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Home , Cheetah, Cheetah (rapper), Kakamega Forest, Leopard, Ol Arabel

Hamilton PH. 1981. The and the in Part 1.

Keywords: 1Afr/1KE/ jubatus/cheetah/Leopard/status/survey

Abstract: Part 1: Report about the leopard. The related "endangered " the Cheetah is included in Chapter 4 and 5. A survey and discussion about the status of the species.

, I (tt t\ THE LTOPARDPanthera pardus I -t AND THt CFEEIAPAc illllif jubatus

IN

KENYA

ECOLOGY STATUS c0Ns€RvATl0li fiANACE14EIJT

R€PORTFON

THEU.S.FISH 6 VILDLIFES€RVICE

TIE AFRICANVILDLIFE LEADERSHIP FOUIIDATION

Al{f)

THE 6OVERNHTNTOF K€NYA

P. H.HAI4 I LTON r98r

ALL RIGHTS RESERVED includingthe . ishi of reproduction in wholeor in part in any form without the permisslonof the author or other the colyright holder for the t imebeing. t' -t_

TAELI OF COIITENTS

L IST Oi FI GURES I ( LIST OFIA8LES

ACK}IOI'LEDGEI'IEIITS

!r4Ilr -I INTRODUCTI ON

CHAPTTR? ECOLOGYOF THE !EOPARO 4 2 .1 | tiTRoDUCTI 0ll tr 2.2 STUDYAREAS L 2.3 }IATIRIALS AND XETHODS 2 .\ P.ESULTS 6 2.!.1. captureand radio-trackingresults 2.\ .2. PoPuIat I on densi tY 2.!.3. Reproduction e of riovcment 2.\.\. Extent tc 2.\.5. 0ccuPancYof the hc'nerange 1l 2 .. .6. ActlvitY Peraods ll Z.!.7. Rest irg Pleces 2.{.8. Feedi n9 ecologY 2.1.9. P-rterns of dlgPersion 17 2.4.10. Pat terns of association 2.!.ll' Soclalorganisaticn rB t9 2-5 DT SCUSS I0N

ChAPTER3 TRAIiSLOCATIOII ?l I l{TR0DUCT| 0ll tlEiH0 D s ).2.1. CaPtureand taanslocation 3.2.2 - tffnobiI isation 3.2.3. Captive 3.2.\. Radlo-tracking RISULTS 3.3.1. Radio_tracking9!cce9s j.t.2- f{ovetnentsof trsnslocatcd leoPard5 2t) 3.3.1. Othcr traislocations in Kenya

5.1 Drscussl0N 35 3.1.1. Translocation: criteria for succes5 the lleru study 3-\.2, 39 3.1r.3. Reasonsfor fai lure not do it t.t,t. Translocationi hoh, to r13 3.\.5. Ttan5locat ion i guide-lincs i.l.e. Conclusion: to translocateor not? 7:.

CHAPTERtr STATUS0F TH€SPECIESi A SURVEY {.1 tNTBooucTt0N \.2 r.rtTHoos \7 I 4 . ]. STATUSSURVTV 4.3.1 . tormat I 4i \.1.2. TurkanaDistrlct tti ,t.i.3. 8erIngo Dist r ict tt.3.It , Vest Pokot t Elgeyo-llarakr'retDistrlcts .{.J.>. UasinGirhu, TransNzoia, 6l{andi Dist.ic!s 5: {.3.6. Nyanza6 Veste rn P.ovinces \.3.7. 6 KerichoDistrlcts 5? 4.3.8. NarokDistrict Kaji6do 0 is t|'l ct !.3.10.Nai robi P.ovince 5t !.3. . llachakosE Klt!i 0istricts \,3-12.Centra I Kenya L1 4.3.1).ia it. Di5 t. ict 64 4.3.I!. TsavoNatlonal Park 4.3.r5.Galana 6 Kulalu Ranches 65 {.r.r6.Kwale,Kilifi, tlombasa,6Lan! Dis!ricts 61 4.3.r7.TansRi ver Distric! 6e {.3.r8.Carissa Distrlcr 69 4.3.r9.Vajir 6 HanderaDistricts q.3 .20. lsiolo Distrlct 71 \.3.21.Samburu0istrict 12 harsabit 0istrict TBELTOPARD AS A STOCK.RAIDER 4.{.1. Introd'.rction 4.4.2, Theoccurrence of stock-raiding 75 4. {.l. lecpard control 7t 4.4,1. Sex .atlo of stock-raidingteopards 1i it.4.5. Discussion

CHAPTER5 STATUS0F THESPECIES: DISCUSSI0N

5.1. THEI.EOPARD II{ KENYA 5.1.1. ThePasr 6l 5.1.2, The oecline E] 5.1.3, Extentof thc Decllne 84 5.1 .4, Where,Uh€n, and Uhy? 5.1.5. P.egcn!5ta!us 5.!.6. The Leopard'sFuture ln Kcnya 5.2 THEIEOPARD III 9: J.3 STATUSOF THTC8EETAH IN KENYAAND AFRTCA CHAPTER6 THELEOPARD A5 A HUNTINGTROPHY O TOURIST ATTRACTION

I NTROOUCTION r02 LEGISLATIVE HIsTORY 102 lc3 ( BACKGROUNDTO THI HUNTINGIIIDUSTRY IN KENYA-, 5 .lr THg LEOPARDAS A IIUNTINGTROPHY 105 5.5 THE LEOPAROAS A TOURISTATTRACTION 107 EFFTCTSOF SPORTHUNTIIIC ON LEOPARD POPULATIONS 108 Drscussr0N 6.7.1. To hunt or not to hunt? ll2 6.7.2. tthcre? ltl 6.7.3 . llhen? I llr 5.7.t,. Howmany? I 1/{ 6.7 .5. For ho!/much? 124 6.7.6, By whomand under rvhatconditions? t2l 6.7.7. u,s. legislation t2J 5.7.8. The problemof stock-raiders 12\ 6.7,9, Concluslon t25

CHAPTEP7 CoNStRvATl0N o I4ANAGET4ENTOF THE LIOPARD IN KENYA A PO!ICY 1)1

REFER€NCE5 f1

APPEI.ii'IX I LIST OF FIGURfS

observedranges of tcn radlo'collared T3avo leopards, as determinedby the nininufi area ( oethod uslng all locations. - t Ft6.2.2. Adjustedhome ranges of nine radlo-collsred Tsavo leopard9. 9-10 FtG,2 .3 . llovernentsof adult nale Leopard3 during the peri od l7-21 June 1973. Facin9 I l

Fr6.3.1.l'leru tlational Park 6nd ru.rounding area. racrng zo FrG.3.2.l,lovementsof Leopard99 In first fortnight after release on 22 Dec 77. Facing 29 FI G.3 .3. l,lovenent5of tuo translocated leoPards re- leased in Tsavol/est National Park in 197C. Faci ng 31 FlG.r.l.Schematicdlagram of the digPersal of trans- iocated lropatds rel€ased in lleru National P.rk, Faci n9 39 - Frc.!.1.Adninistrativedistricts of Kenya. Ir9 5o FlG .Ii.2. Huntingmap of Kenya(1970) showing controlled area block5. \9'5a FrG.{.1.The leopardas a stock-raider. tac Ing /o

FrG.5.1.Reletivedensi!ie5 of the leopardpopulations 87-88

F1G.5.2 . Relative densiti€s of the hlnan Populationsof Kenyain 1980. 87-88 FrG.5. 3. Distribution of the leoPard in Kenyain 1963. Faci ng 88 FrG .5 .4. oistribution of the cheetahin Kenyain 1953. racInq >o

:{ L IST OF TAELES

TASLE2. Observedranges ahd hooe rangesof ten radio- col I ared Tsavo lcopards. Facingl0 _l TABLE!.r Numberof leopardsshot on licence In Kajiado: District in 1965-1973. t7 TA8LE \.2 Leopard5i9htin95 recordedin National Pa.k 1912-75. TABLE1r.3. The cheetahpopulation of 1973-75. {c 1ABLEtr.4. Stock-raid i n9 questionnai re survey of tli I dl i fe conservation and l'lanagementoePartnent game stations. TASLE{.5. camestctions (.r'ith correction for !eopard) I ist ing carnivoresas gtock-raiders. TABLE4 .5. Leopardsghot .nd traPPedby GovcrnDentagencie5 1957-55and 1977-80. 1ABLE4 .7. Analysis of carnivorecontrol. IAELE4.8. Sexesof 83 leopsrds trapped for transloca!ion. n1-n! Status of the leop€rd in the distrlcts of l(enya. IASL€5.2 . Numbersof special licences bought for leoPards in 1958-73and nuhbersof leopards ehot in con- trol led aleas. a5 TAELE 5.3. Asses5edstatus of leopard populations in Kenya groupedio Censi ty categories. 88 lABL€5. {. Assessedgtatus of the leopard PoPUlationsof Kenyarsdistricts groLipedin density categorleg. Faci n9 89 lABL[ 5.5. E5timateddistrlbution of Kenya'9leoPards. 89

TAEIE 5.1. Contribution of the leopard to sPecial licence revenueand controlled area fees in 1965, 1972' and 1973. 105 TAB!€5.2. Relativecontributlons of different 3Peciesto revenuefrom ggeclal licences. r 05 TABLE5. ]. Reletivecontributlons of different Speciesto local authoritv revenuefron controlled area fee5. 106 TABLE5.q . Con!ribution of the Ieopard to control led area fees earnedby Naroksnd r€jiado county councll5 ln 1965. 106 TABrE6.5. liumberof general and special huntlng I lcences issued.nd revenle derived therefrom in 3elected betvreen1959 aod 1974. 107 TA8!E6 .5. Numberof perrnits and licences of various cete_ gorie5 i95uedby the GameDepartment between 1950 and 1974. 1ll ACKNOVL TDGE}IENTS

