987; Schaller cephala, Anoplocephalidae, Capillaria, Denodex, Eimeria, Hab- 1972). ronema,Isospora felis,I. rivolta,Isospora,Trematoda,andTrich- The cre`che is the socialcore of thepride and facilitatesma- uris) occurred in fecal samples from Serengeti National Park and ternaldefenseofcubsandjuvenilesfrompredationandinfanticide Ngorongoro Crater ConservationArea,northernTanzania(Bjorket (Cairns1990;PackerandPusey1997).Cubsattempttoobtainmilk al. 2000). Celland blood parasitessuchasEimeriaandBabesia, from any lactating female in the pride although females preferen- aswellasgastricspiralbacteriaandintramuscularsarcocysts,have tiallynursetheirownoffspringandcubsofcloserelatives;females beenfoundinsomeindividuals(Kinseletal.1998;Lopez-Rebollar with small litters give a higher proportion oftheir nursing tonon- et al. 1999). Endoparasites of the Asian subspecies from zoosare offspringcomparedwithmothersoflargelitters(PuseyandPacker similar to parasites existing in African populations (Bjork et al. 1994). 2000).Ancylostoma,Spirometra,andToxascarishavebeenfound Lionsexhibitdiversepatternsofbehaviorbetweenandwithin in both wild Gir Forest lions and those in captivity (Bjork et al. prides,anddifferentpopulationsvaryinfeedingandhuntingpref- 2000). Antibodiesto theabortifacientparasiteNeosporacaninum erences and methods (Rosevear 1974). Most studies of lions are occurinlions(Ferroglioetal.2003). observational,andsomeexceed40years(Hanbyetal.1995).Other Mortalityforlionessesdeclinesat3–4yearsandthenrapidly methods include censusing, DNA fingerprinting, immobilization accelerates; no lioness has survived beyond 17 years in Serengeti (with ketamine hydrochloride–xylazine hydrochloride—Herbst et NationalParkorNgorongoroCrater,Tanzania(Packeretal.1998). al.1985),marking,and playbackexperiments. Maleslive an average of12 years and up to16years(Hanbyand Lionsaremainlyactiveatnight.Alargeportion(ca.80%)of Bygott1991;Smutsetal.1978).Cubmortalityishighinlionsand thedaily time budget(recordedinTanzaniaduringa24-hperiod) is linked to periods of prey scarcity and infanticide bymalelions isspentsleeping,lyingdown,orsitting(Hanbyetal.1995).Males during pride takeovers; 27%ofcub mortalityisduetoinfanticide usevocalizations(roaring)andscentmarkingtodefineboundaries, (Packer and Pusey 1983b; Schaller 1972; Van Orsdolet al.1985; although these behaviors also function in communicationbetween WhitmanandPacker1997).Cubsurvivalisdependentonmaternal pridemates(FunstonandMills1997;Grinnelletal.1995;McComb survival, although juvenile survival is unaffected bymaternalsur- etal.1994;Schaller1972).Roaringchorusbylionshasamaximum 6 MAMMALIANSPECIES 762—Panthera leo intensity of114 dB at1 m and may have an assemblyfunctionto on the number of males in a coalition (Packer and Pusey 1997). which stray members respond and join the group in response to Male pride tenure rarely exceeds 4 years before another coalition territorialdefenseagainstotherprides(McCombetal.1994;Peters takesover(Bertram1979; Packeretal.1988; Stander1992b). and Wozencraft1996).Lionsuseurinespraysorscrapestodefine Intheabsenceofapridetakeover,malesgenerallyleavetheir territories(Schaller1972). natalpridewhen2–4yearsold(Bertram1975b;PuseyandPacker Whenactive,lionsengageinhunting,communalcubrearing, 1987). Most females are incorporated into their natal prides, but and territorial maintenance (Heinsohn and Packer 1995). Lions some subadult females (33% in the Serengeti) emigratewhen2–4 normallywalkatca.4km/handcanrunforshortdistancesat50– years old (Pusey and Packer 1987; Van Orsdol et al. 1985). Evi- 60km/h;activityintreeshasbeenrecorded,althoughlionsarenot dence from Serengeti indicates that dispersal may be disadvanta- adeptclimbers(NowakandParadiso1983).Wrestlingismostcom- geous for females because of decreased survival of their 1st litter mon in cubs but is rarely used by adults, whereas stalking was (PuseyandPacker1987).Femalesthatremainintheirnatalpride infrequently observed in play by cubs (Schaller 1972). Self and haveareproductivelifespanofca.12yearsbeginningat4–5years social grooming are frequent activities within a pride; forepaws, ofage(Puseyand Packer1987; VanOrsdoletal.1985). chest,andmaneonthechestaremostfrequentlygroomed(Rudnai A lioness may attract a male during estrus by urinating and 1973b).Variousbehaviorsandsoundsthatmayberelatedtosocial