Deakin Research Online This is the published version: Costa, Daniel, Weise, Michael and Arnould, John 2006, Potential influences of whaling on the status and trends of pinniped populations, in Whales, whaling and ocean ecosystems, University of California Press, Berkeley, Calif., pp.344-359. Available from Deakin Research Online:

http://hdl.handle.net/10536/DRO/DU:30008326

Reproduced with the kind permission of the copyright owner. Copyright: 2006, University of California Press.

TWENTY-SEVEN

lotentiallnfluences of Whaling on the Status and Trends of Pinniped Populations

~NiEL P. COSTA, MICHAEL J. WEISE, AND JOHN P. Y. ARNOUlD

Although this volume focuses on whales and whaling, the depletion of great whales over the last SO to 150 years per­turbed the marine interaction web, thus influencing many other species and ecosystem processes (Estes, Chapter 1 of this volume; Paine, Chapter 2 of this volume). Such interac­tion web effects have been hypothesized for several pinniped species. For example, the reduction of great whales in the Southern Ocean may have caused seal and penguin pOFfUla­tions to increase because of reduced competition for their shared prey, krill (Laws 1977; Ballance et a1., Chapter 17 of this volume). In addition, pinnipeds share some of the same predators, especially killer whales, as large whales do. Declines in whale populations may thus have caused the decline of certain pinniped populations because of redirected predation by killer whales (Springer et al. 2003; Branch and Williams, Chapter 20 of this volume). These purported indi­rect effects of whales on pinnipeds are poorly documented and controversiaL Since most of the arguments are area- or species-specific, a global overview of the known patterns and causes for pinniped population change is topical and relevant.

Because of differences in body size and life history, pinnipeds are both easier to study, and possibly more sensitive to environmental fluctuations, than most cetaceans

344

are. Pinnipeds are I to 2 orders of magnitude smaller in masS than whales, which result in greater mass-specific rates of food consumption. Thus the pinnipeds have physiological and environmental scaling functions that must be consid­erably different from those of the great whales. For example, although some pinnipeds have remarkable abilities to fast, even the most extreme durations of fasting in pinnipeds fall easily within the abilities of large cetaceans. The relatively small size of pinnipeds compared with cetaceans results in a much higher mass-specific metabolism and thus a shorter fasting duration. These differences should constrain pin­nipeds to operate at smaller spatial and temporal scales than the large cetaceans, thus making pinnipeds more sen­sitive to variations in prey abundance and distribution. Smaller size is also linked to a shorter generation time in pinnipeds, which makes their populations more vulnerable to environmental disturbances but also affords them a greater potential for population growth. All of these char­acteristics suggest that pinniped populations should be more responsive to changes in their environment than the large whales are.

Pinnipeds have a nearly cosmopolitan distribution in the world oceans, although most species occur in temper­ate to polar regions. Abundances range across species from

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Tfl/Jt1(! hy a'w lil\e .l~)!iHh (lvliy,wd,l 2tJfl2) Thh 'lweIP'; i,

f*',\j'nOy cvmj.dl'f"d tu be ()l1;:, j,)[ ll\(" tWf;nlj' milst lfll.\'aklwd

HlllrJm~ nwmmah ill til{' wmld. Tlw Caspian s.?,]! P'!pii!Hth)Jl d,'cfiIW WN; ,1 .r;')f\'<;{]\"'H(': \')i O"I'.l'C"ph,Hati'Jl1· 115,000 t1.,l J74,OllU I la,,: beVllllill'VI";t<'ri \1'1(' lho"' ,,[n!y ilim~k!;'nlh cuilmy \\1jv~ll;Jkj );U02) , fl],2' lkdinc 01

C:aspian seah 'iNH$ C~~~h\<\'ft)a('{;d hY-;1 l,{FI~~\ HIOd~1iity k:Vt1nl uf "P!7.uOlh m!t\ll1 In 19'>7, w!ltd.! kllkd Sl.~\:nil! lhomand all1"

mnls, 'HIt' l.k1ik,;! Will hij, ilb;(l (kdilll'd !>t0MJi1y in H'(\'H( Y''<li\,

from ahOll1' 7(),OOO ;]!1lfn;;b ill l'Ilt lYlU:; to ahout :'>,UHO ,mim,lls

IT'

11.0 h:ntly, 0,ji\>~ ml.'l't,ditk, fflnn nwri)iilivims (LiI,tH'jshw;lj'

t't aL 19E9) nCo:JHTdJ in l'.lg:?~,l',if,.s ".mi i.n 1.998,

HadlUr ~pajs OCClll Iddt'ly in ('oasta!, (',tHClr!!'!,,, ;md o(~[l$ion·

;dly frohwatcr h;INt"b iKl'm" !h'" Norlh Atlllntk ~ilid P;lci!k

U(.";ws, Till' nOIJmigratory fWtl)l'(' uf thi:> ilpp;mmtly hm, [,;sull(xl in (omhltt"bk r"!llomll llcncti( diffNenUatl(m,

1 i,lt'i)oJ' se:~11 popul;!tion Vt'nds YiH)' h'lddy t'kpi'l1ding upon :ll('n and !'!"bitil(, POj.1ulatJollS Me ;vt:;u' in C1:l1Howl<I, Of('gon, and

at J,y,);to fIN, Pl'!

<",'jn"""'",, (,Idfri<'S <:t ai,

l'Jlf?; Cal [ell" ct ;;t, :WDll. TIl(;,,' illn~(r~i,') (:1)lill';1St wW, tL.:di{le~, of 6.;;:~!}it if) }?')!;'-;j dl~ri~lg the "I 9/t)·:; ~llHj 19B1b

ill ttl<: (;\llf o{ Ali\',k,'l, Prinei;' Vv'iHiam Sound, Mld 1h .. , lh;ring :)(:;\ (pi\dh't 1990; Small d al. 2(J(H; ~<:,: Hgml' ?J.:'l), Il;lrhor

5('a1 nmnh(,h HI ttlt' w('stern NOrth i\lhIlHi( !Iii'll' gcm:uJiy

ht'{'n !;ignH!uH'!1 ){'t<lf ,kdiu;:" For tXilHlj!Jc, il)<: JltllHbt:1' of

l'im'IMI' ,elli !lujJ~ bt'fn on SahlI' Isl;md de\linl'ct hy;)bon1 '15%

hl::hv",'!l 19WJ ;mlll <)~)j {1\0,,"'('11 ,'1 ;,1, 2f)(H), <tpp,)ft'IlHy fwm

predation!;y slwrb (L\l(:a~ ami StGbo (md olmpHilio!1 with i\ rilpWly growing gr,l)' ~C'd! pOpU!il!i!)J1 (jlUWC)1 Nil!.

:rmn), j'brbor ;;cal mHlih\'r<; f1i;\\,'t,' ;.;hv dedlrwd 5ubst;mtial1y

in tlw Fil$Wf!l ;\.tl:mtk:, but lJ\'f(', (h" "p]I<U'ent GlUSt: w,~" Co

phm.·iIH·' 199;';

"pi(li"ll1ic {1kld(·-.Injw'n~en Gnd r!ilrk(lJwll "\t\;,~!",,,'''i'1''i,n N ill, I')'n; Thompwn "1 aL :>}')02),

.l.o~ <)].1 dUling Hw dghk,mih and nin;'t1'('nth

n'lltmi;:." By l'lO() [he '1""";[<" hnd bN'rl n:,du('ed til /0 lo

30 in(jj\'iduaL (llol.'l?(:! d ai. :i 'hn; )iol'lz<'l J 999), l)espi\l> ttl\:

rc>,dting l'tdu('tioll ill g()H0Hc <l1\,C,l'Si1y (Huelze! 1<J9'J), Jloftlwm df'phant w;\I, hi\v(' f<.'uwered 11l an i)$limated

2biJlJ{)

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flC:dJHi:-, ;):? ~~ FOp\~LHfCln tT(~n~h (UJ P~tcUjx h;~rhor sfrd ih"iPUt:.l1iolb n{f UK" Cai~f{)rnid ~'1)~~\,1 (t ~;HTd~g L'i. at 1002\ ~tH' C!.d~ 01 i\ia\k::i O~.;dbk bland!,. ~rHtdl('ilt,h;'fn Absk;1 (S~:k~~ amI l(dcll,Lm} {;,!mdl i'l iii. tiltH), ;,nd ll1giflal-, hl:md 0'[1.(1,.01 I'}\)m.

X, i dml1!~,hnu! thi2 "petie:,' ldn);{' lCoup~r ,md 'i{'{'\\';jfl 19t1:l; SttWMI d <Ii. )\i~Jol), North!!l n ch)ph~m\ $,~,lh In Cal> [(mil" W('It' I;,Aimah:d :'It lOi ,000 iadivkluilh in ZOO1 (ell'·

,.,;\til d ,,1. .?OO2.L ,\hllll, ,,:; .. 1> "n" ti1<' "nly lloplnljf"tihlropic\!

f"0p'Hlh1.~niJ:+-' t)t' tht:.:o:It~. s:.p(;cks 1"BAy f'H'\T'r b;1V\> beeB nUHlcrOHS

h'C,iW," {)j lllC' g~nni;lly io\\, pH)dunlvity of U')jlicoll (!Ct:.:fl5,

IkC!lm: ni their ,wIlH pl)pulinkm 51::(', mOllk \('ah ;lB"' vnlm"CllJll'

to dic-'oHs f11)-IH dj:}(:us('~ ·~~}br{;e(Hn.g f,nd jo\'v R,,10s.:!tlC

\'in!ilbUHy, ,md hnman disiurb,me(', 01 11K tb't<'i: .('('('nt 'l't:dl'S,

!h,c f!;;Willi,m and h1.cdikmmeii!l ul(mL ;i(',.!s il)'(' ,:nd:ll1bcncd,

"!lei ilw C;IdiJiH'im monk ~l'a! J'; cumidtled ('xtfnd ij(('IlW}il

f977). AIU1nHg1'J o\.!{·roH ~'HH1'l~Wr;, ~H"t< (t~t:n:-ndy )l~!hk, -t:(")HW

hny;tii;'~u n)oJlk $t'~d {:I.)}n~1iP\ ,an" i}I~Ti).'"lj \ifl~ '~vl):ai': ()th0:f~~ ~th.'" h~

lk, lipt:, R0;<SOIl, 1')1 \1\1.' d,'dim"; mclmil' ItOIlE1tl dhtmb:Hlti",

h'lbUal bs\, dht'il~c', c()nrp('lition wHh {"hn";'>', ,harK pl'nlilti()n" intn'1=-,p(-i;l11C aggn~Y~hn1) and cnt.l~~g·t~·nlr'nt \vit"h h..,het it::-x ge~if­