So manypeople have glven n|evalued help one way o. another durlng my leopard gtudies ln Tsavoand l,lerulilational Parkgand durino ny leopard gtatus survey 3lnce then that lt ls dlfficult to knoel where !o begln. 8ut as full acknowledgementsto a1! those whohetped ne in Tsavohave already been glven In Hamilton (1976) lt would be superfluous to repea! themhcre, and lack of space lmposesfurther constraints. I must, holrever,record once agaln rnygpecial apprecae- tion of the help and co-operation recelved from the 0irector and Trusteesof the forfi,erKenya l,lational Parks, the Vardena,]d staff of Tsavol,/es t NationalPark, Dr. L.D. flechof the U.S. Fish andVildllfe Servlce, Dr. J,f,1.Klng of Nairobl, and my tracker/assistant Elui Nrnen!i whovras vltal to the difflcult task of 3tudylng leopards. The Tsavoand l.lerustudles were both hade possible by ttre genercJsfinencial support of the Afric.n t{rildlife LeadershlgFo!n- dation of Vashingtor,0.C.i supportwhich not only Drovidedne nith a vehicle but al50 the use of ltr Cessna185. I am grateful for help received fron all the Foundatlonrsstaff In !,/ashingtonand Naiaobi, but eepecially rhe late J,E. Rhea,the late F. l,tinot,the late R.K. Poole, R. l'lcllvaine, and SandraPrlce. Thelr constant support ano encourager,renthave be!n deeply appreclated€nd I amonly 50rry that not all of themare here to 5ee the fln6l results. Their contributior,s ate not forgotlen. TheU^lted States Flsh andWildlife ServiceOfflce of En- dangeredSpecie5, to b/hichthls report is contracted, provided funding for expaision of r,rystudies to include e detailed survey of the stat.r5 of rhe leopard and has patlently awaited the results. I em grateful for both its financial supporrdnd lts patience. In KenyaI om grateful to the office of the Preiident and to J,K. llutinde, fortner Director of yildllfe Conservatlonand lianage- nen!, for permisslonto caray out this research; to D.|.1.Sindlyo, presentDirector, for his co-operatlonIn organislnga questlonnaire surveyof the Departhen!r5game stations; to P.R. Jenkins,f.l.B.E. who ra5 Vardenof ileru Natlonal Park during ny translocation gtudy anC r{hoseco-operation, advlce, hospltal lty, and frl€ndshlp are especial!i appreciated; to F.V. Voodley,f4.C., 1,1.8,E. and P.M,Snyder, who as Vardensof the llountain l'/ationelPirks at ltfieigaprovided vlrtually al I the leopards I recelved for translocatlon; to V.H.V. Ngonze, Vardenof the Nairobl Anlrnalorphanage, and all the Departmentrsother wardens! too numero{,rsto oention by narne,who provlded infolrnation on leopard5tatus, depredatlons,anC translocations; and to the Vildlife FundTrustees who handledmy researchfunds, lam Indebtedto the followlng past and presentofficers, nogt of thah very senlor, of l(enyaNationel Parks, the GameDepartftent, and the lrildllfe Conservatlonand ]lanagementDepartnent fo. provlding lnformation on leopardsfor ny gtatus survey and allovringoe to bene_ fit from their experlcnce, ln manycases exceedingthlrty years, of all aspect5of wildlife conservatlonand managenentln Kenya: G.A.G.Adrn- son, 11.8,E.,J. Barreh,0.8.t., J, B.xendale,itajor R.T. Elliott, E.L. Gos5,l4ajorl.R. Grip{ood,l. Hughes,P.R. Jenkins,r'1. lodhe, l.S.C. Parker, K.A. Slnlth,14.0, uebley, F.V. Woodley. | .lso wiEh to thank Dr. 0.R.fl. Stev/artof the Natal Parks, ce$e and Fish PreservationBoard and Dr. S.C.J. Joubert of the Kruger Nar.ionalPark, Repuhlic of SouthAfrica, for the lrouble they took to sendne Informationon the trenslocatlon of . The infor- mation frornDr, Joubert arrlved too late for IncorPoration in this ( report but i! serves only to relnforce llly conclusionsoil transloca aon. Diicussions$itt' Dr. J' Andersonof lhe Nat.alParks 5nd !r. A.D. Hall- ,lartin of the Kruger Park vrerealso helPful. liy survey of the status of the leopard In Kenyacould not have been donewithout the excellen! co_operationI received from the colniry's profesgionalhunters. Theygave generously of their tine and knowledge,and Chapterlt ls as mr:chiheir vtorkas it i3 mine. Chap!er6, on , alto benefitted froin thelr soundand construc_ tive criticlsm, SeveralProfessiooal hunters had al50 beenin the GarneDeoartrcnt a! one tlme and were usefully sble to vlevr the hurtinc Industry fron different aspects. 8ut v/ithout any doubt the Profese:ot wa5ry mostvaluable source of informationon leoParddistribu!ions and relative numberg,and I record v,/lthapPreciation the contributions mBdeby the following nenrbersof the fortnerEast African Professioi.l Hunters'A5sociation:D. Allen, A.L. Archer, R. ' BaLault,J.t'{. Chef- fings, l. Crai9, P.R,A.D.vey, s.P, Dolrney,A.D'G. Dyer' H.H. Henley, .1.Hessel, R.A. Hurt, t'l.St.J.La!,Jrence, D. flccallun,J. tlcKeand,T.0. Itathev/s,l'1.G. Pretlejohn, A.G Roberts,A. '0. Scth-Smlth,J.C. Su!tr' ' i.F. Veller,0.fl. Villlems,V.H. Vinter. llanyothcr people also helped me in various waYs. I owe a special debt to the farmers and ranchergof l4eruand LaikiPia who weregenero!3 not only in Providingstock-raiding leopards for treas- lo.ation end hospitality for mebut also ail ing sheepto feed the leopardsheld In captivi'.y before release. I efi onlY gorry that thelr co-operationha! not been rewardedby a happier ending to the tranilocetlon3tudy, foa noneof thehwanted to kill a sto.k_raiding leopardif they couldavold it, and their exemplaryco_operatlon did muci.to offset th€ lack shoernby the GameUarden (Nanyuki). I would especially Iike to thank the l'llcholsonsof Bu.guret, the Vebbsof Cedarvale,lhe Fernandesof Kl!ima, the Cralgs of LewaDowns, the l.{innsof Loldaiga, the }lurraysof Lolomarikend llarania, the ValI in9- tons of llbori, ihe Dyersof l{gareNdare, the carneglesof Ngobil, th' Stonerriggsof Sugurol, and the Fernandesof Tharua. Their friendly help ano supporti5 greatly aPpreciated' I amalso grateful to th. following for their help anc in- for|ration: T.J, 86rnley,D.E. Brown,C.R. F;eld, B.A. Heath,P. Henp- hill, Alison Harris, A.D. t{ackay,P.S. l4awson,Cynthia l{os-s, N. l1vers, C.E. l{orris, B. Nyebet,R. Paterson,9. Roberts,Karen Ro;s, J.5cort, O. [3stern, and numerousgame rangers and herdsmenin variou' Parts of th! colntry whoscnarnes I cannot list here individuelly. Hamiltor' Harrison0 f4athev/s,A.l'/.L.F., and the Vi ldlife PlanningUnlt kindiy providedoffice spaceand facilitles. Stetionary0 SystemsLtd. and !he AcmePress Ltd. of Nairobi producedthe report. Finally I nru5tcspeclally thank all of the twenty or mor' pe.rle namedabove vrho ktndly agreedto read varlous Portions of my iraft and greatly helpedme to improvelt. I amesPeclally grateful, however,to lan Parker, Rodney€l1lott, Tony Seth-Smitli,and John Suttonfor giving up 9o muchof thelr time and for their invaluable cri!ici5m and conrnents. ThesehelPed to glve the repor! any nerit it lnayhav€, but any rraors tha! remainare my responsibillty. alone. t alro wlsh to thank D.. D.R.t4.Stewart of the l{atal Parks, Gareand Firh PrcservatlonBoard snd Dr. S.C.J, Joubert of the Kruger ti3:ionel Perk, RePublic of SouthAfrica, for the trouble they took to sendr.e inforBa!ion on the translocatlon of carnivoa€s. The infor- mation from Dr. Joubcrt aarlved too late for incorPoratlon'in lhis r€oort but it serves only to reinforce my conclugionson transloca'-ior. Diic,,rssionswith Dr. J. Andersonof the t'latal Park5 and Dr. A.D. Hall- f1artin of the Kruger Park were also helPful. l1ysurvey of the statua of the leoPard In Kenyecould not havebeen donerlthout the excetlent co_oPerationI received fron the coun:ryts profe5sional hunters. TheYgave generouslyof thelr tirne and knowledge,end Chapte.I is as mr:chthelr brorkas it i5 riine Chapter6, on hunting, also benefitted from thelr soundand construc- rive criticism. Several professiontslhunters had also been in the cane Departfi,entat one time and were usefully able !o view the hunling industry fron dlfferent aspects. 8ut wlthout any doubt thc Profcsg:on vr6sry most valuable 5ourceoi information on leoParddigtributions 6nd relatlve numberg,and I record wlth appreciatlon the contributions nade by the following membersof the forner East African Ptofestiona: Hunters'Association: D. Allen, A,L, Archer, R.|'1.Babault, J.H. Chef- fin9s, l. Cral9, P.R.A.Davey, S.P. Downey,A.D.G. Dyer' H,H. Nenley' ,1.iesset, R.A. Hurt, f.St'J. Lavtrence,D' l'lcCallun,J. ficKcand'T.O. fiathe!/s,I'1.G. Prettejohn, A.G Robetts, A.H'0' Seth-Snlth, J.C' s!ttor,, i.F, veller, D.H.Fllllams, v.H. trinter. llanyother people also helpcd |ne in various r0ays. I ov/e a spccial debt to the farme.l and ranchersof t4eruand LaikiPia vihc were generousno! only in providing 5tock-raiding leoPar,rsfor trer9- lo--.tion and hospitality for me but also alling ehecPto feed the leopardsheld tn captivity before release. I amonly gorry that their co-oper€tionhas no! been rewardedby a haPPierendlng to the translocetlon ttudy, for noneof themh,anted to klll a stock-ralding leopard if they coutd avold lt, and thelr exehplery co_oPeratlon cld rnuci.to offset the lack shownbv the Gamevarden (NanYUki). I would esgeciallv like to thankthe Nlcholsonsof Burguret,the vebbsof Cedarvale,the Fernandesof Klsima, the Cralgs of Lcwaoonns ' tne llinns or loldal9a, the fllrrays of Lolornarikand Naranla, the Vall ing_ tons of l4bori, ih. Dyer: of NgareNdare, the Carnegle5of Ngobit, the Stonewiggsof Sugurol, and the Fernandesof Tharua. Thelr friendly help ano support i5 greatly aPPreciated. I amalso grateful to thc following for their help and in- foriiation: T,J. garnley, D.E. Brown,C.R. Field, B.R. Heath,P' HeftP- hill, Al lson Harris, A.D, f'{ackay,P'S. l'lawson,Cynthia },tos-s,N. l'lyerE' C.€. t{orris, B, Nyebet,R. Paterson,V. Roberls,Karen Ro;s, J' sco:t, D. llestern, lnd numerousgalne rangers and herdsmenIn various parts of the country uhosc namesI cannot ligt here Individually' tlamiltor. Harrison6 tlatheus,A.v.L.F', and the vildlife PlanningUnlt kindiy provlded office spacr snd feclllties. St.tlonary E SystemsLtd. and the AcmePrcss Ltd. of l,laI rob I Produced the rePort. Finally t ..utt rsPeclally thank all of the tbrentyor mor? pecple namedlbove $rhoklndly agreed to read various Portlons of my irait anagreatly helpedne to lmptovelt. I amesPeclally grateful, however,to lan P.rker, RodneyElllott, Tony seth-smith' and John S,.rttonior giving !p 30 rluchof thelr tlme and for thelr invaluable criticism aid co'nnents. ThesehelPed to glvr the report any fterit it fl'ayhave, but any errors tha! remainare my responslbillty alone' -1-