spraying scent from her anal glands along travel routes (Rosevear maintenanceandcommunicationincludeclawing,crouching,duck- 1974). In the Luangwa Valley in eastern Zambia, copulation has ing,grunting,headrubbing,rollingontoback,scratching,snarling, occurred between females and males from differentprides(Yama- stretching,andtailflicking(Rudnai1973b). zaki1996). The 1stmale to find a female in estrus willguardher Territorial defense involves male, female, and juvenile lions and attack any approaching male (Packer and Pusey 1997).How- (Grinnell and McComb 1996; Heinsohn 1997; Heinsohn et al. ever,lionesseswillmatewithseveralmalesduringestrus(Bertram 1996;McCombetal.1993).Malesdefendtheprideagainstincur- 1975a;Bygottetal.1979;PackerandPusey1982;Schaller1972). sionsbyothermales,therebyensuringsomeexclusivityinmating, Matingoccursduringdayandnight.Thepairremainclosetogether, andfemalesdefendtheiryoungagainstinfanticidalmalesandtheir and eithersexmayinitiatecopulation,withthemaleinitiatingca. territory againstadjacentfemaleprides(Packeretal.1991).Male 40% of copulations during courtship (Rudnai 1973a). Ritualistic lions have territorial patrols that protect the pride from others displaysassociatedwithmatingincludevocalizations,suchas‘mat- (Packer and Pusey 1997). Females protect denning sites, hunting ing snarls,’ social grooming, and following of the lioness by the grounds, and water holes from other prides (Packer and Pusey male(Cooper1942;Rosevear1974;Rudnai1973a).Crouchingby 1997). the female generally indicates receptiveness, leading to the male Members of adjacent prides usually remain severalkm apart mountingwithhisforelegsandcopulatingandendingwiththemale (Hanby et al. 1995). When neighboring prides come into contact, bitingtheneckofthelioness(Rosevear1974;Rudnai1973a).Cop- lionesses typically attempt to expel intruders if they outnumber ulation lasts for ca. 55 s (range, 30–154 s—Rudnai1973a).After them (Heinsohn andPacker1995;Schaller1972).Somelionesses copulation,thelionessoftenrollsontoherbackandremainsprone playagreaterroleindefendingtheterritorythanotherfemalesin (Rudnai1973b). the same pride (Heinsohn and Packer 1995). As they approach Lions are opportunist carnivores that usually hunt in groups; sexualmaturity,juvenilelionessesbecomeprogressivelymorelike- maleshuntlessfrequentlythandofemales,butmalesarestronger lytojoinadultfemalesinterritorialdefense(Heinsohnetal.1996). and can gain access to kills made by females (Bertram 1975a; Increasedaggressioninpridedefenseoccurswithhighliondensity, Scheel and Packer 1991). Prey selection is related to seasonal such asin the NgorongoroCrater(Heinsohn1997;McCombetal. weatherpatternsandthemigrationoflargeherbivoresinsomeparts 1994).Territorialdisputesoftenendwithlargergroupschasingoff ofAfrica(Hanbyetal.1995).Mostspeciesofpreyarelarge,with smallergroups(Hanbyetal.1995). a modal prey size of ca. 150 kg (Packer 1986). Average food ac- Malesdefendtheirareabyusingcooperativebehaviorthatis quisition ranges from 8.7 kg/day per lioness in the dry season to not conditional on either kinship or behavior of the male’s com- 14 kg/day per lioness in the wet season in Etosha National Park panions(Grinnelletal. 1995). Roaringmayfacilitatecommunica- and 8.5 kg/day when prey animals are abundant in Serengeti Na- tion within prides as well as discourage the approach of nonpride tional Park (Packer et al. 1990; Stander 1992b). Males may con- membersfromaterritory(McCombetal.1994).Femaleswithcubs sumetwiceasmuchmeatasfemales;cubs(,1yearold)consume candistinguishtheroarsofresidentmalesfromthoseofunfamiliar one-thirdasmuchasadultfemales;subadults(1–2yearsold)con- males that might pose a threat to their offspring (McComb et al. sumetwo-thirdsasmuch asadultfemales(Packeretal.1990). 