M~'{Ul'Tr;B,,~an monk ,NIh ;m' pn,,<;nHy 0.'iHmatNi ilt :lSi) tn

~(tO jnd!vldunh. 'Ilk }1H't} .. ·~{ knuwn a}~~g~l"f!,~~1Jnn t~f tbis :;O;Pi:c[t.·'S)

w)ll<;h 'X,Uf, at 111" !,,'lIill:;uhl cd O'p;.' Blanc in aw \V('S!NIl

:;ah"fll, "utf(;rcd a n)iI~' mOri;,!lity '.',\'';'1\\ ,;,j HllNJ()Wn ()l!gin ill

1;)('7 \I.h'\ 1\;'I,i>.I(('d I;' nHmiH,r,; rmm ,HI' U, 10'1 11ldividu:l!s (j(m:l.ld'l ,,', )1. 1<>')'1), .\ j'litiWf!i> ~m!;mwm chlollY \\';.l' di"()ve,,,,! jJl lilt' Ciiit:i;m n"y nft TlirK,':' ((;1.!\:U (:\ iii. :!!l(!'ii

Uk.: Ihe Iwt1!'dn $(:at I'll" ()llCl'''l1hulld,wt ,{mU'WI"Il

(-kphanl "'ill 1.\1\5 IWi''''!!Y !~)(plodt"d dmi!lg Ill,: dghi0i:Tdil

;;Inti nin('+",~~:~rl\:h \:ultarlCi<, \¥\?h~ Si.:V{<ii"')Y f\~dlj(·i,ld

at ;;illlldj{)f brcw!ing sites, Although coHtrnlled h,HV{;"\> \Wii'

pnpU'ialiol! is r."lit1),\lNl :11 64{i,OOO lmlivkluJi\, \\'i\1\ maj.(J1'

!Jrm~.dlHg ,:dnnics IO(i!kd (ttl 10\1111 (;,'orgb (ll:tJ10{l pup"

p(~r ~~I{'<~nf th0 K0Jgu{:i\."tl Ar('hlplC'ht~~<J !4~1J)GO pHP~ pur Y'·}~H")t

M,\('qnarh' J;;bnd (! 9,01)0 jWp' pC, I. k;.mJ 'hlaml n 7,OO{}

pups [ll'r ),(,arJ, am! J'<:nimu111 Vilk\(,s 11,1,500 pups pl,r yt',ar)

l'~i[<1dm;kal!y, d("pjlt' lilldl fh.OVUy f<,HowJng comrnHd;il

s>c,;difi,'i illld ;1 J,1tK n! ,W'lS1:!tjl1t'ni bunting p;es,U{(', ,oulh<'1'!1 <:h)ph"n\ ,<'I:\! in \lw l'ndlk ;md lthIi,;tJ Hi ,:,W, dn:liw2d (';0% to HO':r,) i!dwC('!I tile l';lSO~ mid t~90, r.Mc:'l;;lJOil "t ;\1. 2!1(13), 'n", <';!lI'("d}llhcw~

dr.'U'l~ii~d.~~ alt.' poOrly "llnd<n-AD1.XL SOH~<: aHt'hH1'~! h<{v-<., ~lnrH'j. ..

qt:-.,·d Hwrt~ h) iOUt> .. ·tf'nu l.'nVifOnnH"'jHat dllHlg~.' h'ad~ng :(H

l't'';Otm:i' Ihniia1i<.i1l (Hud.nfl d "I, 1997; ~i(:~d;Jhon d ,It :clln:n, VVht."f<~:1~ Oi hvr~; ha\'(~ prnpu~·ed Hil.} {, inJ\i~'k~ \'1(: }n~,:n:·a~I,,·{1 kllh:'r

wh;lI(' (B:I!T;H and \1oHgln lilfk: (;11ln('\ d al. ) '192;

Jlrimdl and \VWhms, ('h;rpk'f .:20 oj lbh V,)!mlH'),

Ill,' f/.;nlidll!t1fk "m!ih'm h('Hlhf1h~'r(' pfwcirh (vW'ddr'H, Ho~~) ~~ad)('ilt(:r. ;"H:~d k-(1fXri'd o,;l·.\;j~9 <HV n:"~H'k'l~:d tu lh(~ Ant;H~.'tk

cuntinent ;WI' its "mmwHJlng p;w]; k,', :'ltlHmgli ill"O'",

'1'('d", hf,Ve' 1>(,1,':1'1 hniV(',tNi irr('ku1ar!y 111 p:!St )'e;jfs, tilt)' weH'm'Vl'f >0 <cxt.:mivdy :1, pHllIjrx'd~ dWWI1(,H: ill

Ill<' world, Climcot h.H'it',( h,e,,!>; ;m' w"Hl.;kd ,md{:!. the

C<m'{('lltl,),1 ot! th.:' t,;OlUI.'fVaUOIl of /llllilKlk :iva):; (CCAS),

rOf v.:ujUU$ ~o~ist k ~tt (~~\~5!)n:-;,. 0SUnt;~1'(-'~ of ahDn\..h1lk~\ f(u

1Iw Ant:nr!k hOlIS'.' 1l,'0!l dlfihuli to <lbt.llll. (:mHoni

",hlll;!th ;,ri' 22n,Olll] !H +m,(,(lfj for kop:m! w;d,; SOu,om)

~o ~ ~nHHur1 {~)l \,o,i('ddf,fj $(:~~H;; lO H) l~; !ldHk>o f(~r ('rtJhf;;~~ler

",-,!\;,; 011»,1 iOO,tIO(l 1" bSn,f)OO ("I' I,,,,·, ~";jk (;!\,(:ll ILl'

.;·dy';.(<;n((~ of ht~t{Hk;~~ i.;Snnl~~t("S lind tbf' h~)t(, ~'1~H·(~:rtaintl:(:~

~t~$nt"i(th~d y.:~th nH)d(.~·r!~·Hday i.::':>~iln;J't{~-i.l it ~~ 1101'

m ;m:t'!\,j[n \'\!fl'i,'nl population In.'mh lor llny oj

[h{~ Anlar,',\(' l,hod()S, NOlH:lhdt':"', t-ill' rdallvdy ,mall ntilnbtH ;:sf anirH,{j<; harvested (z;a. 39 J)on shtc(' i H92J o::;'V'(.'f a

\f~dt~ g('()gl~~ph~cal }';Hlgt1 unHkc·ly to bdV(' lld't){'r~('ly

;IH\~{'1C'd por~llLHj.o.n'{ of a~\y of dH';,;e- s"pccic'~; (hu$, (h"Hlgf;':-'

In (l)s1.rlbmioll ;ind ,1!jl.!!l(bm:e ;Jh; iil(ely du(, jo .. U]('! (;'1(.,

Ion. 'J\:\L1 d ,11. (1991) dO((llHCO\t:d cydit' p;,tkm~ in Hw

.'lg(.'sln;ClW(: ;111<1 <'<Ihmt ",vognl of (';tlwakr, W,,:{hJell, ;illll

j{'''p;m.J ~l';ils lI;Ja((:d to th(' S()nthnn ():;dll:ltkm Jnd<:J,

\son, J:mi'lH:'rnhlt(;, Hcngl'<)£l ,bel Laws {I 'JKS) ami HaEiHlcc (.[ ill. (Ch"pln 17 ili thj, \'<:>hnn{'j sHggested thai" repmlpd

(iN.'lhw in ttH; ag(' uf I'm;, H'pmdl1<:linn of ('fiibc'aJ(:! sr:;lh Wi"

causN! by iCI1 inu(;llo,,:(! <w,\1\\i\)Hity ,,( krill, whlfj; nSl\'!'):'.i·

hi}' resu]kd frum tll1' dlcpll'Hun ollJrd,':eI) ""bilk!;, '1'1", "W' oj'

i'irH rq:trodul'lhm inn("l,,,,1 :lgilin bio'lWf:t'H 1963 through

1·9/6.j ~~

:1v1iiblhillty.

UK" tt:lll[)!,r,d,'~Ii\ti(\lCk pll()c!ds, moH (It'.ldkl .'pI'd,,, "',,'f(­

IW<u!y ()vcrIiMv{'stlng iw (he end of til<' Him',

t.N'nlh terltUI}', 'Nilh the "c·',at1())1 d iHlrvcn, rno,;! ,hod;,

r<.:wvereJ td viHying d"~;fd')' 0\'('[':111, fur ,ea[ populiuinm

,wp~'lr to h!!vi: rcmv,'lW! fliOIl: HlpitHy \I!;m \"a lion popula.

tions. hu $.<.;,d, «bo outmlmh,T '\(';1 H{m, inl mil,;! oj m<lg­

ni!u<ie (nC;lf!Y g mH!\(.n hu: :;(:41;; wprldWli.\(:, in WUn'>ttn

illS1 OVN 6{lO,(lOI) ~i'il hon~), V;lda\IoJ} ill did,

:md ft'giona! pf<,;,hH.'ilvi.ty may ",;plaiJ) 1.11" Jiff('H'l1tY, '))

ah~lt'ldnn((~ and fi~C1lVeqi' ratf.l~ b<~~tv<'<'~l) lb(" h\:'() Bn .. HJPS~

!\niarctli. 1m Sfal~ weN: l'hotight jo ilt' (,X!illC( Iwiil a lC'HJ'

wun populatilil1 of 1 ,UOO j(l ],IUO ii1)lllWb 'Ni,!; dls(Ov,"Il:<i

;)JwHwr :diU ,.;linn!:, Yien.' dh{,'Wdf'd (,ff Hird 1.;[;,nd in dw

ti)30'; (Laws 19!:lL Sin",: \lUi UHH' lhe ;\nlarCik 1m ~,::al

IlClPuhl\\\m lIas l\rcWJr\ ",' '.1110\.\1 1('%),1 1:\) H 199\1 (,',limah"

of 1.6 mH1h'll :Juim;;J!, 199.'\) i\UJlOugh AOlardk fur

s!2'al~ hxt.,(C' ~nCf(\s)cd HH<~t1ghout UH~tr r~H1g1;.\ \;l dL

(19Wn that Uk' ,!UWN H'('OY(:,fY r,ln~ on dl(' South

Sh<:thmd hlnmh h a rrsult or kop,lId ~1C,11

l»(.frtli(;rn flU St'db illtmhh lhf" North I'neilk O';;';ilfi imd

til!' Tkling SCil, wlth prhnar>,' l!r;!;'dlllj!. i'''(l'l\('rie; on til,' !'xihiloj

iiHd CA"mm;lIHi<'r i5hnd~ ,nul smil!lt::r {'olol1j(·'~ in trW ~('a oi

Okhulsh. ;wd Kmih hJ:m,h, ,\1' (l\l1lying u.,[nny ~llsi) on~m:;'

tm SanMJ.\;ucl hLur£! '>ff th(~ wi"l of ~'>ilthcm CZtiJlomiii,

l"oliowlllg til" ,wahMV('~l ,jill! dejll~UmJ of N(lnhl~m lw

;iICnh dm.ing till' righttCnUl (md ninc\(.";(Jllh cen!urk~,

were' P 1'\.1 (,; (:\\;([ ill 191 l Imil H'CI)",'twl during ttw

litst half tit dw \ 'SGl!!dh ,:.Uti.u\,y, \i{)WCh'l, in t:'\l1\ta,t wilh

Alllil),Clk I'm \\,;Ik llr)lJi'lCl\l itfi :,(1;11 nUmb(cf, \0

th;'dlIlC lq;'cilll it) ttli: j 95\k ii lrf'od thill. h"" cLllltimwt\ W 1 ill.'

present dilV (,!\njJ:lhs ami Lfll!;'W ~:OO:l,; LtH,'[lll d. 11!. ZOO;?), Ti!H (',;HP;'-i; ·of i'his i)ng~)ini:; d<::\Jin€ 1,) unknov .. TL

Tlw ,.{'biNi Cllw (\f1~11I\l,tfaliaH I'm ,eah rWf,' both

grcilHy \kpldt:d by comnwt(f,'\l ',,,,\ling, H,)wt'ver, tlwil' pop'

tllatiorh hilYl' ~in(,; t,Ii<<::1l rntlwr diff,:rl'IH 1.,ljr'Ciock,.