CHAPTER

I NTROOUCTION

TheleoDard Dardus i5 the mostwidelv distrlbuted of al I the world's I arsETFfi-(!E[iT-1975) . lt rangesover most of Afr i ca and nuch of Asia, as far to the north and east as l'lanchuria and , and it is probably the most numerous. However,because of I ts secreti ve, sol itary, and lergely nocturnal habits it has been one of the last to yield to 5cientificstudY. Until the spoor tracking study of snith (1977) in zinbabvreand the radio-tracking studies of collared leoPardsin Tsavoand f{eru National Parks, Kenya(Hariilton 1976i this study), the National Park, (Eertrem 1978), and the Kruger Natiooal Park' Republicof South Africa (Bailey, pers.convn.),almost all recordedinforrnation on the leo- pard appearedin popular and semi-popularliterature, most of which Turnbull-Kemp(1967) has surmarised, In the scientific literature the leopardhas featured,usually only lncidentally, in studies of other anirnalssharing the sal||ehabitats (Schaller 1957, Schaller 1972' EisenbergE Loakhait 19/2, ltuckenhlrna Eisenberg1973,8ertran 1978)or in broad scopesurveys of basedupon carcaseanalyses (l{right !950, hirchell, ShentonE Uys 1965,Kruuk e Tumer 1957, Pienaar 1959, Hirst 1969). only onestudy, by Grobler6 yil5on (1972),ha5 specificallY beendevoted to the leopard'5 diel. Indeed,our lack of kno{ledgeof the species in 1967 ls well illustrated by Turnbull-KemP'sbook "The Leopard" Nrhichis laroely devoted to such topics as hunting and photograPhingthe creature,and consideration of it in captivity, heraldry,s!perstition and!ritchcraft, and as a man-eater, In l98l, hc*,/ever,ne are further forward. one reasonfor this has been the developnentin the 1950sand improvementin the 19705of radio- tracking systems(ltech 1973). Another has been the stinulus for study provided by increasing concernover the status of the leopard In the 1970s, and the realisation that not very muchwas knownof the sPeciesr ecology, particularly its movements,populatlon densities, and social organisation. In a paper presentd at the First International Symposiumon the €cology, Behaviourand conservationof the vorid'5 , llyers (1973)drew attention to the massivevolume of the trade in leopard skins and to the sifluitaneousdecline of the species in Africa. This led to a U.S. Fish s vi ldl ife service reoort on the status of the leooard (Paradiso1972) which r€sufted in the leopard being llsted in 1972as an EndangeredsPecies under the United States EndangeredSpecles Act of 1959. An 'rendangeredspecies" is one defined as'rin dangerof extinction throJghoutall or a significant portion of it5 range" (Federal Regigter 1980). Thi5 classlfication Pro- hibited the importation of any leopardsor parts thereof, dead or alive, except under permit, and it effectively stoppedthe heavyflor of leoPards into the united states fol the trade' lt €l5o Dreventedan Anerican sport hunter from bringing hdne a leopard lroPhy shot legally in a country where licensed leopard hunting was permitted. The follolring the leopardwas also placed on Appendix1 of the Conventionon Internatiolal Trade in EndangeredSpecies of vlld Faunaand Flora (c.l.T.E.s.): ApPendix I included all speciese,hich are threatenedwith extinction and are or maybe affected by trade, and required lhat val id permits be issued by both exporting and importing countries. _ Controversyover the status of the leopard contlnueo, hovever, and ln 1975the U.S. Departmentof the lnterior inltlateo its orn slrvey, whlch $rasactually undertakenby the U.S. Stare Departmentthrough lts embassiesabroad. The results were of dosbtfui valldity as usually only one personwas contacted In each country, blt they led the oepartmentof the Interlor to contlnue to clasgify rrendangered,,. the leopard as Shortly aftervrards Xyers (197.6b)publ ished his painstaklng survey of the statug of the leopard ln Afrlca south of the : a study conmleslonedby the Internatlonal Unldn for the Conser- vation of ilaturc and the World fJlldllfe Flnd and pattly flnanced by the International Fur Trade Federationwhich voluntarlly ilnposeda three- year npratoriumon the handling of leopard and cheetahsklns. I,lyersr feport concluded that althoogh the leopard was lrnder heavy pressure frofi I poachlng, lt still occurred ln reesonable numbers ln sorneAirlcan coun- trles and was not in Inlnlnentdanger of extiDctlon. Thls Has "[ew Inrerpreteo by rhe york Times,, (2v2/7ql and the ,,tntematlonat Herald Tri6une,, (22/2/7\) to nean that the leopard 'ias ,'a thrlving species', Hhich had beelr lin found abundant numbersln every African country golth of the Sahara" - which was not ar all what t'tyershad saidl l{evertheless,this Increasedthe pressureon the u.S. Flsh and Vildlife Servlce to renovethe leopard froln it5 endanqeredllst: a move strongly advocated by taton (1975) whose leopard statris survey comissloned by Safari Club lnternatlonal flatly contradlcted oany of Iyers, assess- rrthe |hents and concluded that leopard ls not (nor was) endangered or threatenedIn the majority of sub-SaharanAfrica...and in rrlost of lts range ha9 a satlsfactory and pro,nlslnggtatus." The valldlty of thls assessmentin general and its appllcability to KenyaIn particular will be exaolned later, Eatonrs survey was fol I c*.redby another by Teer 6 SvJank (1977r. thls was comoisslonedby the U.S. Fish and Ulldllfe Service and was based, llke Eatonrs, on a brief questionnaire, with the addition of a numberof intervlews which relieo heavlly on governmentofficials. Teer E Sr,/ankalso concluded that the leoDard shor-rld be deorlved of its starus as an endangeredspecles, and recofinendedthat it should be reassioned to Appendixll of C.l,T.E,S. This covers species whlch are not yei threatened with extinction but which could becomeehdanaered if trade is noE con- trol led. only an export penrit fror the cou;try of origln is required, and there ls no aestrlction on use for cooriercial p|.,rpoi.r. Finally, pronptedby the feeling that none of the surveys cited had been able to makeuse of detailed knc}lledgeof the leopard, the U.S. Flsh and ulldllfe Service askedme to submit a report on the ecoloov and status of the specles In (enya. This was conplementedby requests- the Kenya Governhentfor a survey of the leopard's status and for assess- rients of the speciesr lmportance i9 a stock-ralder and of the value of translocatlon as a conservetlon and managenentpolicy. lly o,vn research on leopards began in l97l i{hen KenyaNatlonal Parks requested a fol I crr-up study of t|,o fadio-collared translocated leo- pard5 which had already been rele.sed in Jsavo Vest ttatlonal park. Realis- Ihg that to study translocated leopards before we had any detailed kncr,v- ledge of an undisturbed porulatlo.r of resldent leopafds was to t6ckle the problen the *rong way round, I persuadedl{ational parks to a o{ a study of resident leopards ln Tsavofirst, Thls they agreedto, and the Afri- can lrildllfe LeadershlpFoqndat ion of Vashlngton,D.C. generooslypro- vided the funds. The results of thls 3o-rbnth study were describedby Hamilton (1976) and in abbreviatedform constltute Chapter2 of thts report. A Seco.ldstudy, of tranglocated leopards in l|eru Natlonal park 5- id 1977-1979,was also fundedby the Foundatlon' The results of this .i" oi'".""iJ in chapter 3 tJhich then examines in depth the whole oolicy and practice of translocation and concludeswith somere_ com"ndationt addressed to the KenyaGovernment' Chapter 4 Presents In the results of a detailed survey of lhe status of tne leoryrd Kenva.financed by the U.S. Fish and Wildlife Service thr+gh the founaition, and includes considerationof lhe 5peciesr lmPortanceas "-iioir-r"iaut. Theseresults are discussedIn chapter 5 $rhichuses in it". to at"t conclusionsabolt the current status of the leoPard ienva in p".ticut"r and In Africa as a whole' and concludesi'ith iecot-ena"tions addfessedto the U.s. Departmentof the lnterior relatina to its current proPosals (Federal Register l9E0) to amend 5 ,ii-i""iti".i"".i".r. oi t;e leopard in the unlted states' chapter as e""rinir tte importanceof lhe leopard es a tourist attractlon.snd " ttunting trophy and includes recoftnendationsaddressed to both the Kenyanaid the iJ.S. Governments.Chapter 7 drewsuPon infor'iation tor oiven in the precedingchapters to present a suggestedPol lcy ionservation ind managementof the gPecies ln Kenya, with recomen_ dations to the KenyaGovernment. "endangered Finally, becauseinformation on a rel ated speci estr, the Cheetah (Aci nony!-iqq9l Schreber), has turned uP so often durin9 mYresearch on the status of the I eopard, and because manyinterest ing points have emerged,I have includedthis 5pecies in Chapter4 andChapter5, The bulk of this report devoted to the Ieopard. CHAPT ER 2