1993). Capture of prey includes searching, stalking, attacking, and Asmallproportionoflionsarenomadic,includingyoungand subduing (Elliott et al. 1976). Cover, group size, prey-group size, adult males without a pride. Nomadic lions follow the migrations andlight(duringnocturnalhunts)affecthuntingsuccess(VanOrs- of prey and hunt and scavenge cooperatively(Bertram1975a;By- dol 1984). Lions stalk with their heads low to the ground, moving gott et al. 1979; Schaller 1968, 1969; Van Orsdol et al. 1985). slowly and graduallytoward prey(Rosevear1974).Smallpreyare Lionesses are highly philopatric, and few females are nomadic; a struckandstunnedorkilledfromasingleblowofthepawfollowed solitary female usually returns to or settles near her natal pride by a quick bite if necessary (Rosevear 1974). Larger prey are at- (Packer 1986). Nomadic lions, which have large ranges that may tacked typically at the shoulder or flank, the hind claws deeply overlap with pride territories, are commonly found singly or in embedded,forepawsattheneck,chest,orfarsideofthebodywith groupsofasmanyas5,withmembershipchangingfreely(Schaller a seizing of the muzzle and twisting of the head (Rosevear1974). 1972). Preyisdraggeddownratherthanknockedover(Ewer1973).Ata In a pride takeover, a coalition of males that generally are kill, lions eat in the open or drag the carcass to a more protected relatedeitherattackandkillorotherwisecausethedeathsofsmall locationwherebloodislickedfromthecarcass;thebellyisripped cubs(Hanbyand Bygott1987; Packerand Pusey 1997). Withina open; and intestines, internal organs, connective tissue, muscle, few days of cub mortality, females resume estrous cycles. Mating bones, and skin are consumed (Rosevear 1974; Van Valkenburgh activity is quickly initiated by the newly resident males. A syn- 1996). chronous birth event within the pride is the result (Packer and InAfrica,lionsaremoresuccessfulatcapturingsmallerprey, Pusey 1983a). By removing cubs from a pride and stimulatingfe- suchasThomson’sgazelle,whenattacksarelaunchedatdistances malereproduction,malescanfathermoreoffspring.Subadultmales of #7.6 m than from further distances. For larger prey, such as usually leave attakeovers; subadultfemalesbecomeperipheralor wildebeestsandzebras,huntingsuccessis50%atdistancesof15.2 leaveunlesstheymatewiththeincomingmales(HanbyandBygott m(Elliottetal.1976).Of61stalks,only10weresuccessful(Rud- 1987). nai1973b).Variationinoverallhuntingsuccessratesreflectsqual- Cohortsof$3malesusuallyenternewpridesasagroup,but ityofthehuntingenvironment,suchasopenorshort-grassplains, cohortsofonly1or2malesoftenteamupwithsinglemalesfrom or food abundance (Elliot et al. 1976; Eloff 1984; Funston et al. other prides to achieve successful takeover of a pride (Packer et 1998;PackerandRuttan1988;Schaller1972;Stander1992b;Van al. 1991). Bachelor groups are formed as coalitionsofrelatedand Orsdol1984).Onein3attacksonThomson’sgazellewassuccessful unrelated males; coalitions of unrelated males are not larger than inlonggrass,butsuccessratefellto1in6inshortgrass(Schaller 3individuals,whereascoalitionsof4–9animalsarecloserelatives 1969).Huntingsuccessgenerallyincreasesasymptoticallywithin- (PackerandPusey1997).Amale’sreproductivesuccessisdirectly creasing group size when individuals are expected to hunt coop- related to his length of tenure within a pride, which is dependent eratively (Packer and Ruttan 1988). Success rate in capturing 762—Panthera leo MAMMALIANSPECIES 7 Thomson’s gazelle, wildebeest, and zebra ishigher when.2 lion- may presentnew conservation challengesbecauseoflimitedterri- esseshunttogether(Schaller1972).However,theaveragebiomass torial space (Singh 1997). Lion–human conflict will continue to of kills made by groups of different sizes or the hunting rates of challenge management in the Gir Forest, where lion attacks on different-sized groups is not consistent (Bertram 1975a, 1976, humans are associated with prolonged drought conditions(Nowell 1979; Packer1986; Packerand Pusey1982, 1983a,1983b). andJackson1996;Saberwaletal.1994).Prohibitionoflionbaiting Lions exhibit individual preferences in prey selection within fortouristshows,consolidationofreserveboundaries,development and between prides in the same area (Rudnai 1973b; Van Orsdol of movement corridors between suitable areas, arresting the deg- 1984). Prey selection is based on sexual and social differences radation of peripheral forests and wastelands, and more equitable within a pride (Funston et al. 1998). A complex division of labor compensation to villagers for livestock destroyed by lions could was found among lionesses hunting in Etosha National Park,with lessen lion–human conflict in the region and improve lion habitat individuals repeatedly playing the same role during stalking, am- (Saberwaletal.1994; Singh1997). bush, and kill (Stander 1992a, 1992b). Group hunts