1m 11,;;1 popll!;Hinm, 1':l';He ,mbjtcld (0 contnYlk(! hmHs,

ilav(:; incr(';lsHI tf) an ("tliIFIt<:d 1,7 milli()n indirkhwh and

I.H'<: \lHJwilljl. at ;\% \,n H'i\! (\liltkl wOfll\ 1;;\ ;.\\' 1995), In C{l1\­

(tiesi, AU~,lld§;lIl hl1 ~eal F0j>ulatloH1, whkh bw,,: not been

hnnt(!d )h~u~\ ]~)2.1, h~~vc ha~.f',,\J~{:'d onl,y to an f"Hhnaic-d ("lOOt'! indi'c'i(l,,';1h, Vidi \)1.:1\:11',' lil0. (:;,til'i:"':' l,i

17'.\,000 10 2ZS,UOO Imlividnak Th('w dHk:I'JiI

gf~nvth fidC~, ilHLI \_d)ntHl~ln{,'(t!} ~laVt; ~X'Ol ,~ttdbutn:l to ViHld:

bon iu rood and rjjHt,t'-ing, LJr~1!!,ln4( '\tr;~t~;gl(~

'Ai nOIJk) and 'WHr!'it'iH' ;~O()?.),

i'xtindl()!l during: (he ClHWllCni;)! w,lling ('I'a but have 6im­

Hill'ly nlhoumlnLI \k ",\\) .. ,",nt;HUk 1m ';i:ai, whwh br,;,,(\,

iil~lll()rt!i (if Ihc Ant,lrrtk polar from 1.>1'1 >ub··i\ntmnk ,m;l

hj,;lrl(h, i1tlmb,:(\ tlHln' (him Z:~:;,Df){) t.)

2KS,OOO imlb;id1)~I" and I, hlCf\:il~ciHg by ,b nll.lrh ,I;; 9% to

\-1% 'il' j in a f,'.\\' ,'ol;mi,."> (IV]((;('n, ,mel 'I'm\; 1997'). Ttl<'

South Amclj,'<H1 fur .,,~,l!, whic'l1 (\(CW'~ ;l!O!lg Up;; l'aGifl.;: lCM,j

of :"omh All\('rio, ill tht bland, w<";! of Ikn';' dd FW,;go, .ill

th{: FillkL:H~d i::;L~l1hh, ~tnd ~H Un.tgn~~:,": ({rid ~tlOt~g th{~ t;\.nHh~

,;m <.:0<1,1 oi llfiv.il, v,.a~ hlll1li'd timing 1h(; ,;:iding 'iT;l :!lKl, 1l10t(' r(;(cntly, in it.;gU!,ll SIlMn, nmlrnllrd }wrYl;,(, in

ullHi the 19\1(1), W!lik~ 1I11ll\b!'r« (If 50\\tll AnI.ell

t:rig,l flU .. ,t~al4.; ~n'F H'H:~y t1.-'fnalu d~~HiHhl1(OdJ ~qqmf'"

CHI.ly dtK' tu tit!' rl'Ci'llt c,)mm('rd,d h;u \",',1. Thc N,:w

Z('atmd 1m !;""t :;n ,,,:tlmati,lt\ 13sJ>on ;mimills, h

,mu inn"cw;lng t)lHXlglwUi Hs r;mg('in Nt'w/:'i'aJ;md and

j'tUri!.fiilid (Widwm ilnd Ym);; 1 ')'17). Till' (;Hi!d,:jh\j,l~ 1m $\'al.

now 1\),' r;m~~l Ft nl! im :)Ci.ll 'pcrln (iihovt 7,000 imJiyiJu· ;~h;)! y./('t) pr\;~Hnh;d (~xtlncl unlit ~~ s~H{d~ g~\('tUp

\Va', discOVHNi ill h\,' ',k <';ni.l\.!dhijl(! in 192f{ rf<.lwnsrljd

! ')1.8). Tl115 ,midl cnlony W<1'~ rb<:n (xh:.rmil1,j{,·'d hy

rn\t~C"lH (011\:, hN~ (HIIl'tboiorm:w 1950,: ! hibbs 1956). The

FIJpn!itt!on bnith:l{i,:(,:k p.'::-rUH),N! ~n ~~ S;Uh;<;t'i'HHhd ios~~ oi

gUiHit' vmiill>i1Hy (Vv\![j;'.1 H aL ZOP'l), Tlw (;,ll:,p,tgn, fm

<~C'i)t, \'d~ich h Hn!~h,d to the G{7:l5pi.{;:;~,:i~ hlnnth ~{nd w;:~~~

d('-pk't<:d dudng lfu/ ")p;-dil!g t~~a, H~{'C~\.!I"'t(·d if'> H:; t":U

Hlatcd pnpul.fHio~,~ h~v~:d cd ~lih}~H ijnJ;(H~ .:~nlrn~lb

by 19/1 HI/S. !ki\'1(~\,l'f, \i?t' in IUs "l','<:h",

(JUdl.!dW" l;olbJd.:rnhly in f('i\lililse 10 n ;0ii!{) <:\'"n1;

(Trillmldl d ,1\, (991), For 2X;!flipk, 'ill',;, pup rl1.0ILllit),

{;slimMed 6,000 h,OIJ{) ilnim<1ts fO!iowJng thc' )997 i99g

H l'\i!\o ('VHI.! {$,,];,;c;u 200Zl. I he' ,hun Fcn\(m<,!v nIl' ""II,

which h qlililrll'{! (p lh" hLmds nff (11(' r(n~! of Chili.', nm·

I,dw:d;m bl'imilL-d r"HI milii(j.l/ aninwh bd:o!t.: the s,~alinj(

ePl, Tlii!; \jl~·:(je" v) 1)1> {'s!me! !mtt! a 'inial! popu·

apprDxiU1;:11'fily 19.0nO anhUdbi. ·and gn~\\'~ng (\;Vrd<{"H~; :~nd

)'Dr}: r9~r?).

In (OPtl,'~<';~ ii,i lh~' fur \hds~ n1().l;1' .\;{~;+ Hon popob,H(~ih all}~.n

d+:,cHnf'} itnd ont' sp\~u\!~ (t{H} Jap(tue:"i~~ $0.3. !ionl h

!':"tim:L (,diipd/!,O', ~(\l lion; lM'vl: cl)JIrlnul'tJ \'(l dt'l'ih\i;' from

anHuopu:l",Hk j!!llmc!~, I'.l Ni:'w l:V~'H(S: Mld {(i'il:llSi:' mlllJH'aks,

with ,t mOSl n:<:crll CULm! in 2002 oJ bctwz'u) 14,000 ;!!1d

16,000 ;minl;!js (Saiil1,iu 20(2), Ill" N{'w ;!'<~;J!and [.or r looker's)

,,:il HOi!, wblch nnc!' orrtHn:,(j ill! ,lfotmd New /:(;)11;111(\, W~,

:(k·plt.,.tr'1:j l.l}' ~u['{I~t~h~~H:l' ~uhJ CU1Hnh~!l'i:~1. ~HJ;')i:i; tg. !fl,e5e~Hry~ d~ .JH~ (.:d[pJ!,' f1Uti~~:' lS rt.'\hJ\:·h'{~ io ;:1 hJ'!''l suh··l~n~oHC~k bkHld.')

{C~~k\ ~Hhj ftt!t.dH:I' ~999} . .I\U:-.na!Lln Si};~ Hon~ .. ibo %~t:'['('

i.k-pk .. t,-ed hj' ~~(;ali~~~~> t.:vcn the' poput~!,(ni\) ·of

;:ymp:,tfiC fur M':l!, \'10[(' ,he p,lm;,ry tMRI2lL lilb ,tk'Ck" nnw Olli: "I,ll(" world's rarc", ;;nd nlP';! Ilm!,ln! 'in; liom, Oll

~Hcnplll ot l.t~. I ;,i:-IH~lO~h hfi."i.!l..Hng ~'ydl' tH~gyjns 19Y3)1 ill

((.HltHl~! ~() lht: '1 ;t<H(HHb <.:ydt: iYP!Ld~ -of ~b'h grnup. h'lpu!a<­

[10m UH'(HtJ 0kw L.:"L!ml ;md AHsu:,dllm \(;alilm:;. wllik lili

:md IO,(!I1U {(;;,k, d ;It 1994: F: D. Sh;lHgh!lNSY t,t a1. 2{)IY;!

l'e\pedh-'dYI arc pn\\{lH'Hy ~~ablc 14nv..'{;ver .. unHkp tlu." New Zb1h~Jld ,:,t'-ii j~!Hl! Hw An\~~.dliall \1..',] lion h \'dd.;,-"ly di~11 ihuh~d

a~nnng 6/ "C;tJU:'-h'~:d {(df),nir-..;. in ~o\tfh~'!n >(lJH1 t,Vf'':;h>rn i\H\~

tHilli', ,md mil) in f,j,'t bc: in decline. The \tHJth lqll('Ji('~~n iCi1 Hon, 'whkh wngcf fn,.;rn BrazH tn

l\'w; w:,> d"l'litM!;'d in tlw hum l\)! niL P()[lul~Hj()m

"wnd al appm"imJ!i'iy ;W';-;; of thdr hbtorkill tHUll''''"

tt:Hppn~~I{) 2001:) .. \~n!ough Po-p!l~inior;~ :dO}~g th~:' }\It,'i~\~'

CO!\:\\: of South /\rncriGl aH,l pnorly knnwn~ nl~Hly aniH)ah

,:PPilfcntlyab:mdon till> [(:glOB <lm!11)\ "w,'re {\1 NUj() ;:;V{:nts,

Thumpson d <it. \LUUS) 1<op01t.;:\1 lJOj)ulHtion (]cdll!i,'~ in tll\!