ECOtOGY OF THE LEOPARO

INTROOUCTION

Betweenl97l and 1979 the novementsand ecotogy of resident leopards living in the wild were studied in Ts6voVest and |ieru National Parks by the use of mobile radio-tracklng systemsoperatlng in the 148.350-148.62,IHz frequencyrange. Th- Tsavortudy, whi;h took ptace between l97l and 1974 and in whlch twelve leopards were caught, was more l'detailed and successful than the tleru study, from 1977to 1979, ln which becauseof the scarclty of leopardr as a result gf only three were caught. As the methodsand results of the Tsavo study have alreadv been described in detai | (Hamilton 1976) | proposeor|ly to sunrnariserhen here, with reference to the |leru leopardswhere appropriate, For the convenience of American readers I have accomDani ed metric measurementsby their U.S. equlvalents whereverI haved edned this helpful, but I have sometimesro(lnd€d off the latter for 5inpl icityi the metric neasur€rnentls alwaysdefinltive.

2.2. STUDYAREAS

Tsavotrest fbtional Park covers 9,065 knz (3,500 sq.mi.) in south-easternKenya about halfway betweenNairobi and Hombasa,while fteru National Park, covering 8/2 km2 (337 sq.mi.), lies on the EquarorIn central Keoyaabout 200 kn (124 mil es) norrh-east of N3irobi. Both study areas I ie within the arid eco-climatlc zone of Pratt, CreeftJay€ Gwynne(1955). This is characterlsedby a nean annual rainfall of gene- rally less than 700 |''n (27.6,,)and a high rate of evaporaticndue to a combina!ionof |or elevation (rnostlybelow 1200mor J!40') and high ternperature5. The characterlstic vegetation is dry thorn-bushlandwith Acacia and Commiphoraspecies predonlnating. Thevegetatlon of the tno gtudy areag is girnilar, with the exception of the western portion of lleru llational Park where Acacia and Ccrnbret!,nwooded grassl.nds, intergpersed with svramps,predomTiE i- (A,rent6 Gillett 1975), But their top€raphy ls dissimilar. Vhereasthe Itlerustudy was centred on the baslcally flat plains arolnd Rainkombein the centre of the Park, the Tsavo study |{as centred on the rugged Ngulia Rangewhich .ises iteeply from 750 rn (2&50.) to a height of t820 m (5970'). Prey ln the two areas were rllch the same,wlth dlkdlk and gar,re birds conmonand lhpala 6nd lesser kudu present in both, but were noticeably nnre plentlful in the rocky itgulias.

2.3. TTATERIALS ANDIIETHOOS

Resident leopaads were trapped In metal box traps measuring 2.0m X 0.6mX 0,9rn(6*' X 2, X 3,) to which they were attracted by a bait hung in a suitable tree abovethe trap. Baits consisted of an entire smalI , such as a deBdgoat or ,or, rnoreoften, of a portion of a l.rger anioralsuch as a Grantrs or ihpala. Theyvaried in weight from 4.5 kg to 45 kg (10-100lb) and were usually placed in shady evergreentrees slted .rearwaterholes or gametrails kno,rnto be used by 5- leopards, or at the foot of ravlnes, rocky outcrops' and passes between hills. After a leopard had fed on. b6it for one or, preferably, tero nights, the bait $rasDoved from the tree into the trap. A pull on the bait by a feeding aninEl releasedthe drop door. Trapped leopards i,ere trahsferred to a snal ler wocdenbox of kn.i{nweight for immobili5ation, as describedby lamiI tor} (1976). The box was then weighed from a tripod, and the I eopard was ihjected by hand with the desired amountof drug, either directly into the rumPthrough strategical ly sited holes in the end of the box, or into the root of the tail which was seized through an adjustable 2.5-7.5 cn (l-3") Sap lnder th€ door. Trappedleopards were inmobilised the norning after caDtureand were al lowedto recover alone at the site of capture- Eight leopardswerq illrnobilisedwith Sernylan (Parke-Davis E Conpany,Hounsl o,,, ,.liddlesex, ) and Acepromazine(The Boots CompanyLtd, Nottingham,€ngland) on thirteen occasionsand ten leo- pards wirh Cl-744 or Telazot (Parke-Davis6 Co.npany,Detroit, t4ichi9a., U,S.A.) on eleven occasions. Twoold rnalesdied under anaesthesia,one with Sernylanand one with Cl-744. In both casesdeeth vrescaused by overheating resulting faomdrug-induced digruption of tetnperature regulation and unhelpfully high ambientternPeratures. l{evertheless Cl-/44 was consideredto be the drug of choice (Hamilton1976) as Ser- nylan had an undesirably long narcotic action and recoveryPeriod, and a greater tendencyto raise body temperature,dePress respiration, and ca;se convulsions. Thesedrugs are cor''lparedin detail by Hamilton (1976) and Kin9, Bertratl|E Hamilton (1977). tnnbbilised leoPardswere fitted with acrYlic or machine belt radio-collars emitting Pulsedsignals frorn transmitters Powered by 2.8 volt5 sr.ippliedby m-rcuryor lithium cells, as describedby Hamilton (1976), The col lared leopardswere redio-tracked by vehicle, fron the air, and on foot, using a single portable tlodel LA-12receiver (A.V.11.Instrunent Company, Chdmpeign, lllinois, U.S.A.). llost radio-locations were obtained from a fourl/heel drive LandRover or a Toyota LandCruiser equipPedwith a twin_yagi receiving system consisting of t$,o three_eletnentantennae. The5ev,,ere mcunted with their elerents vertical on a tubular aluminiumboom which seParated thqn horizontal ly by half a !,ravelength. The boomwas suppo.ted by a vertical mastlrhich held it 3m (10') aboveground level and could be rotated throlgh 3600 faorninside the cab. Signals froo eachantenna were brought by 70 ohmcoexial cable to a conbiningcircuit contained in a peak/null box where the sumor the difference of the incominqsi9- nals could be selected by throwing a s$/itch, as describedby Andersol6 DeHoor (1971). The location of a leopard$,as determined by triangulation fron tt/o or more kno!,/npointt, or radio_tracking stations' which had been plotted on l/50,000 rnapsby plane table surveying (Clark 1955). At each point a bearing of antennadirection wa9 taken with a prisnitic compassabout 8n (26') frotn the vehicle, fer enoughto avold ihter- ference from the latterrs nagnetic field. The indicated direction was also searchedr{ith l0 X {{) binocularg, and this sometimesresulted in 5ightings of leopards. The true bearingswere plotted on l/50'000 maps overlaid by a I km X I kmgrid. fhe Point at which two or more anter_ sected representedthe leoPard's Plotted position, which was exPressed to the nearest l00n as a six figure mapreference. Coll.red leopardster. llso located froo ghe air, using a slngle yagi antennaprojectlng for*ards, rJlth lts elernentsverr;cal lv in llne, from th6 wing etruts of a Piper Cub (pA- ) or Supercub(pA:tB 150) or a Cessna 185. The aircraft },.s flc*,,n slor.rly tovards the soorce of the signal, often in a poreredgllde, until the strengrh of the signal droppedoff sharply a5 the aircraft passedover the tranimitter. Anoiher run from a dlfferent direction confI riled the anlmalrs?pogltion but was usually unnecessary. The tnethod,s potential accuracy:was proveo Dy slghting collsred leopardsfour tlmes frofl the !ir, but turbulenceend auggedterrain sofietlmes lmposedconstralnts on accuracy, particUlarly In r5avo. Leopardswere also radio-tracked on foot, using a single I'hand-heldyagi, as described by Hamilton (1975). The main objeciives of this were to locate thelr resting places, which were investigated the day after belng vacated, .nd suspectedkll ls, A .4!8 calibie Itannlicher-Steyrrifle was carrled for self-defence as , rhino, and buffalo were often encounteredat close quarters In.densebush.

2.4. RESULTS

2.4.1, Captureend radio-tracking resultg

Twelve Tsavo leopards were caught and recaught a total of 22 times in 26 months. Of the twelve caught, one old mal; di€d during Sernylananaesthesia, and the redio-collar of another functloned for only two days. The ten leopardswhich provided radio-tracking data carried functioning collars for periods of 54-554days (mean= 264 Oevs) but only three transrnittedfor more than six dpnths wlthout inter- ruption. Six of the ten acrylic rlng collars crackedand broke; on at least three occasionsacts of violence, such a9 st.uggles with other leopardsor large prey, precededbreakage. Tvrocollars which were put on too loose cameoff intact after one and 158days. Twoacrylic ring and four rnachinebelt collars malfunctlonedaftet 2-167days. Forru- nately collars could usually be replaced 6s lt provedpossible to trap five I eopards hore than once and one €s manyas flve tines. In l{eru only three leopardgwere caught: the nale that died under anaesthesiaand two fsnales, an adult and a subadult. 7he adult female's collar year 'Radio-trackingtransmitted for over a and the subadultrs for seven honths. p.oved nore difficult in l{eru becauseof the flat terrain and densebush.