ofteninvolve spreading out around the intended victim, with each lion takinga REMARKS. Variation in size of teeth, auditory bullae, and differentroutetooptimizecatchsuccessifthepreyanimalattempts othercranialanddentalfeaturesmaynotprovideenoughevidence escape (Bertram 1979). However, highlydevelopedteamworkdur- forsubspecificdesignation(Allen1924;Hollister1918).Morethan inghuntingisnotobservedinSerengeti,whereindividuallionesses 20 subspecies of the lion were based on zoo specimens, but cap- mayrefrainfromhuntingdependingonthepreyspeciesbeingpur- tivity influences skull shape (Hollister 1918), rendering zoo spec- sued (Packer and Pusey 1997). Lions do not adjust their stalking imensuselessfortaxonomy(Smithers1983).Geneticmaterialfrom behavioraccordingtovigilanceoftheprey(Scheel1993b). lionsineasternandsouthernAfricaissufficientlysimilartowarrant asingleAfricansubspecies,P.leoleo.Iflionsweretobeclassified GENETICS. Lions have 38 chromosomes. Estimatesofhet- as a unique subgenus or genus the name Leo would have prece- erozygosity vary with method and genetic structure sampled, and dence (Hershkovitz 1966). However, lions have been crossedwith includesvaluesof3.7%(Menotti-RaymondandO’Brien1995),ca. tigersandthehybridshavesuccessfullymated(Guggisberg1975). 26% (Shankaranarayanan et al. 1997), and 0.75% (Spong et al. Sperm from wild and captive Asian subspecies have a high inci- 2002).Isolationoflionpopulationscausesalossofgenetichetero- dence of abnormalities (O’Brien et al. 1987a; Wildt et al. 1987). zygosity in some areas (Wildtetal. 1987). Low geneticvariability TheetymologyofleoisLatinforlion. may be characteristic of the Asian subspecies and not the conse- C. R.HaringtonandC.Packerreviewedearlierdraftsofthis quenceofinbreedingdepression(Shivajietal.1998). manuscript.ResearchassistancewasprovidedbyDanaSanchezat LionsofNgorongoroCraterinAfricaexperiencedapopulation theUniversityofArizona.SusieGillattprovidedtechnicalsupport crash after an outbreak of a biting fly (Stomoxys calcitrans) in increatingtherangemap.Photographsoftheskullwereprovided 1962.TheNgorongoropopulationbottleneckcausedlossofrestric- byY.Petryszyn.TheSmithCollegeinterlibraryloanstaffprovided tion fragment length variation in the class I genes of the major excellentassistancein obtainingobscurecitations. histocompatabilitycomplex.Thesegenesplayacriticalroleinthe LITERATURECITED development of immune defenses (Packer et al. 1990). Decreased genetic variability of lions from Ngorongoro Crater did not affect ALLEN,G.M. 1939. AchecklistofAfricanmammals.Bulletinof thereproductive-endocrinesystemoffemalescomparedtooutbred the Museum of Comparative Zoology at Harvard College 83: counterpartsfromSerengeti(Brownetal.1993). 1–763. Asianlionsaregeneticallydistinctfromsub-Saharansubspe- ALLEN,J.A. 1924. CarnivoracollectedbytheAmericanMuseum cies, although the difference is smaller than genetic differences Congo Expedition. Bulletin of the American Museum ofNat- betweenhumanracialgroups.Basedongeneticdistance,theAsian uralHistory47:73–281. subspeciesseparatedfromtheAfricanpopulationca.100,000years ANDERSON,J.L. 1981. There-establishmentandmanagementof ago(O’Brienetal.1987a,1987b). alionPantheraleoinZululand,SouthAfrica.BiologicalCon- servation19:107–117. CONSERVATION STATUS. The Asian subspecies, P. l. ASDELL,S. 1946. Patternsofmammalianreproduction.Comstock persica,waslistedintheConventiononInternationalTradeinEn- PublishingCompany,NewYork. dangered Species (CITES) Appendix I in 1977 and is fully pro- BAUER, H., H. H. DE IONGH, AND I. DI SILVESTRE. 2003. Lion tected in India. The United StatesEndangered SpeciesAct(ESA) (Panthera leo) social behaviour in the west and central Af- andtheInternationalUnionforConservationofNatureandNatural ricansavannahbelt.MammalianBiology68:239–243. Resources (IUCN) recognize the Asian subspecies as endangered BENNETT, E. T. 1829. [unknown title]. Towermenagerie.Volume (Wilson and Reeder 1993). A census estimate for the Gir Forest 1 [pagesunknown](notseen,citedin Pocock1930:660). populationapproximates280individualsinandaroundthereserve BEN SHAUL, D. M. 1962. The composition of the milk of wild boundaries (Khan 1995). Other extant subspecies were listed in animals.InternationalZooYearbook4:333–342. CITES Appendix II in 1977. 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