F;dkbnd lsbmh 10 "L5% of t!li:1 en? 'l1;lIltl,bn('l.' ,·,tim:'\!.' h(:lwf'i'll 196:' ;md 1990, for n':1i01b tlmtwmain nm:INl',

P~lll prodw,;HoJ1 jg th~: FaOdan-(.h ~hjn.l 1990 ha~) tncnj~h·vd at

:1.2% to 1';'5% per year, thus jmHing this POlmlillinl1 on it

simibr gn>'1ilh tl',*:c!()rr to th,: lldlaccnt ArgoHi!!lCllll

j:lo['l,bii\.H l,

TIl\' lwdlH.'ln or Stdler Wi' liull (SSt), which un!;", from (:ilHhHnj;~ t(J J;1PHtt~ ... "\1S huntc.d f.or 'i,-';1r!nus r{~as(;:n~ ~lnH~ thp

enrly 1 \l:;()'o.l'lWIl' h \tW it )milH 'Hh~ibh:m:<' mh lly AI1l'i1·.il

~~;11i"{tt'~. Two !1<k.ct;~. af'f; i:'ufrt'ntly I(~~,,'~gnti.\;{~ il'j u.~, \\'i'ih~~l~,

with (;;'1)(' St.l(kllnll Abslq \ l-l,'nv! 'the t1t'nlJl'OUml point

b\'\w\~en \h~ e~,st,("In :;,tTH..l v.;"~'\tt'rn ':1t\1CKS, th>~pHx' ~hr.l (es~<Hlr~n

of lno\Iln~.~!l~d Pf~>f{'"\'\kH1 [;ron') ·oth~~l" dhturh;tnn;~; tlK' \\:e'Sl­

{Tn q'nd; !lVgilll. 1l !1Tvdpitom de.dim' ill 111(;' hlp 197(h 01'

*"H'ly l.'JSUs "LOllghiln d ;)1, ]992: Nl(C :'.OO!lj, 'i'iw W<.;"I(;;,11

\\Ocl': is nlfr\.-:nt1y lb.h~d d],'; End~ln~~i'ted tHtd th(", (~<'hit'tn sfnr·k

,1STiln.'iltt:·ned under thz' U5. EmLml\Ufi! Specie, .\ct, The

\(!J1.'1·"tl M ~lhP(:r1:(),;1 ('en",;:; of SSl. lll()t\i,lIlylndutl(: iml<!CH­

tnf Jn~~l)~ ~n h'jH"rij:~. g(~(n( l..~ut;~nJJknH~n~ ~n m~tfln{' (h'bris .•

:'"~h{.jt~~in(~, i'Huq,wtitinn 'l,vHh lh~w·d(-'~ for ItH.>tid ~l{'l'<Ul \:Jhnah-l

(han}:('. am! predn\l.o1l by kWn Whilk" SSL imtllhei! ""ed­dwd ('tn\y:--;"t~/n~ huvc hi.~dl thf9 objC'ch. gf hHf'U'i:i~/{" ri:.'''V4!1t h)

VI:! 1\1<')'( h ';dllnowld('ly anept<'d C~pbl1:ltkjll lor k;J Hons'

{.\·~"ni tkriim: (Nile .zOO:.h 1'1 H' ~ ~;~Hfon~i;~ :-'~.~~.I tinn~ \vh~dl f~H I}~t.''' frl)Jn ?·A~:\in7 k~ Brhhh

Cn~n(nl1ia, h dw ;rH)~l ;1hufHluH (~1 ~11! \(':1 Ht.il ~ ~,p:;:~ i\~<,_ i :tjlH:'Q~.

iargely b'L'C~lH~k: 4~1 pe(cL:t~h"{'d {J;,MU::.~g~: to ~,:A~lUt~kr(ia! (~lU.j\(~

;d~d. ~:(Hnpf:HH(ln for ~.~dfffonid .l:{')Oi.!k.,":\o H~'VLrHt Hnd

Ih.C{;·rh j9B2)~ n .... d~;c(:.'d tb(' ~thundanG' uf CaUf~Yint~;. $t:f.t lh)f),s ~n

"'Atl.!wm CaW'owlil ;Illd !Vkliic{) W 'tPFw~[mn\dy ! )SOO hldi·

vidH,~h hy thl' ~920~ .. ljH· ~lp~'.d,,·~ ~\~"I~' H\rH~~_\ed ':itt;~;'jt,H~y ,1\ S~;'<t

to 6.;l<'~<") y~ t f'hr~Hi~h lh\~ liHVt ph.n c.{ nJf: h\t~~(H\(·th ~>,'~ntHry {i'\Ml,::\ !''l9n l'r",,,n1Iy, llKle al'~ ;Ul ",limClh,J 2CH,O()O to 2Ji!,{lOO C'~ll!fm!\i,l "I'd hon, in C.S. Wilt,'" lC;lW'(!a <)\ ,11,

~?002j~ .:L uJ .. 0 I 1n.kh~inrwi 44JH)O to SjJllJU ~rnin~~~h sf1 \h.';'{ic(.)

C!<"'i-t.l'".i\1l{-"~·{ ~\tH~lJ\ ~d ~a! l.d ?.rv,-lLI·L;~ H uait'/ J 994~ .

Potential Causes of Population Declines

in fh~' prn'(ding \\'O~c'fh of 1hh J,j),:lph'f W(, h~rt'>c prH\'~di:'d ~.,

hrO;ld O\,i:f\'i~~;.v ?)i th~: popiAb"don stntus nod trdHh ~}f pin.,

:ntPb:b~ w!'11ki\'vide. ~ithf \Ilvck~; V~'f're l(~(tUft:d \Ub~t(lnt1~i!t:'f'

,Iwing t]l" e!'~ of c()nlnH~r,,!al ',>;liing. Tlk oxpbniHlull 1m­

Ib~~,~ "nrly .:ledin,'!. h (hn nnd tTd{lill''' lmmal1S k!lkd them. I !<)wcwr. populallOll$ haw not illl f""piHldi:d tf! .h(: <:~s'.'di"n

of cOimllnd;ll ')(,Flling jn tll" $!tm" \\,;ly. Some 'I)('<'in or iOfill

pupuh"lHtHh. 111{'H'ast'(i t (~Hf'H ntpi.1,tly. J h;<}'t!; ag~illl t~h:r{~ b HHh'.'

nl~r:\t('ry ,~"\: (r) vvh~'" n"dlJ<~''(l n'lPf~-\ili{y! tHg{dh\~f 'i,-'dth rtt~u\~ro,.'

m!l)lut~, n(:~lll'd liy !lH: "arEu lh)j)lliillion l'<:<llACiiml, PH'\)­

ilLI)' fuduJ pll))lJi:llioll gmwt!lln " m,Hlllel exp,x\\-d imlll ~,jn.l ph~ il nd flIt·or;.:, t"h·,:l~A'~:V\« cd }"i(>(

,peck> and populiltion<; dtiH."r did 1101. u:covu from ,('.:dlng {n dId :~(O\,{'f bUt hH\'~ ~ul.r$\:ql..WHtly ~tl{~liu {kdincd, Tht,%,

bH~:r Gl~(~\ (t~(' }nO{t~ di1'fKH!1 to :Ul'Ukr"li(LHd. ~!t ~h~.'· nH;d

>'"diem 01 this dliipj(~r W{' <l!.tcmpt f." o,1'l\"d 1Igh; '~'Jl Hlv"~

pspk\inlJ ~;'Clld$ by flllipping knowll Of :\mp(~c!(:d lM\l<2Wi.

<)f V;'IL,tk>ll in lil<o, hhtmy, bclmvluf, ,md 12!!V1Wllmcn! with r;,pnrtN! r(jpnf~ti(1n Ir;ljN·tnri('so Om synthC51s f(jrl\~e, on duel' Imttl:m:; :111<.1 proc"sscs; dlfkdng {(,pmd\I(~tl\l\' ,h'11h>gks bctwc(ol'l phoc;d<; ",Id O\'!Hi1(\S; physi(\f(lg,<;.Ii. Wni­

f.iltiD)]' ~~md'lt,~d with pMtkta111f !'0ri,ging ,mel diving behavior;

d!H! tbl' lCWWflg suscepllhillty 10 dhlmhal'lo.' In pH,y n.',otl!ce, ,Hili predation.

Uf" HMof,! <Inti Behilviof8! Correlates

It~\>dd~ and (~t;hitd~ hav(.> ~~f~I\'('d Uh~ C'{)-))HKl!niL~ th'UI<UH'h 1)f'

kn('!;\l'inl p:l1'hniH()J) and I1Hli!k In tiif!(,,'1ll w;'Y" {!bnh':}.bm.<!w 1<)70: ("'it;l i9n). ;'I.h:ol plwud<; <II<: i,'ilphii;

hr=E'{~d(~{~;i ,o;;~()rin~L prior to p~!rHlrit10n.' suffkicnt c·n('rgv for

T-rlc t'T*iiF-{~ bi'1'Hi ion pedo(t {}tarH-ds~ in COiHt~1:\1, tU(! HlCCHH<'

hTt1erlY!C" fr/f{.Hn~~ Hll,"ift.-> or k'~") c~)"1tti~~~~~}P${y dUf~n~t kR't,~~{t~)~t

fi'n>!il j 9illa,b. 1.99:1; [loyd :!O{tO), TI1,'£':, dli'kH",ll !.(.,;,10f;1,!,

<Xmf"l dlifHil1g b~llC'f\\, ;md (0£1', to plHKids and oiaJiid" Clplt<l! hn;('ding dt~il''i(Kiil1r:, ,,,prvcincU\,\' :)\i<;,'(S, Emili !('·'.'ill

!lwei ;lv;lilal>i1i!r Tlw mHri1kml1! jll'ov!511)ning of pup, by

jJhodd mOl!!':" h HUh largciy Um'fJ!l>1I:ilhwd liy !.[;lv<"1ill1:: 1~~!~(:' in: H'l'1d hi)nl n~(.' jni;~!!ing J~;lnHl~h, :dH'i-d~y ;.111ovdHg

thfH1 h";o lHiHl,l~ pr{~y thai :HV rnnre "\.H<;p(~)~;(\t p;;H, 'hy, ~*n;~p'

di.(~;~bk.: Of ljtstUHt troH) tilt' r~Jo.kny, 'H1\.' ;h~f{~~\Hy oi !6'd~

~ng dlfrfnx tl!.. {{tHon (Y~))jn{tFUi o~.ur~rd i,:nHks 10 t'\lUlg(

do\~.~r ~(J {hv 1h~h r~,'prJ)dH{;th~l..· ~u'ct:c";,$ <And

h)(~a~ ph:Y ~lhtH;~bn-C,' (CO~~~i'l 199]) nnd ~h~'r{'hy connF;_ ,tb~g pi ~pt1hit itlf) '~t;rt t1'). !J) Jn~:;~lb"<d i:nvin ~rnl1j/n~;,~1 changes -such LiS LI Nit)(~}~~)lfth(-H) {ht':1l1:Hk;.i) ('\'N'~~i,;