2.4.2. Populat ion denslty

0nly in Tsavo were enough leopards trapp€d to give an idea of the density of the populatlon. Ihis had to be basedon nrlchspeculation as there was a striking disparity in the sex ratio of the twelve trap- ped leopardswhich comprisedten adult males, one subadult nale, anC one adult female. The probablllty of trapping nales and fenales in a ratio of ll t I was less than l1 lf snifi|alsof both 5exeswere Dresent In the populationin a I : I ratio.nd wereequally susceptible to trapplng (Binomial tt.o-tailed test, P c.005). But even individuals o; the samesex were not equally susceptible to trapping, .nd although trapping slccess srJggestedthat most, if not all, of the adult males in the luare area were caught during the study, n|ost of the females knon,n to be Present evaded capture, apparently becauseof thei. greater wariness. In fifteen renddi encountersin rhich adult or large sub- adult leopardsire re seen in the study area purely by chance,and their sexeswere determinedbeyond doubt, eight were malesand sevenwere fenales. There is no signlfici.lt difference betteen these figures and the I : I sex ratio they suggest (Binomial two-taited test, P ).05), although the samplesize of l5 is too sniall for the test to exciude sexratios of 2t1or1t2, I The leopard population of the vihole Ngulia arda, sone 400 kn:2 (154sq,mi.), wasunkhown. But it was possibleto estimatethat of a gnal|, welI defined portion of the study area conrprising130 kml or 50 squaremi les centred on NdatreHill. This !censusarear coripletell contained the hone rangesof seven radio_collared leopards (five adul: nales, one adult fernale,and one subadult male) vAich betweenthen t provided over 75? of all radio-locations. Radio-trackingpro/ed that five of these leopardsu,ere present in the censusarea in early June 1973,whi sightings indicated that the other two were al50 preseni although they h,erenot then transmitting. 8ut these were not the onlv leooardg in the censusarea. Uncollaredfemales r,rete seen there on ten occasionsand it is believed thesewidely distributed sightings representeclat least four differ nt individuals: maybemore. Although it is not knor.rnif al I were pre5erii in Jtrne1973, it is likely that they b,ereresideots. Uncollaredfe- malesand their spoor were seen in the censusarea throughoutthe study and in the samelocalities more than once. I suggest therefore that not less than five adult females, including the radio-collared Leopard7, were present in the censusarea in June 1973. Thesefigures representa density of oneadult regident leupard per 13 knz ll/5 sq,mi,), but the numberof subadultsand dep!n- dent cubs cen only be the gubject of specuiation, lt is not kno$nho,'r manyof the five fenales bel ieved to be present had cubs, althorgh Leopard7 can be discounted. Hanilton (1976) sugge5tedthat at Ieast two of these adult femalesftight have had e total of iour dependentcubs, and kie$/that there were also at least tvro independentsubadults present. Thesenunbers (five resident edult melesi five adult fe.nales,two inde_ pendentsubadults, and four dependentcubs) represent a density of one feopard/8,1 knz (n.1 sq.mi.). This can safely be regErdedas a miniiun density for the Tsavocensus area and is rnuchhigher than some of the densitiespreviously reporred: I residentadult/30 km2in vil- partu National Park, Sri Lank. (Eisenberg6 Loclhart 1972); ! residen! adult/40 krnzor I resident leopard/29k# in the Serengeti National Pdrk; and most previoug guessesat leopard densities. lt is lower, however,than densities of I resident adult,/5 k# or I ieopard to tl.5-5.0 km' reported from the RhodesHatopos National park bv Smith (1977),and of 1 adult/6.1-10.5knp in the KruoerNational Park (5ail ey, pers.comm. ). 8ut it is possible,probably likely, that the true density of leopards in the Tsavocengu9 area was greater than Hamilton (!975) suggested. The Tsavostudy was discriminatory becauseit largely ex- cluded fsnales although these rderep.esent. Eailey, working in thc Kruger, was moresuccessful and c6ught them in the ratio of 1.8 adult fenEles per adult nale (Baiiey, pers,corm.). In vierv of the social organisa!ion of the leopard, discossed later in this chapter, a pre- ponderanceof resident females ls to be expected, This phenomeiont,: alscrbeen reported in studi€s of (IakachaE Schaller 1959, Rudnai 1970,Schaller 1912, Eloff 1973a,Bertram 1978), (Sch6ller l!72), tlgers (Schaller 1957)and mountainI ions (Hornocker1969, Seidensticker et al. 1973). 8y contr.st the majority of nqnadicor rransient indivi- d.ralsussally se€ftto be rElcs. lf lt is assumedthat the ratio of resldent adult females to resldent edult Dales was In fact 1.8 : I in Tsavoand that, on the basis of data from other large cats, 551 of these fenales h-addependent cubs, the resident population might have had the fol loring tornposition: 5 re5ldent adult nlales 9 resident adult fernales,of which: 4 had no cubg 2 had lltters of 3 t 2 J had large dependentcubs (2 + I + 1) 3 indeDendentgubadul ts 'lq resident adults + l2 inrnat!re. total of 26 leoDards.

This wouldrepresent densities of I residentadulr/9.3 tm2 (l/3.6 3q.m:. or 1 resident leopard/5 knz (1/1.9 sq.ni.) |rhich are equivalent to the high densitie5 in Rhodesttatopos i,tatlonal Park and the Kruger. In conclusion the density of leopards in the Tsavocensus area vrasnot less than I resident adult/l3 krn20r I resident leopard per 8. 1 knz but could have-beenas high as t resident adult/9-3 knz or I resident leopard/5 kmz. Densities of this order rrJgt be regardeo as high for an essentially solitary large . There is rea9o1to be- I ieve (Chapter4 and Chapter6) rhat softe leopard populations in Xenya m€yattain densities of up to I leopard/2.6 kmz (t leopard per square ftile) but this is likely to be near the llmit andof onlv locaiised occurrenceand p.obably includes transients.

2, {.3. Reproduct i on

The Tsav. :ind Xeru studies provided r€rettably I ittle infor- rnationon reproduc. , In Tsavothe only collared female, Leopard7, was a youngadult t:: :ad no knovncubs during the study. Although the adult female in t. .- did have cubs, probably in ttarch 1979, attempts to find themproved frur!iess. lf, like Joy Adansonrssemi-tame leopard in Shaba, she noved her cubs every 2-t days (Adamson1980), it i5 nor surprising that they were so difficult to find. I believe froflrspoor that two were allve and accompanyingher in July but thet by Septenrber only one large cub, which | gav,r,remalned. Scanty information frorn variors sources suggegts that there is no fixed breedingseascn for leopards in either (Turnbull-Kemp1967, Eertrarh1978, Adamson1980) or South Africa (Pienaar l!51) although Eisenberg6 Lockhart (1972) found sqne evidence for seasonalbreeding in . The gestation period is about 90-100days (Zuckerman1953, Sadleir 1966, Adamson1980). Litter sizes range from one to six (Turnbull-Kanp 1967) but data f.om captives 9u99ests that tt{o or three ls usual (Zucke.mant95l; Reuther E Doherty 1958; Eisenberg6 Lockhart 1972; Harris, pers.corrn.). This eeemsro apply ln the wi ld too (Pien.ar 1963, Eertram1978, Adanson1980). nor- ever, the observationsof Turnbull-Ke|np (1967), Schatler (1972), ltuckenhirn6 [i5enberg (1973), and Bertram (1978) shc{ that usually only oaleor two well grcrh yolng are seenwith their mothersin the vrlld, Cubrprtallty r|lst therefore be hioh. The two cubs of Joy Adanrsonrsleopard were born in an almort inaccessiblecave high !p in the rocks and openedtheir eyes after a week (Adamson1980). At about three mcnthsleopard cubs are weaned (Turnbull-Kenp1967, Adamson1980) and by four nonths occasionally accompanytheir nother (this study). Although there ere records of leopardcubs ki I Iing smalI prey at five monLhsof age (Tqrnbull-l(emp 1957) the cub normally remainswith its motheruntil at least one year old (Turnbull-Kenp1967) and probably for as loog ad l8 rnonths to two years (Turnbrjll-Kemp 1967, Eertram1980). At 2l rnonthsJoy Adamson'sleopard cameinto oestrus and at 26 monthsstarted to mate. In thc Serenge(iBertrem's radio-collared fenele cub left her hother at the age of about 2l months,when her rnothercaoe on heat again and rnatedintermittently, but rernained at least temporarily in the sane area. By contrast the Tsavosubadult rnaleleft hi5 juvenile ho.nerange at the eslinated aee of 30 monthsand was never seen there again.

2. c. {. €xtent of moveirent

The nlovenrentsof radio-collared Tsavo leopardsare desc- ribed in de!ail by Hernilton{1976). Thefo)lc'ring is merelya simPli- fied sufirn3ryghorn of s!atistica] analysis and other complications. The knounextent of nbvementof ten radio-collared Tsavo leopardsduring the study is sho,n in Fig.2.l. 6rd Table2, although it almost certainly under-representsthe movementsthey actually tnade. I have used the terrn robservedrange' to represent thc tot.l area over which a I eopardwas kncrrnto roamwhile its col lar w:s functioning, lt !.Jascalculated by the mininumarea method(!:ll'e 19q2) in which the o!termost points oi loc6tion are joined together. Observed rangesof the ten I eopardsvari ed I to-'r22.6-120.5 kmz (8.7- 46.6 sq.ni.) anderere not necessarilythe same6s tlre anin'als'hor.c ranges. The 'homerange'was the area over $rhicha leopard nortnally travelled in rtersuitof its routine ectivities anddid not includ! forayg or journeyg outside this area. In nost anirnalmovement studies the irinimumarea fiethod i9 used to determinethe size and shapt of the hor,eranae, but it is to someextent €rtificial and inflexible be.ause it is basedon georietric connectionof the outermostPoints of location and nay often include areas where the aninEl was not known to go. There is fiuch to be said for applying a methodbased upon consideredappraisal by the researcher: one which, although baseJ on the minirnumarea |nethod,takes into account the spatial arrangeroentof all points of location, topographyof the habitat, and all aveilable knovledgeot the aninaJ'stravel routes, habits, and bchaviour, This is essentiaily an expangionof the ratypical habit3t el imination me!hod' suggestedby Ables (1969)but I heve used the ternl !adjusted hone ranger here. Providedthat any tenPtation to mekethe boundarresfit in r.,ith preconceivedideas is resisted, I bel ieve th6t edjusted homeranqe more closely reprcsents the size and shapeof leopardsr homeranqes than other nethods, Criticism that it is too subjective can be counteredby oointing out that the geonetric meth.ds are artificial, ioflexible, andwaste information. The adjusted hon',eranges^of sevcn adult male teopards rangedin size fr(]x 19.7 to 59.3 knt t7.6-22.9sq.mi.) with an ave- rage of 30.5 k'iz (1i.8 sq,mi.), as shdrn in Table2 aft Fi9.2.2, The hr':]eranges of the only colla.ed lsavo femdl€ anCof the suaadult ?<--t- -{'?'\

'/'' '/v \ (P \--"------_ t. zZ--=, /zZ-=,-:z*.-j ;-\;;;.i.'"-..:i\3 l+b= '*:-r\-vl 7l ;f;F: /, \ 7 li'H:f-j\.Ji O) I J, 12

F \<1\r=\S b..aqP= \

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r€

a o - l0 -

malewere much sBaller: t3.9 km2(5.4 sq.nri.)and lO.2 kmz(3.9 sq.mi.) respectively. ln. conparisonthe ho|nerange of the lleru adr.rlifemale coveredZb,7 km. (10.3^5q.Illi.), while that of the }leru subadulr fe_ (5.9 1':al:cqprised 15.3 kmz sq.rni,) fron October-December1977 and 32.5 k'n,z(12.5 sq.mi.) f rornJanuary-4p.;1 1978. I believe theie airre- rences reflect the lov{erdensity of leopards in Neru. I l.{otall the Tsavo leopards rernained$,ithin thelr orn nome rangegduring the study period, Three adult alalesand the female hade detected forays outside. Leopard2, for example.made eioht kno{n forays between flarch 1972 and Septemberl!/1. itrese varied in duration from one day to two monthsbut usually lasted frorn 1-l weeks and N/ereseparated by intervals of from foor diys to elevenweeks. F Theyusually follaned a sifiilar pattern of movement,taking him west- wards to the l€ul;as. 0n at least two occasionstrli forayi imnediatety fol loJed hostile encounterswith male leopards8 and 3. iwo other males, 8 and I, borh.madetarge kiJls duiing their fliays, wbicr,nigf,t h.ve .l Deenmade for this_ purpose. The female, Leopard/, madeonly tio Knctrntorays in lq months but in both she eccompanieda neighbouring"she adult male, Leopard3, into his hdne range for a fervdays; was probably in oestrus and nay havemated with him. Neither of the t\,volleru females madeknern forays outside their homeranges. Hcnever,the homerange of the subaduli, whowas living on her ovn, did 5hift from the Rai;kombearea, where she was probably born, to the eastern portion of the park: ; changethat may nave representedthe dispersal of a younganimal seekinq to establish her adult.honerangu. Similarly, Leopard6, the Tsavoiubadult mare, jlvenile_horne left his rangeat the.9e of abour30 nonrhs, He initialty novedsome l5 km (9,3 miles) to the northwesr, but failure of hrs radro_ coll-, halted further monitoring of his search for a nev homerange.

2,4.5. occupancyof the homerange

Leopardsh,ere not found in all parts of their homeranges equally often. Therewere usually sectors of concentrateduse: on averageabotlt 75Uof a leopard's locations fell within about 50t of its hone range. 8ut although leopardsdid spendmore time in cerrdrn areas, in general they coveredthe greater part of their homeranges frequently and thoroughly. This is well iliustrated by Leopardl, the best studied Tsavo male and one whose movementsapoeared to be typical. Fi9.2.3. shor.rshis Dovementsby night and day over a period of 95 hours during which he rravelled a lnininumof 43.i km (27.i miles) or an averageof 10.9 km (6.8 miles) per day, hi5 moveientstaking place |nostly betlreen1700 and t000 hours. In those four davs he madeno le99 than- three ci rcuits of Ndawehil l and coveredih" o.""r". pa.t of his 30 kmr ho.nerange. Similar 3-4 dey studies on orhe; occasionsrevealed mrch the semepattern. The;e flndinos are at variance v{ith previous be!iefs, such as rhat of Astley iabertey (1962) I'l'lal who lvrote: es hunt over a wide area of their chosen ranoe: seldomrsnaining In one portion of it for fipre than a ft* dais, :ay a week, et a time, before movingon, and so, in the course of a monrn or so, patrolling the wholebeat.tl Radio-collaredleopards sho,redfew differences in their seasonaluse of the homerange. Therewas a slight tendencyfor rnost of them to increase the size of the homerange in the wet season,Dur in the case of only one leopard, Tsavoadult mrle 5, was there a ['

d\ EE

= hUE

o E,,

;

; stati5tical ly significant dlfference. But although he extendedhis rangeby about 408 lp the wet seasons,there was no narkedshif! th location and he remainedin nrch the samearea. SorneTsavo leopards also sho.reda tendency towards more even use of the hofie range in the wet season. The trees and shrubs on the plains r,verethen ln full leaf and their foliage providcJ innumerablerhady resting places that probably encouragefthis, for the leopards then had less need to seek shade and sheltei amonqtne rocks and evergreenson the hil lsides and kopjes.

2.4.5- Activ i ty periods

Radio-collared leopards rnovednostly at night and rested dsring the hotter hours of the day (1000-t700h) when in both Tsavo and l'leru shade temperatures were usually betereen29-33oC (84-92op;. 8ut, as in the Serengeti (Bertram1978), they eooetinesmoved e. an) tir|rc of day, and ofl two occasions I sa\,,,leopards hun!ing at 0930 anC 1500h on hot days. usually, horever, there was little movenentbe- tween1000 and 1700h, a sharp increase betkeen 1800and 1900h (dusk), moreor less continuousnovement throughout the night, although the animai might stay in the sane place for several hours, and a gradual reduction between0800 and 1000h. Similarty although Tsavoieoparos sonetinrescalled at any tine of day, they did so rnostlyduring the night and arounddusk and dawn. In lleru they were geldomheard-

2.4.7. Restinqpl aces

Soneauthorities (e.9. Bere 1962) havewritten of leopards' 'lairs', a lair.rrmally beingtaken to meana resting piaaeuseC frequently and regularly by the sameindividual. But my radio- col lared leopardsusual ly rested in a different place eachday excep! vrhenthey had large kills or, in the case of the l,1eruadult female, soalI cubs. Theydid, hoi/ever,sometimes return to the semetree or rock in the courgeof time and seenedto have somefavoured resting places, just as they preferred certain gametrails and vehicle tracks a5 rout e9 of travel. The straight-line distance betweenresting sites on con- 'daily secutive days, the distancer, varied frorn leopard to ieopard moreor less in direct proportion to the size of the animelts . For examplethe meandaily distance of the subadutt Tsavo mele,vrith the smailesthome range, was only 0.9 krn(0.6 miles). That of the adult femalewas 2.0 km (1.2 miles), vrhile thoseof five adlft mafesvaried from 2.3 to \.2 km (1.4-2.6miles) Hith a mean for ill five malesof 2.9 km (1.8 miles). Themaxiftrm daily dis- tances recordedfor each leopardlrithin its hone rangevaried in much the samet{ay: fror 1.9 km {1.2 miles) for the subadult male and 4.9 km (3.0 ftiles) for the adult fsnale ro 8.4 km (5.2 niles) for the adutr male with the largest homerenge. Thesefigures suggest that adult male leopardstend to travel over iarger areag than fefieles or subadults. Althouqh|teru provided no comparativedata for rnalesand females the samerelation- ship frobably applied there too. The hean daily di5tance for the 5ub- adult fefiale before she movedeastlrards vras 1.2 km and for the adult fenale 1.3 km. The meanfor the edult fenale is I or,/erthan expected

and_probably reflects a reducedtendency to travel while she had her a teopards, hc :nl]-:rf:. .For ever, the da y distance disguises of ttavel actually undertakenduring lhe night, as Section z!{.r.5nc*rs-:n::r*:t

Radio-collared_leopards in both study areas used rocks, as resting places, ln Tsavothere were interesting and.9tatisticatty:l:":: :i9^lrtl:. significant seasonaldifferences ih their use of rocks and trees. In the dry 5ea9on, when the deciduous trees €no 5nruDs-hacttost their leaves, leopardssh@red a significant prete- rence.for,rocky resting places wherecaves, overhaigs, and d'ense gro$/rns or evergreen trees and shrub5 provided deep shade and con_ ":-l"ll soodvantase pointi andsafety'fron Jiitu.u"n"u uy :i:]11l:- other.:s big game. :::?:"1:::11 -The leopardswere then atso straregi_ - in the right ptace - ::l]y_1::":1 andet the right time for priyins ypo:r rocx-hyraxeswhen they eynergedfor their eveningfeedinq period rJare r)o), and I saw them take advantageof this, In the wet season leoPards were more often seen in trees ihan arongst rocks, d;;pite the difficulty of seeing themamidst the densefo aie. itreii i"riu.it" choices in Tsavowere the deciduousgaobab, Adanionia diqitata. and both.of were then in Tiif-TETI--fi-Heru reopards Slts_yrl_l9Fj,were _which most often seen in Acacie tortills, the conrnonestlarge tree in the study area.