Ohthnkh (1 ill. J;'I91.; s,," ng,\H'.~ 27..1 to!' n\,~ C.diil)fni.1 '~;3

Jk~n}. SIJ~Dfba~'Htt ~~at'!at~(lD~~ HI hip dtn~lth.}n" ;.:~mdl,;'· lU}1{H.,.

th:;H, h~n~ndi{l,/f Fl~P8H)\Vn~ ntt~\' ~UH! ~Ul'Vh~{lt'in n;spf.Jn\{1 1':)"

IHlm[lims ill pn')" aV;jilahH'lty l'allwd by dunging IlCc;;WO­

grilpllk l'i.il'Idruml$ im:(mmH()1I ill ()\;uiid,; !C<l\\<l l'la!. 1989,

>to

x ~

'1:;3 <II ,S 'l)

.~ c

c ro ;:l <Ii 3D

::I 0 0 0 Q.t: ::I

u.,

21)

10 ~r-'~~~~~~I~~;k~~~~~~~.-~~-4~~r-~'-4'

1tl75 '!!lfH} 19811 Hl90

Year

hGlH,L:'i7A P\iP p,<)(.\tKlil'll of Cllilm nb ,(':' Hoth off!!w Cdtlmnla !:<1int {C(·liTt'!!i.

<:, "L 201m. f'otl\e!lw dliiv!<:II{ dl"'~i of Ut(' !W>:) ;)nd )995 "Iron;; !':) 1\:il\ufSonth,,1'O OS(mdicm {i'N,\(}j ('';{1'1I1'%.

H:s«1 (,tal. .hN!.; !ri)imit'/l H ill, l'Wf, lloyd ,-1 ,lL !()i).!;

;r\\r\ l"IlHfi\)' ?()(H). On ~11t.' {'lh!,,! hand, !i%'\\m;t m- )",.1\1".,(\

ieedjn~ dlHInR b<...lntlon hndh nH~ iotal ,'HHounl flf {~H~>.rg)/

{md ('J ()h"in tlJ"t Ulil b" im.'{:sidi in phoctd young, n;~!.ilt!ng In s ~tnaik,:r n:JaHue pHp lnJ~S ~u tHHriti('h~~d it!dependi.~HCe,

OV('( the prt:vhlib ~,,rt>ar; po~~~w·t{HUng ~lnV~\~:a~ h r('l;1h"l;,~ 1n

bo~h \\"(~':.lnJH?< JH~i~:\ (~,·{H.'r):~y f\~Sef'j/(~S elovhh'd hy rjH~

m()!hl'l i ;md l'Ostwt':ming r,,$OUl:'::\' ,1VG.ibbilily (:;1.,',,;111 ;md

Ln'ipw HK1) j'mH",;'Jli:)i(:, w\,';nlillg in plmdlh h llhmpt,

\'illl, pn:v"nting pups frum !('iludng IIOW l!wlf lt1{,Hla'; hOI,!

to fO!<Ig:"" ".aptli<'Jltial (\h:\cht;ll1\il"W in tinw5 oj lood shOrlilg(',

j~ oftr·n ~yl~rjHnnGH~ wHh.IH ~PC'd('S1 \,,~llt~+h nH:~.~~~~ hH.ge HtHUher\ -lJt ineXpCtk'l1(¢d lBd~viddab 'i\'~H two .s~frHdt;l~ ll\,.<ot1":,,11' tt':}ling HiI.' v/~Wl'r{.:, fh:!-ltf hf.;~~{ding (nh:w~k·~;'i lb.us

rrH~1jng HH:n\ tn pn-d;tffun. Thl.~ \ypit ali;- ~hnJ1

p"n<'d in \lh('!tkJ~, ,lbn :,;I!,W{'> Llwm to uH1w<, un""" Lile hl't",.:dhlg ~d~lhU~Ht:5" :;uC{) :1\ F4~{ k h.\~ (SUdH1F,; 19/", It)fP,;

COq;1 1 ')~n.j.

fmp!j( altOn, • .>1 the MOI\~m\:HtmrWli (iiHi'H'llCh in '('Pm, tlu~-HDl~ ~1nd ptlO~.:l{b anti Ohlril(15 Hf( pot.: Il·,

tl;lUy t~xt'mpllfwll bv \11(' linj!\llll; (\Hj,:r!8Tlce£ In I'H'O\'l'l) I,!i('~

01 I'm' :WIli;., ek:ph;mt s,>,ds, Mld ,,'<I Hum on C;Had:l!uTW hlami,

f"lt'Xiccl> Wl!ik lw :,,'ilh ,l1ld .wars iJ()1h incw~s"d fo!~ lowillg OJ(' o;,.,:,Hon of !WOtilll\. 11K' elephant ~eal r,'ciw'i,y

hM brc<l L,l mow dlilll1illk (HI(!.m· 2/,;'cL !)\1lill~ Illb ,,;mw

pvriud, St'" li\m mmlh~l':; at (;u<ldalnjli' hland !l,lVP n'l1l:LirH,'rl

.I,)w iY1111'i)),,1kdy st,1blt', Wh,;l{"ll;: pO[lul;)ti.(ms d~('\"!Kr,' h;lV<:

inn,';;SN! A:> (t!((mrt: iJrv<';/t'r" CilHornill ~ea How,

;\!hl GHiHbll1Fe iur sl'ab lHmt rUHuln wUIl 1hZ'll P'!!)' f()r

ill,!" ;lImo.lt ,myw)ww In lilt' Nonh l'crdik Oo.'aH iSfc'wmt ;\Hd

I)i'! .ong '~9~r{; J ,t:~Hn\IHf l."t ;i~L 2(1)QJ> thus j)nl\'~diug thit; spi:ri;.\;;

\"-l~th HU .. 'itter {H.;~;e\~-i tu f.>r~·<}; H·\nUrr(~,;,

Vmii\!iol! ill dl\'c h(:h;lvior <;Ill ;I!K< l·llt.!l.i.':IKc' \('\;I:<;tng ,'m­delll y 1111(\, '["", the pm;:'1111;<! 1m rill'), iiwltatinll ill j!1ii.!Uj;,·

tinn n:'.«.~gl{H"ion, 'lTwt h,+ pinnip\:'d~ dna {Jf)QW1l..? iii ~~.}r n~:~~.H'" 1 h{"ar phy:,]ojngh,-,J! hmih ~Ilouki ll;i\'<C litik.' ,',rp;;cilv to ;,;:Ii'."" 1'\:11,

,lging dforl m h'sprHlW In food ;w;lil"billly, \,,<lk[\~;l'> Hhh('

lilut "pn,!ti' hdow llidl' Il!lysio/{lg)\:i',l hmib ,hrmld noi i){' so "nn~lFlln0d. TiHh, one woul,1 cqll'C! slrorlW'? COWH;lHion

1,[:IW,'('11 POI'\Jl;l!!r1H In'od, and IOnlI nvldl;,hilliy [Dr ,pl'f'i," io

rile' riltll\H tbilll in lil(' hittt.r glOHp (Cmla did, 20m: Co:;t"

;md (;;lh', ?O[H; ( :');11;) ci .d, 20(H /' Thb propos,-,\! )d;J\i()n~hip

·Jwukl be pin\[(:\I!arly <.'vkknl ildw('('g b(,l1i\lk ;\IV\ W'.IH'I

~.ohHlln ~Oi(i:gi'J') bi,'ChHS~~\ iK'uHiir ka·;.lg(q'~~ nT':l~l t)0 "fvud~]ng ~11

~.I.~yd:; ,"k):;('1'l,;'~ th(ir :l}'~:~X"kn\B"n VhY.'jlulugi(.1; .. ifvJnf.~ r<~p~h.Hy,

H\).pthk' r:),H"t\:~{~HP, r<'.F.i;h.~;s It:nj;,~(~r hd:O);j.t €it-1t\:~i-~ thus n:duc~n~i !1"u' -On\(\ bt:n\.'~dth the· %Ot"oi\,.l'· th~tt :;-; Hv;,dbbft' {O ~<-':H(h fot PIC),." ({ JB{~t £1iH~ VViULHHS ~ 999). Het:~~u:-'<'I ;HluH':' oj h(:j~H:dc l~)f~

nglng ~p\.'{~.('s ~:rt: lV'orkh~g tilt 0]' .1\t?dH ~ht'"it ph\-~~oJogk;-d Hndl(

tlw ,malln' IIIWnilI'5, \viHi their ;,,~<hH,'l'(l pl'ly,;bkll1kal ("1'<>­biHli(.'<, 8nd {)xYiWrl ~to!'c" ~hi)Ujd fit' pmli.:uJdrly vulrlf,l'Jbll, t"

,""'JUra' iimitatil'm, Smvival uf luw'nil,<, ill [;,;111fJic loYaging

sped"" mIght thw; lw ;, m;*'J[ <id>trmimmtn! Ikmq.;mphk ttt'mb. Fmt\i{,nnuw, !wmhic fowgluR ~IW('ks might I>(~ P;H' tkllliE'ly wminVl' tel bottom traw!>:>", wl1kh disl uP! the llc,bi·

t;\t ;md l (:!YltWC tile' Jar;?", ~ize·da\",:, of jhh{', upon whk:h Oky

ofh'll dcpl,:ml iTlrrmh ~'t ;11, 1 99B), eill,l", nme! h;md, lil('

tWill ho, mil\, Ix ,I lllW,' jlwdicl<.hl., ~'A!U.'''' of PI",')' th;w the

w;lk!' {'Dlumn, i1H(.I (1m, \J('ll!hic IOJilKil!g ';Iwei,':i may be k"

14000

12001l

'HlMll

!::

:8 IlO(J(l

J)J ::;;; n BOllG 0

fj,

"11M

20M

(j

16tH, lSi)() 1920 '1940

N afep:hant }i8ai

I I I

J

I

I

GU:;1ti:a!upu ~ur s~ai I

I

I I

!