Bushesand thickets v,,erefrequently chorenas resting places in both Tsavoand l{eru, but the I eopard;were then seldomvi;ibte and their presencewas determinedby d cor,lbinationof radio-tracking on foot and investigation the follo{ing day, after the anirnalhad moved on. In the vret seasonalmost any bush in leaf sufficed but ii tne day.seison there was a preferencefor evergreenssuch as goscia "19 l'laefuafirFii. Leopardsresting r.,nde.u,s neiiliil ry ++::99curreo up cto5e to the base and were well concealed. Kills eaten on draggedinto bushestoo. In fact.onii.ry .o popular_bel:!: liorld.y.:",rsually (e.9. ief oorst E Dandelot 1970, yeigel l97j) less tnan one- third of the large kills I found in Ts.vo and li.u we." c"..i.a up Into trees: probably becauseneither hyaenasnor jackals were conmon.

2.4.8. Feedinqecoloqy

.Analysis of the changesin day-to-day locations of Tsavo I eoParcts shcir/edthat on only l3Z Of all consecutivedays on which indi_ vrOuar teopardswere located were they found ih the sane place as on the precedingd6y. This In itself inplied 'large'klll that they were nrakingfew large kills: a beingani that detaini tt., fo.rnor" tnan a ctayand occasionally included aninals as small as dikdik and durker. Radio-trackingdata given by Hanilton (1976) indicated that the average,adultoale.Tsavo leopard madeabout lg large kills a year {range 9-33) and that individuels sometlmeswent for p;riods of 2-3 weets, If not npre, witholt feeding on large prey. In lteru ihe kill rale eppearedto.be_rather higher: of the order of 2g large kills a year tor the edult f€rnale: a difference prob.bty related io difference, in the availabil i ty of prey. The suggegtion frorn novementdata that Tsavo leooards must have been subsisting largely on small prey r{as confirmedby faecal dnalysis. Thls reveatedthat ,9t of the 5l sanples containedthe rsnains of smalI rodents such as and Arvicanthls and insectivores 5uch-e5 _Jateri lrocidura; 278 conrained the hesygs;l;;fE;;d hai r of smatt anteropes such as dikdik, grey duiker, and steinbok; and 27t contarned rne rematns.orground birds such as the yelloi_neckedspurfoul (pternistis leucoscepr.rs). others conteined the renains .f ,.;k-;y;:;; ;;.ri;ffi 13-

baboon,small snakesand lizards, and 6 variety of arthropodginclr.lding tetigonid grasshoppersand the large turqoise centlpedeScolopendra. In l{eru so f e!, teopard droppings vrere found (another i nd'iiEi'i6i--6i a sparse population) that faecal analysis was not attefipted. lhereas no less than 822 of the Tsavo faecal sBmplescon- talned remnantgof rodents, hares, hyraxes, small , and birds, only l22 contained identifiable rernainsof the larger aoielopes. The3e animals, however,forned the majority of the thirty knorn leopard kills found during the study. 0f the 23 kills, elevenvrere , four were bushbuck,three $,eredikdik, tr,/owere les5er kudu, two $lere grey duiker, and one t/as a . The other sevenkills comprised two lian cubs, a younggiraffe, a wartSog, a vervet rrbnkey,a rock t hyrax and an African hare. The seven knownkills madeby l'leru leo- pards included impala, bushbuck,lesser kudu, Grantrs gazelle, water- buck, and a baboon. Noneof the kills I examinedappeared to be in ooor condit i on. It is clear that in the case of the Tsavoleopards exani- nation of merely the kills that were discov€redgives a rnisleadingin- pres5ionof the animalsr diet. ||ost of the large ki lls r.rerefound vrhen I inve5tigated a leopardrs tetnporarycegsation of dai ly rnovsnent- Small kills such as birds, hares and rodents that did not require days to eat stood no chanceof Deingdetected in this way. lt i5 undoubtedly becauselarge kills are nEre likely to attract attention that nost papers discussing predetion by leopardstend to list the larger i t€rns at the expenseof the smaller (e.9. Wright 1960,Hitchell et al. 1955, Kruuk€ Turner 1967, Pienaar 1969). Neverthelessthe diet of leopards do:s vary fron place to place dependingupon the prey available and proba5ly also thc size of the leopards,whoge adult weight can range in Kenyaalone from 30 kg (551b) to 95 kS (209 lb). In the Seren- geti, fo. example, the prey species nrost coflrsbnI y caught by leopards are impalasand (Bertram1978) but that is an exceptionally rich habitat with an abundanceof large prey end the Ieopardsthere are also larger than those in Tsavo (Eertranr,pers.comn.). I bel ieve lhat tbe feeiing ecology of Tsavo leopards i5.probably nore typical, at ieae! for rhe smallish leopards In the 30-50 kg size range that in- habit the Kenyanbushlands where large p.ey are less abundant. lt is interesting that leopards in the Rhodeshatopos National Park also seemto feed extensively on rock hyrax, kl ipspringer, duiker, hares, rodents, and gamebirds (crobler 6 tJil5on 1972, 5rnith1977). It is also of interest that baboorsand the various species of wild pig that are so often supposedto be the favourite prey of lec_ pards (vright 1960, Eere 1962, Snlitherst966, Dorsr 6 Dandelot1970, Veigcl 1975)seldonr featured in my Tsavoand lleru studies or in the ki ll recordsgiven b) Vright (1960),14itchel I et ai. (1965),Kruuk E Turner (1967),Pienaar (1969), schaller (1972),and Bertrari,(r978), The leopard maybe the baboon!schief natural enemy(Vright 1950, Smiles 1961, Astley ltaberley 1952, Eere 1962) although this is de- batable (GaneDept 1953-54,Guggisberg 1951) but it does not nece- ssarily follc^r that the baboonis the leoDardrschief prey, and fe$r professional hunte.s in Kenyaeven consider tlsln9 baboonfor bait. tor it i9 seldomsr.rccessful. Uhile someleopards do 5pecialise in hunting them, as on llount Suswaln Kenya(Sinons 1965), this is probably for t.hevery good reasonthat little other food is available, for - and pig - are forrnidableprey whosecapture is not without risk, IndeedEertrar0rs radio-collared leopards in the Serengetiand Joy Adamsonrs sejni-tame fenale ln Shabal{ational Re5erve,Xenya, usual ly avoided baboonswhenever the two specles,net (8ertram 1978, t980). Adamson