Your

FlGURf ?Y,iL .l'Dj>I1i.ntlnll t.rends from th~ due" ~p;cdl:' I"f p,iHnip,'d (lid' Me fouml on

GWI(!;dupc htnl'ld, l\kxk,o: (;al-itornja ~L'a Hc)]), Guati/l!upe fUr seal, (lnd N,gtiH!W deplml!1 ,e<J li.l'r(;, from J. P,,)hlo.( ;jli1ll, ImJl\JbH~!wd),

d!f.zct,:d by U('''<JIl-<''W'aph'( p';;J:tmhiHIOll-; '''Kfl iI" 1'1 Nih,)~ 1"11',,1\

IVdtr'f t'phlj\ljl feed!'l" n';Hlk! ilild 20tH). ()h,l:fVl:d n:lil\Ji)I!~,hjp$ 1),'11;'(('0 :ll'i'ilhj( dll!(' Hm"il (AIlL),

wi1id I j, a nl(c!hm" 01 phy~iolL1gkill divi;; Ulp!Jhilit ~!, iHld foraging

bell;wie, iJ(J(IS;, Ai"" nLlrlid ~p,'Ch', (j\m,.!'(jk' ;md !,ush;;Han 1m

sc'a!s; ;\ltSilalial1, C:llifoiilia, it!lt! New Ze:lIaml >1.:;1 li('!J>J pj() .. vidlJ~ ,m iniliiit I\;',{ of thc,c j.Hctli{!ion'i (Custa d ;11. l.OO-+) , i'!w

An\ilH:l[c ita s{'iJi milk .. , ,hmt, ·;hallow diVl."", while 11l~: "\15'

'l~\li;m ;md New Zc;dmll.l ';tJ HOlls and (hr AU:llliili,m \1)( :;>,,)1 milk,' {k>:!p j1'fllong.>d dlvc.,,, to HI(' lwnlb(l, (lloyd et ,d, 1991; Cosla ;md Gales ;WOO, 2X)0:1), C,alilornlil $\';] Ilons ill{' C,pi'('l,llly int"n:stinf: bC'C;1l.l'C they (HI' kw;! Ow fi.'nmJr', ,wd jt!\'lcnih·",j diH~

"pllwkgjc;:dly Qr! tlw cml"t ,.If ,{nl\l'l,Hl C,lliii"mli1 d

ilL 19i:\'h Wht,'I:{',1~ mnch {jpepef 011 nw.';oppjagk lJ["(,v ill tf1(';;<~a "fCmr.'?, (C 'E,l<uhn, fl, AU! [ojcs·(;mT,i'lo'l, Mid D, p,

em, (<I, illl.jJllb)i,twd), TIn- lW;H'\\If(;Wf"\"l:ding Antardil fur ,,,,'ill ilnd (:,,\ifIlWi;\ ,(,<Ilioll in ·;onthern O\lifu[ll!;! well wiihin their c;llwhH(\ Ant (cAUL), Wh~'i'"n) t-h.;' tllr,s: t",mhk;IHy into

"c(iog ~;p'}d{,:'ip{Jrm!alhm:, routinei)' excIO"d dwir \,;\DL {nglw~

27,6), Fpr butl! [HD\;llis "nd ,(,I Hom, 11-,(' k'EIllie

,"jO'X, of the-if time at sea diving, W[IlTC;Jl' illt; t'pipeliigif" fmagus

'1J<'ud<1O% of lhCtl lim," il! :-;"" <.living (hgm,' l"1,/), 'lJ"',,, fwd" ing. mgynxplilin why poplilili ion~ 01 many 1m s13:lllVecit"-,, (ami

lhe pAilgic Cidif",m!i, 51':,lioll) hnvi' in<Tf'ilt'i'tl '1N1'!\'r(~M

j}lQ;! ,l"il lin))., ((llilll}' of which occupy Ib.e sam13 tifl'iI ll~ W,'iH"

fur wal ,pc',,),:'>; CO:'(,l .'mll <';;lkt, ZO(d) lliiV\'

r.:-mCli!)e(\ M;lbh· I)r d<:('lirwd de aL 199,';; Sydi:'m;lll ;md

Al1,~n 1999; Gale~ ami F!et(iwl !!j1l9; (;,lit',,"! ill. 1 (04),

Hisii' o! Predation

l'iHn:j>~d~ alt IJri:Y-(';.l (,Il by il \',n-k-IV nf 'l/('(:;('S, Including ntlln pll1tli[lNls, ilnnmm, l.!\:<I!"y, \'\'\\kL'~, l()x':~, (oyd~'"

bye",,, awl jar!.;,]!" t'iigk'i, ,hilJ);s, ,mil Rilk,[ Whilks (.Vdkr

L',GOl), ill lh,: norrl"itHl Iwmhpiwn:, limd- or Kl"fmocd pruj ..

;i1Di'~ I)wopk, lw,i1", t\'iU,,;'} iWi' PiHliui!luJy llllJ,Ui\im1.,

whc IC;)S ] n til(' ,m ill Wf!l k,~ ","jl" ~tH; irel' frnm

\l',!Tt''i(fial hut ill"':: 'l1L'i~'llt:d V! ~"'h;'Wj ili,I\"a,i<

pn,(!iHOl), riw d(:Mty dll'l'lgCllt prr-liator ilvoi,iiHK\' til( lie,

bdwteH "'nic MId i\n[arclk piHllip';(.\;, with Artli( ~lkcih

Ik.;jng iq({t tlK willn io !'-'fape predation and Antarctic

,peck, :i!'Z'killg wfng{' Gil thoc lUi (Stirling 1975, l,983; ,\VdleJ

Z002), ,,(ksl: to "ihe ~lt"agtl:t Elml ifnpmtillK,; of 1Iw'w pf('i.hH"o(­

Ph'Y int1cr'ldinm.

Kilkr wh:t!cs {Old/IUS (11m) Me pwbably 1'h~ m,oBi impor­tant i!QI1Zllk pn>dator of marIJ\(' l1HHllfllllls. IJarlml' suds ,He

tbe mu\l commonly n:porkd j.lU-y or kl1kt Wi1ili("; in the H()rtj'Wf(1 hemi,phen: (klfn,oFl {'j (d, "li)'il). llowt'vt't, killer

Ivlldh,,, ,H'e "ho knowJl tel prcy Orl m,my uth"r ,pc'(iO (>1' !m:ludiniS \"he (j'ill)(-;i\'(,l" s('<\f C.,lnith ei aL 19Hn

"/HllIwm Wil lion din'! \()ullwrn t'lcpliMli 'fill Mid

FIN'>; (;ulnN dill. ?{)DfI), l1ilrtilcm fll!' :,wal 1M, t;ndltl jwr.){mai communkmiolH, and C;\litmnia >;IC/( lioll, III \OJ"ll('

;",a" killvr whilln lnklltl()flnlly !>him\! tlWIl1S(:!Vl.;' In m(kr to ,~,'i;;,' plll1li]wd prey, 'lKb iI,: ,()UHH:nI ~,:n EOIl) ,:mel 1i,,(lj h' em ei(::pitaot ;l'dh, .. Hl !I,l' lKad, (L6p(;7 Zlmll.('jI(:/: 19[;5).

Al!hough pfcdatm ,'(mtwl of pimlipt<.! h tlHfiu!l\ \n v('lif)" ,,"verat ,Iud"". t'iHwr (j('nH!ll.~t!";;ll" or

,a,l~g('it Sit:nificunt pnpulati()H ll;'v,d itn!Xl(:lS ()[

Hmb(.[ ,('i,l pnp prndlld\on ;It Siihk hL,nd declined [rom

60H if! ]9R4 10 ju\li2 in 1")')7 (tiKi)';' ,'II)\! Sloho 7.mm; EUW('ll "I ,:II. :!.O()3). :\n i~~Hnuj(ld 4,',% of l"lw tota!pup

lti'01.h,dl(ln h'''" kil1c>d by Bll,lrk" durin;;- 19\16, ThE jnGc'''~­

ing [!ll'y ,eal popnLllioll may hdV\' imiin'U'ly iliijuem:~d

harbor E~~:ih by ,1wrk.:. W Uk' .rq:iUH ami ,-('lI!,

pCl.lllg ,virb liar],,,r '(',ds ffir pH:Y (ilo1,Yl"n dill. 2n03!-

fiJi AU13(mll;!sP.IltioJ1 :1 " ,;}

'I' HooKors. SGa i/Qfl

!I! @ 0 AnlarcHc lUI seal ",J !iii <1/ Austrrd!an- fur se~1 0 III .:r. <) CA sea Hen n~eS{)pOh1g}C 'j ;:,(j , f!!i!t (,'J, CA se" li<m ,,!!lp(ll~ll!" 1) [l!i

~ iii 'i' <) ill iii Jill

;;:; ~ 'L5 ,0 '1r V" :::> 'W .JI> '(;l

>l~ "! >-

" 'I' "II'

'5 -',.' 'f/ <> .~ (;iI.DL

c: 1,() ifP <Ii> t~ {§) @

E {> ,,-Q

() <) (> '.> &/,6 '0

"L,-ftl r..:

.. , , r~

i) 2Q 4Q liD SU '100 nil' 14(} 100 180 ;lOll

mve depth (mettlrs}

fj(~t)HE ;t'/"i,L nin,~ twdorrnan({\ ddlnciJ .~..:. thi:- i1-tti01)f ;lVt~r~l:ge dh·'l~ i,.hHdUnn to t h{~

1.'nl"")"I(:<1 ;wro\)ic ctlv,' $,imll ;('\VL,t ;" d i'IllKl\on ilf {Hn' depth in fIV," pil11l1p~{! $jJl"d~""

Hang!! 01 ,,;\ill. ()\ltUm:d hI' til(' !lox g the: (I\UL ph*, SO':;, to ;tcnlH!llftH fhe III I'MR c",§itll,'k\. NoUn: Owt !.I'ltll lilt' diV(' thwltl.m ;l!1d r<::'lldcIKY t<,> \~){C!,:(d ill, .. ,:ADL ~r" J{f~;H.{'r. .in b:nthu; f=OT;tgf:ng ?:;prch'\.

70

M c::::l Epip0lagii: en

L !ZZll Btlnthic '> ~ 50 ... c::

f1>\ i!:l. 4(J ~ IfI re t» rJ!

4-' 30 ro a;

E ~'::J 20 1lJ >-:;::; ro

1i.) 10· t:rt:

FH,;URi: )J 7 l~h' 1f"hh\-<~' hn1\~ ... p;~.n~ ~'Dr~)g~ng \\'h~~r' ~11 S(~D, f(~rn'p.H;\,.·<"~ ~l\' rOh i,.':tgh' $f'~fCt~'.\ 01 ut,M~kb_ 'j he· grdop if) U~(' j(~a (~n~;;';b ('IJ h.u ~(>~'bh ~~nd t-h-e gUHtp to tili? t~gh~ {'(1osbts

,oj ~\"~l lion',_ ( H dM~ ~i.:h Pon.;, H'ii~ CaHforni:1 Ht';n \~pjpi.'!1Fgiudly. lvh~rt',d\

on~~s 0n!~>. hH :,\.).,d f()f~~B-(?) bvnthic:dlyo i i:ltt-l: f4:d Ar\UH(th hu <."c,kh i,'Olrh-'l froHt {_apt:

Shirr-..Jf fCS-} !1nd Hi~d hbl%,d iBH ~CO\\11 ~lWJ Gdle.1 20UTt

Spiln~('r ('j a!. 1200J) recl'utly ~f!~'~\llakd WM 1:111<;1 wh<ik jil<~datkm Wii~ ptindpally rC,p()H>ibk f,)t Wi(1{"PT('<1d Impn­

'lalhw d.\.Tlln<\' or 5(';, oHt'{, lind pinnipl'zh, in 'iDutll'lw;;t

r\t.a:\:l-;i.L T ht-jr i"H'gHHH:iH ~··('iL} b..:t5-(:d lrfrg('~y on f~:';.tsH)tLHr

rmaly,e, from \kmngr;,phk and "nergdk mod,ding (\Villi!Jm;;i'\ ill. 2DO·l" Stffmg <:viddKt that kUkI wllal\: p"'e d:,.t\Oll Qll",J !he wa ulkr d<ccfine (bl{~:; d aL 19!)i:\i, ;l11d

V,tHOU, lw:ull,i"\('liclc, in tiW avaJLli)li! !nionnatiOll with

[\r;lIlcl! ilnd \VliIi,.nb ((h'iPi(~! 7.0oi t I1h voiumr), on ,hi'

Lkl$i;. (,f ~imiliH t;VI{i,'l'jC(' imd amlly"':S, have cOllclmkd

tIKI! 1dH"r Wililk predatioll might <Iho luvc iig,w:,d pnA.l.ll,

Bently ill tfa' d"cfillt' of vul.IOU:; Soplh"l'l1 (h'i',m !linnip('d

popl)latiol1s< In support of thiq laW:! "ubgii'sUnn, kiHN whn10~ 'NC't(~ llhsN'vdl htking up to 25% or tlu: ~()Hthi'rn

0lcplwnt s('al we,tnJings frorn one lu'adl at Croi',d !slaml ,mel h,Wf' Iw(,n lmpHcatf'rllP til(;' dt.dim; vf dm~ PQPUllJtiOH

(Gnlnft 1992; G1\ind "t ,"l. 19')2;),

l'innlped5 ;;hc, [m~y on one 1H1()1IwL ,\ ~\ngk Hooker's ~<"I

Hnc on tvlanjmlfie hland kilkd 43')/, of tbe 130 ;\nti!1(lk Ilnd

£ulH\lltilfct!c Wt ~o:,1t pup:, tJOrn uv{'[ a ly,U ycar pt'lliJd (EoiJi<bfH1 Cl a]. 19'./\ll< SimH<1tly, kopanl ,ntis kilh,(j lin e~ti­

J1Hh'd :$,[,);, oj the Anjardk 1m ~,e,ll PUl' pwdmlinH ili S[\;I

[slam! Cfloveng d ilL 199)\), f1\ ;'ont;l San JwW(, Pen, up to

1),;:\";, of Sm.lth i\ml'l'imn fur s0al pHpS 1m' H'p,..>[("dly k.ilh:d by Sout!lC:Hl ~\~;l lions \l !;m:UUH 199;n N1.>llll",n 1m ,,';tl pup> 'm'

kilk,d :Hid \c;\h~n

Jnhnsol1 p;gn

Summary

In thi~ dl;lpt\;;t "'ie h;l''l" r\:vi(~\v{:<\ th", (um~!1t status and

tn:n{\s of pinnip,-d p()!1tdatiOIP; \Vori(h'lk!<2., with it tocns on

tll{' imihrt>l',,~eni'< and nalural hk,)ogic(ll PW(;(:iS<:, thnt wight hI' re,ponsiblt ior the:!,,' !i('rHls_ Al1tiOUgh little m nil

daUl an: ;lVililahh' f()l SO!)\(' :;p~:Cte" llSQfu! time s\:'l'ie~ (txht f\)f' ln~rny others, ]n{~h<::\ing pupulations lnt:lnde J))ost ui- the

'tmlht'Pllkm}\pl!NV foH ,{'flh, i h\~ Uiliwrnia sell llon, hM' [:10, ,fol\! pop,dMi()[')'i ()ff ttl" west ('ml~,! of UK Unih:d Stal(··;;.

and Ill,< northern l']('pil;;lIt !iNtI. Foplllfilinns In d(!CJiik

lm;\w:k lli)ulWIH ilnc:i ,01.101('1 n ~['ii liom, Ilw llnrrlwm fur

~\~,tl, till' smillwrn elephallt ot'JI ill parL; <J! the Soutllcul Ocean, lind Hil: harbor 5<:<11 ill ${)\11hw('s( :\;~Sl('L 'Hw hopi­

(,Jl monk $\'(\15 <l«} cJthl:r ~tahk at low levds or ill ut'drm:,

P<)!.lt1!<rtjOH i rc'lhh fm polar "!n~cjcs it! C~ pondy knnwn,

by .md Lug\.' UKS\: <>pc('l.;:, app;:;rr tn in' hot It almll­

dan! and iillrl)! sLlble, uu:on;!)' fi!lltll''''~ or ,,~(C!H p0j}nlatkm ck:clin", me most

(c)fmnoHly al\tibll~cd j[) iniNi)C!iolls wlt.h IishDri('~ and "IlV]

mmllentill cll;'mg~> l'i1!lIip<'d intt'r(\djuiI' vdtil lhl'l('(k, in.; IwK

hc~a.h {}~)~r<l!i(awl ,)nd p(oIogiGi1 (:'fft·trt~ il!~~rvpy li;n{7; \,latt:

,HId fLu VPi" 19f1/)_ r'A:",h'gk~l dkcb «:\ult fn:lrll dtr('(,

(j!l1F~Hlion, tllu;. (>S\cn:,jhly r(··(jlldng hmh pOh,nlial i'1:ilu:ry

and 1ht:envlwmm.'lItai ca!Tying 1:i1.p;iCity lw pinnipi'(h

df(.(l, oc~:m 'N11('.1'l

non~ UHl1C' into dired COn\dC1:< For

MId f,';}wry il1)(c;:l\"

plnnij!<'ds {,1ll be

irH.'JdcntaBy entangled and d~IU1i:lge fi~hhrg gear .Jnd .!C'rnov~ (If

d:mMg>:: ihh ill rwb Pi Clll !,'hlr~g lillO, A (\Hwml)'

c~pl'''ldltlg 01V! I.Hfgrly Ilnrqi,lllltNi tmde jl1 sral !lrod\l(:~s

(n{'i"Vf~S ?'{J(2)J Pl}Of or Jnj,')~J,jitktd .pofH.da1k~1~ j:'jLin,lgt"J'fJ(;,nt j

and ,xmtlmH:d (>Vt;:H")(plnl1i,l\on 0( snow poplliMlml\ ,1f" nm­

Hil!tllill); [,lett}):" }Ul t';lInpll', al1'hollgh o>nmwrti;\! ,,,allil;.\ ha;; ,kdinl'd COlhidH:ibly ,inn: tla: 19iiih, nal.iw lnmkt'; 1;(11

mOI'to \liM! WiJ,()OO rin)l{'(:I, b(';mli:-{l, ribb(,n, l';'''[l, 1'1(",:,d<'(.I, <lnd

St}d~h~d ~{C;1!~ 2inn~l:iHy (Rh"VUs. ~~O(2), 'il} a(lditi(u"t~ id~("r ,i! fi. .. ·dnc ..

~jnn ~n takt~IUflmrp>iS'l!'>- ill ljF" i9S0", H~}1b.~lljnK'n~ St~bsidh"\ ih(V(' r"invijiul';,\cfl til<: CilnndiMI (\)lJHlWl'dai bunl, wlih

"ppwxlnwtdy 3SIl,oiH) Il,1fP \!':.h hdllg lah,l[ in e",l€.'Hl

i:anad;] ;md (;rel'fljand in ]<!'.)l:) (LiVigne l\f'N), hi')!\Wf~i!tn

;.f)d J.(Ll~q;m Sl1ip:! :rloo (~k" ten, ,A IhouS-1mb; of harp and

hooded ~eals ilrlHudlly in ttll' Gftl;!Hlaml ,HIt! nareab ~"'jS, In till' s1)u\lv.:m hembph<:r'';, Sm.HlJ Amc)'iCZrn hll' ,1'111.', 'W(,f('

hMWS(('d in UnlgU\IY mHil 1/K' 199fh_ and till' celHuril.,,-old

hunt Ill!' sll1ih ill ,()Ilthw(;st{;rn AldGI COlltlliUCS to 1,,1«' thousand:; o.! 1m S('ilh <lnm1<1Hy,

i'lnnlped jl()plJlntio1!S abn have WffCH.'d advc-mJ impIJ(h by Inml<1J! 1IlOdH,«lthn,;; (If ('O",!{H [t(,d mnrine tl1Vif{mm<'!H5,

[hw, ('(.'sulling in di.5Wli;,nH;(>, los'. of bIN-din!': and ('sling sHe";,

.11 td :llh'rMjnl'l of 1~,n\gil1g gmlllld~ Ilk(:'\,(;:\ ZOOZ), W

vmknn pdlnU\llt:; b a 10 ,()nl<: "1'1:;15, ;md dhC\)'i(;

GWlJJ(';IKS me oH('n rda1ed 1<', inHflHH<'-ticSPOIlS(; ~llPPH~",j()n

l.ilu\cd by" ViH\(;\Y 0( pnllHli!nl~ <Hid 20()2)'

In addHJoJ1 to ('flvimnmCnl;)! forcing, plnnlpNi jlOpu!a­

twa tl{·n<1$ Mr' jn!ltlN1C('(i by vu\aritm iii iil\~ history, I)"hav,

im, ilnd phy,[(>bgkill {<ilp~dty> Capitlll b""n!u; Ol >pnks th,ll !i;iVt: lift hislory p~Ucrns !11ll! tdlmv ihnn to fmi!!;\"

llUIW~ Q,,;[m b;,~jn, hilW;1 greater capability Iff r.,conery from

,;xploit:ltii.\ll, when:l!'; im.Dflle hreeders iltc' mmr srnsHivr to

io<:<iI vCt:kmogrilpilic VaJ:lil.tiolls Ilnd llS~\)(iar('d UmitMinn, In

prel" H'SO\lfCCS nwy tenwt!!' lHore ~ltm'!y. Similarly, dlffl?re

'"nu~~ il, tilt, ioraging "Id\(;'gy of olilliids milY also be i1 filClm iH lildl abillly to hl I'HVil(Hl(Tl<'ntal f1\l\ tlldtlom,

Iknt));c djving ntariid, (e,g" '\tplkr, An~lr:lIi1m, "OlHllt'rn,

and N\~w Z"il!,md ~\"I H,m'i) IHlli\<i\ lmw'l (\'prot.\urHvvnulpnt

lh;ln (,pip('j;lgk &pech's h,;ClllSf' dwy ,;pi;nd \TWX\' ltJH(' at ",;1

divillg iwd lh\~r /)lHh thwr limib 'rhh h )w­ther compounded by;j p{J(cnllallol H'dun:{l juvI:nlir' MllYivi1!