crvnfeel ing, ard that of nany gamewardens and ressronat_hunters. .lly oro_ l-have spokento in Kenya, i9 that the betief that eopards teed .r targely on baboonsahd pig and therebl keep their num_ 9:I:99"n {Beret962, Dorst a Dandetoii970, c"r" otpr.-ilii, ts:0, 1932-_t\, 1935, 1937, 1950) has.becornepart of the tdoparaii-r,,ytnorogy. OneGame Depa.tmentreport (t930) obseives, fo. e^"rpi.,- illl rnais_ crrmrnate staughter of leopardsduring 1929and 1930has upset numDers the.resutt that pi9..,,have :.i?,1I,.11.n increasedconsiderably.,, 5ucnan tncreasein sucha short period of time is biologically probable! inr- to say the least, and a better explanationiay'le-iounc in the gpread of humansettlement into the natural habitat of the ,verminr and the attractioi exerted on themby cultivation; it"ii numbers"roura probably-still appear to be increasing even in the presenceof a substantial leopard population. Indeedthe lreetop; Saiient ih the ADeroareshas. the dengest population of e,arthog,, and giant torest hog I have ever seen in Xenyadespite hivlng als6 a denie popu_ lation of unusually large leopards. tn conclusionl yrhite it is cer_ tarnly true thal leopardsdo take baboornand pig, particulariy the young whenthey set the opportunity, and ttrerif;ie obviously iavc some errect on their popuiations, I bel ieve this effect bas been e-xaggerareo and is less than popular bel ief supposes. There was little information on the food intake of wild African leopardsbefore iry Tsavogtudy apart from Turnbull-Kemp,s(1967) estinates for the anounts of neat coniumedby Rhodesian leooi.js in 2lrhours: thesevaried fron 8.1 to tl,6 tg irZ.g:jg.8 rti.'-ii rr"uo, hcr,,ever,it was possible to measlre the daily food intake of radio_ ieoe:rds of kncr.rnweight ::lf:l:d feeding on baits of knc$,nweight. of,meat consumedranged from 2.0 kg (4.t tb) to 9.5 (9 il::.::1,:tDt. per teopardper :zv.) night, with a meanof 6,3 kg (13.9 lb).- rxpresseda5 percentagesof the body weights of individual leopards, !ne ar,ounts.eatenin one night represented(-24t of bojy weighi, witlr a meanof l6i. Thirteen (621) of_the meals representedl3_lii of body weight (t9?) and tour exceeded203 of body weighr. As th-e amountof bait remainingwas never a limiting factoi it ian be assumJ iiat tne leopards took as muchas they wanted. Thesefindinqs show that the reopard, I ike the I ion (Schatter 1972)and (S;hatter 1967), na" eat as rnuchas 202 of its body weight in 24 hor-rrs. it was possible to makesone rough estimatesof the.. .. _ln_"9dition daily food intake of Tsavo leoparls of kncNnweight feedinq on therr o!,/nkills. Thooghthe kills could not be weighed, tfreir aooroiimate e,eightscould be derived fron various published sou.ces (".g. Ledge, 1954,sachs 1967) ana radio-t.ackin9 aata s troreu t*-io"g-iiu i"opuro. spent with each. Data from six lnpila, bushbuckand lesier kudu kilts I examinedsuggested that the leopd.ds consumedfrdn 6.g to l0.g k9 per.day (15.0-23.8 tbs), with a meanof g.l kg,/day(17.9 lblday) over periods of 3-5 days. The five teopardsweignea zi-42 k; (64_9i tb) with,a meanN,eight of l/.4 k9 (82.5 lb) ana-tneir-esiiiGtJ aliry too: fnrakes expressed^aspercentages of body rveight rangedf ro{,r17 to 26r, with a meanof 23t. This substantial intake concuriwith rhe therrs tlat.:::l'ivo:es ',a -or -fami";-l..g are adapt€dro feast i,;;i iC.t i"v ".. at..t>o>),. the suggestionthat leopardsare r,asteful feeders (Smiles.Lontrary.to t95l), those in Tsavoand lleru invariably nade the - r5- most of.ny large kill. They continuedto feed .fter putrefaction rra5 well advancedand were efficient.nd tidy feeders that took care to conceal their prey from scavengersand left llttle by the time they had f in ished. In conclusioi the leopard has a fl|orecatholic diet than eny of the other large cats. l'ly studies and a survey of thg llterature cited earlier shcrr that the African I eop6rd feeds upon o/er fifty species of wild r|alrmals(excluding lnsectivores and rodents) ln addi- tion to domestic and, occasionally, r€n hinself. Birds ranging in size frdrl starlings to ostrich, and includlng poultry, also feature In its diet, a9 well es crocodiles,snakes, lizards, emphibia, fish, snails, crabs, and insectg; indeedsofie leopardshave been kno{n to specialise in fishing and crab-eatlng (Turnbull-xemp1967). 0rass is sonetimeseaten intentionally (personalobservation) and wild fruits mayoccasionel ly be taken (Lyell, in Tlrnbull-Kemp1967). l,loreover, leopardsoften scavenge,contrary to the suggestion (Weigel 1975) that they only do so in an einergency. As they r,,lll feed on ani- mals such as elephsnt (personalobservatlon) that hardly constitut€ their norrnal prey. And flry Tgavo subadult filale which llved near Ngulia lodge often usecito forage ih the rubbish dunpafter dusk, after sperC- ing the day less than 200 m fron the gtaff quarters; on one occasion he was foond late one night rudmagingin waste bins in the kitchen which he hed entered through a terporarlly missing panel in the door: Indeedit can truthfully be sald that the I eopard$rill eat alt'lostany animal,and this adaptability ls perhapsthe speciesrgreateststrength apart from its gecretive habits. Eecausethe leopard ha5 such a wide and varied diet it is less serlously affected than other large predatorg by the decline or disappearanceof populationsof any one, or even seve- ral, of lts prey species. This tbuffer effectr, together with the animalrssecretive habits, largely exprainsthe leopardrsability to exist in areas of dense hunansettleneot. lt i5 an importantaspec! therefore of the speciesr rsurvlval ecologyr. Another irnportantaspect is the leopard's ability to survive for long period5 (1.e. several rnonths)withou! water, as they appedr to do in Sotswana(Snrithers 1956) ana parts of Kenya(this stuay). It ls not true that they drlnk every day (Dorst a Dandelor1970). tly Tsavoobservations suggest that wherewater is available they probably do drink every two or three days but do not need to do so. There wes no water in one leopardts homerange ln the dry season and he was not kn6/n to visit the nearest spring. Like the Kalahari which may becofiecompletely indeperdentof rater under desert conditions (Eloff 1973b), the leopardrs prectice of resting during the heat of the da./ and dpving dpgtly at night adapts lt !.elI to I iving in arid vraterless envi ronments.

2.!.9. Patternsof dlspersion

Fi9.2.2. shoJed that the homeranges of Tsavo leopards we,e not arranged haphazardly in space but with a degree of order that vras haintained during the short time gpan of the study. The horneranges of the adult maleswere arranged In a nosaic uponnhlch thore of the eub.dult male (Leopard5) and the adult fsnale (Leopard7) were super- imposedsepar€tely. The [bst interesting feature of Fi9.2.2. is not the generally small degreeof overlap betweenthe homeranges of thc adult aales but the way In vJhichthe boundariesof adiacent hdne rangas corresFond with cach other. Leopards2 and 8, the two adr.jlt gane ftales tha! shared the 6rea, constitute a speciai case thaa lrill be examlnedlater. 8ut cven their hooe ranges illustrate the phcno_ Denonof correspondence, for both Indented;est of xavu hilt in tne sameplace. thig Indentatlon rari rnorc or le5s parallel to Kavuand may have been partly Influenced by topography. gut what is interesr_ Ing ls th.t the eastern boqndary of teopari i,s hope range tilieo into it without extendlngfurther ea;twards.' there alsi "pp"ii"a io o" reasonably disttnct boundaries between the hone ranges of leopards 3 6nd 9' 3 and 2, and J and ll. Vhere Leopard3rs hde ranqedia over- Iap those of the other adqlt lnales, the overlap may be exiaoerated because he provided more radio-locatlons ovea a lohger perlfo than any other leopardahd.l I his .rovements,including thosi on the periphery of hig ho.nerange, are better reDresenteo. The spatial arrangenentof ho,neranges and the dispersion or Inqtvtouats can be expregsedquantitatively by calculatina for €ach leopard the-tgeonetric centre' (flech, 'centre leitei e varner ti66j or of actlvityr (Haynel9t9) of all its locations rrithin the n9...-l:nS" and by comparlng thc dlstances between the geonetric centres of-different leop.rds. Thi5 shd./edthat for six neiqh6ou.iiq aautt nales the averagedistance from the gecrnetriccentre of the iocations of each of them to that of the neareit kncnn adult."i" "", j.i r.rno. 1.9 mlles (range 0.8-tr.6 kn). Ihe distance betN,eenthe qeom'eiiic centres of the adult.f€rnaleLeopard 7 and the nearest adrllt male tr.u |(lnor u.o nites, lressub9tantially less. Anothermeasure of dlsperslon is r inter-lndividual (Hamilton distance' 1976): the linear dlstance betweenany t*o teopirar-at tte sane tine. The Tsavo data shored that an adult fi|ale lras normalry separatedfrom his nearest knd^,nadult male neighbourby an average (2.4 of-3.8 kn rniles). Eventhe adult males2 ind g were on ar..ug" (1.7 2.8 km niles) apart ltthough they shared over 70t of ttreir frome ranges.and.thegeometric centres of their locations were only 0.g km (u.) mrres, apert; In other words they appearedto be actively avoid_ ing e.ch. other. ltips of the lntensity of use of ttrel . toru ..ig., shored thit they did concentrate thelr activities in different a.",. and the separatlon of the two anlmals in tine and space iii;.;.;;;;-' Leyhausenrspoint (1965) that the ccnmonuse of an area does not nece- ssarily nean sinultaneoususe. lt is also of interest that Leopara/, the.only collar€d 6dult fsnale, was normally separatedfrom aduit rnate .t.7 ll by 2.2 kr (1,4 nlles) andfrom subadultmale 6 by km (t.l miies) although her hole range overlappedextensively with'theirs. xo"ever, concentrationof the activltie5 of leopards/ and 6 in muchthe same area su99ests_aconsiderable |tr]tual tolerance.

2,4.10, Patterns of asgociatlon

. I'ly.observat ions In Tsavo and Heru confl dned the flndings of Bertr.m (1978)In-the serengetlrhat leopards"r" U".i"itiy,otitiry aninals and are almost alwaysalone. In Tsavog7g of my l!0 sightings of leopard were of leopards sean alone, and on B8l of the occasions nhen my baits rJere fed upon by leopards oily one anlmal was Dresent. leru .ln too I only saw my adult female alone e"\cept when accompanied by her cub. 8ut for several reasonsradio-tracking did not reveal the true fr?quencyof association betweenindividual t;opards: not att the lcopards in the study 6reas v,ere radio-collared; not all the - 17 -

collared I eopards transmitted at the sameti,nei and the freguency of radio-tracking |vashot sufficiently continuousfor the ctose ionitoring required to detect €ll, or even nogt, associations betweencollered leopards. Eocounters betb/eentherrr, particularly O.t"een ,ai.i, *.r" often brief and stood tittle chanceit O"ing a"i..tJ.- o"loii" tni, the technique providedsorne interesting infirmation *- tf,r-l"i"tion_-'- ships betweenindividuais of both sexei. l The relationship betweenadult males appearedto be one nuruar of Intoterance, even hos!ility. Hamilton (1976)gives evidence of six suspected conflicts betweenLeopard Z "na-i"opi.Oi f-"na g. In three of lhese encounters,which all took place ii., ti" 6ouio".y o. overlap. areas betweenneighbouring hone ranges, ."Oio_i"il".i *"r. ertner Drokenor cameoff intact. Therewa9 also evidencethar mooe_ rately severe injuries were so.netimessustalned by the coflrbatanrs: rn at least tliro encqrnters leopards 3 and 8 were bitien on the heaOand clawed oo the head, neck and forequarters, and ti,o hours after another encounter Leopard2 was seen I loping badty with a cut or 9pl it fore- pdw.. The infliction of deepcanine toothrnarkson the heai'anj tle Dreakage of acryl ic col lars Indicate bite5 of considerableforce. At_ though lhe woundseventually healed, sometlmes.lte., p".iJ ot suppuration (LeoDard 8), it is not surprising thet such bites sor1rlerImes result in death. Voodley, for o(arirp;. ,.poits a tatat encountg oe_ tweentwo malesthat met on 6 forest ga,netrail; the youngeranimal was_killed by a bite rhat crushedthe (}{ountainNati6nat part