in benthk h!;(';m~w or" f!!i~ wdlJced divl!lg

c(lpaiJihty of juvl'llik's< mfkrcm:", in tht: SHdhT:){'\

and tqll()ductlw pltkulS "Is<:) 1l";I}' In;,l"" ,-"rl';'1n plm1!.pl'.ds

metrr ,u,cqHlhk' t+mn others t{) pn:datioll, With bUlb [~PW' dudive o!mlegie, till, young are exptrS('d to pr('dMhll1, hut

ndul! 10malc" oIMiiil:; wi!! iw mort.' stlSCI;,ptlble to im:i:enwd prnlalilln ht'ClllN! il!t:ome bf"l2tlef~ must mak;, mult!pltvl:;Hs

to ,md f'U](l Uk wokery,

i\!t!JflUgh our revi('w pwvith!> ,In <1\'l;'tyh'w of pinniped populallrJll UUld, W(){llh,i<j":lnd ,l svnlilNic:df'ilt to HJJzk,··

,\"and n'<i:\01], tor f««<'il!' l")jiu!m:in!l zkd.in,':> and tilt; !ill];!!'" ()~ vnn(}u~ oi.fll.,'f p(JpUl(ltion~; ~u ~:\lf'n }'CUJVr-f frorn '(j\.'H·l~);:~

plo\i:iliol), in UHth thes,! p:Jtterns an: pourly Imckr~toj>d<

AWwHgh (;apitill br<,<;:ding ""')\lld )«(I1'l \{) ,:nllwy :m ;rdv;m·

tfig(' to ph(h.jd~~ nver oiJri'lrhd~~ H '--rHHIOl ~~xptii.U why bo{h

harhot $h!lv. Jnd ~I:('l!"i M'il lion, iiil'v\' dl'clilled <11 the same

1<l(e imd OW'l" the :,~m(' r~lJi"m of south, "),,,>f,e,,',, \.enh ha\,"(' f(·()'.)vC'i\.::d

~() \peCLlc·lll.ill'Y wherGl$ ftlillly v,'(,t:h,~m ,'kph:ml. ;,;<11 pop

illation.', hiNt' :;imilmly. alth(jugh liivin>; bdliWbl'

!HId ptlyslolog,l,.,d Hmit<itJ(}jJ ".(lHid ,('\;m (0 «)Jlvq. it ldiltive

djs(td)';H]t~)g,~ b('nlhk l~}rag~tl}5 OVP) eFirH~~ngk' ror:lgjng

ur;mkl), 1M" expl<llMi k,,) ;11, IH(' <..11111,,1 ,!u'owjl fOf (ht )1.1('(

(4culm populatiun ('O[J;lp.W of ill<' r(lr,)giIlH :it<.:lkl

~i:a hon, Ch~ HH~ oth<..?f lEtnd r 01nlou(~h then: i~}

evidcm:e that pRdatmli 11<\\,,' driw'll Ihe d"·dil.ie,, of '·:m:IIJ

j,(liitl0ti pimli.ped t:ohmilcs and !ttd'(, !1ilW been ,11'i~Um('llt'

fm: 11l(, !rnp(I\'<anct;: oj kUl"f whale p,hhlirm ill Ih,' b,?:e scal;:

pop1!!atlnn d(~rJin(:", of ,cals, $\:i( !ic1JlS, ,)!'Ie! Oll,""" of iht" !\innh

p;;(:i!lc, {Vndusl\'<' cvitkm;(' i;; larking.

(;iH'1I' ilw ~""i!\:l!1dy dyll:.lBlk n:nure OfjllnOlpe£1 fllJpllk,·

tio.ns USl.d ih;..' hJgh ot uncet over tliliruat{-? CiHJ~L:~

oj p1!111ip.:<1 popntMlf)!'l irmj.ltiD'I15 and d"dinc" it h Hot

mHU1:;Onal'llc I.U iJlLlgill<.' !lld th('~(' dvmalllC:; '",('((' inHu·

t'HCt,d ,Ii led>! !o ,innj: (kg":!.: hv t!1~' t'fkd~ of whaling Oil

,K<':;m {;C;hy"tt'lll':. ;\, Lmt! i'i ill, ICh.lj"*'l' 1 (, 01 this vl}\uwd

pujp! oW, (he gn.'d.'Nhilh:, 1'0 ()jlkd ;1 ,ignH1Ci'm1lirOpmiinn

of tIll' wnrh!,:; marine prol.l\lcti(ll1 bd\.ll'~ wh;lllng, ;wel i! J{"m;m <llId P[,lllml;;'s {?/)()::l) pr('whiding i\bundnnct' ~,;t;.

m2\tc, JJ(' (:ven do:,~ til bt'lng aCClJr(ltc-. Ib' m;ignHlld(~ nl 1 ilL

!.'.ftc"l would l'lat'Q bU'n ",'ell grcater. III a o!lllpiistk: seme, h:'" PWdUC1()lllll'ing ('{,ooph'd by \Vhilk~ me,tll, j)utl.:lll'iaJly mOHO

rill' othc:1' c;c,:;m ,XH)',tmWf: •• );ll{"h ,'ffn;ls haw.' heen

ill tlw ~t'HJ1h,'u, ,',eu",], whi~n' the: N!flO"';,l of krill"(I;lting

·whi1le.~ (');;\'~n!;ihly !NI1\) in('C:1S{'S In ('I(lu:r krill (':om;\HlH:rs,

of tbis V()lmnc). l!OW('VG1, ,1, Pain(' :t <If thi, wilum('l

points om, fond \,'/('1'1 dyHMU1rS ,m' !iUely so simp]". iiitill.cgh

\:uHlpk':c lood web dynamic" nf Illi, ,;on iHC

O(l:'ttll t:('(),.\',;hmu, lh" 'NlY 1;\(;1 tiiill pinnlp('11 PVPUIHiioll

d'jinam,(', ht ,,0 poo;:ly 111'«) \udHl,)fl"\ 11)0\(\\

ub.:.:s UK di,1ind p')'"ibmly ,hai luHutlH.V' ()(

wlwlinl~ i\!(,' (t$s()(itr!Nl '>'lith sO!JJ~' of dli.'> Illln'lldiniy.

literature Cited

Allglios, K. P. <lnd t<. L Lotlg<'. 2(l02, ,\I,,,,/(,, !limb;;' Iliml!1illii silJd:

aSW'.\.'iItI<:ni's .. litO:? I).S, D,'pawm:nt of Omm1('ri:'~, Nt. )(,;\ 'r,odt­

nlGjj !vtvmm;m(lum NMFS .. Al'SC-I.:n. sGlak: ,1,ia,k;;l'h!wr)f';,

S{.'lt:'flle t.:t~t\t ... ~r.

Allh1Il\(L j. \1, Y, :mtl R \'1. VV~\l'nd:.l". :WD.;5. (rtiJ'Nlh arid mndiilun 1H AH,",h:~rhin 11Jf S(.'~'l5-~ (tiJ{ i{K{'ph~~ ,,pu:dllus -;luri/',.:Iu.\) ( .. [~tniv\)r;~:

r~Hnip:;dh}J, lht~j)<ffjdN bn:1r!k!f (}(/a{rl<Z~F 50: 5:-l>()(i

/u~dn!f''\"r ;{~I"!i~>\)n~ D. !~nd A . .z.nv~l~ ,Uonz~-de!" I ~)9,+' Ecn!(lgi>. d-i

t>!.ctnrs th£\t (lct~:nninl.: d.i;,tdhl~l €UU ~Hld ,d)UHdalh),: of th{,~

CidihH n.l~; :~~:~¥ 110H /.<-dophr.t~ ~ afityrnhuJWi in lhc (;H~i of

{'~aljfiJ(!d:/:,. (.'iOtl'idS ~,fd!hid.~ 20: S35·<'S3,

g,nf;\\, /I. <wd J.t. MOUfi,IH. \97<1. The Smlilwrtl \'!"pham ;,,;<11, IcJinw\xo il.'()lliI"I, ('{ i'u"'O,i"H bland, CW1N

46"25' .>(lUlll. :; !,,'o;:,' (';k;t, Mammalia·l?: ) 4:5- i ;>4 (ill Fn,'tH.'h), ibill'l()kIlYt;::W, C./Io USU. A Hmk (;I'S on Sail Nicola, l$i;lIld,

Clllifofnil!. JOImml ,){ MmlmJ(d{(~y:\ 1:17,,"·180,

] 97'11, /, l1wifuj it)! the {'vn)u\iol) of pinnip8.l polygyny.

/iq)iwitm 24: :;'16 S.\<'i, H(\Jlg~\'{11~.j,L ;ltul.H.\J [<;C~\,<t'\_ 19x~, -l'r~~f\d.\ rn \~r{1!);;'a1-(,'t' '.;'}i:;~ ,1=g(­

Lo-hudun l.·nr:{-,ir{'~)p}i(lSUS ~tt ~~~,'\.u~11 nnrhinty~ ~i:n. insight inib

Ant:tH f k n1adn~" h)h'l\ln~t)n~, h) .I-tf){,ttl" tl~ -"Wi! i~'<;;l {n.}.,/!\ Wid

FJott WeJh, ·V\~,1( SkgtrH~d} 1< H. ~ :OI\OYl dud !t "\A. h~),.v~, {>&r.

\\{din: SpJll''';'~l' Vi.:i\h!j, Pi', (6)' iII'S,

g-O~:'{:nn, j>, L'J L }.·L HJndd j \{ 1( ~(h\Y\H~/, a;u.i L L. lh~Hgl,"';{WL

] 99ft P1 tpuhrH!7n grcAdh ('~r i\nHt:al,-th: hg ;-,t"<lh~ IbnHd.dHH hy ,) {Oti pt('d~Hoj;') th\.' ;.;c(?;i;) [;'colo,gF /9; /J{H(., .. ~~g/}-.

HtY('\,i~n( tY.H. !~JWl. H,.ok' oill ~HfUip~· fDmm!8ab In ,JqU~Mic ('reb\,\

h~~"n~c lJ,Hhs( i/rd?),J.Y j'r!'l8h\';\ S~~fk<'~ IS~; ;,?(); Z7·L

1\<"""11, \\'. n .. S, L, filns, :;,1. IVi,'H()J), ;,nd n,,!, HmH'~'. ?1.1t);·\.

)'1httP"'! l\rld t'l('wbom lif"lmtulv trilih (\urln~ TWTi,Kh (Ii

{'Olltr;J,,\ni\ f>')j>\1!;ll,on tr(,n(!:;, J!)ll'liO!iu!!) hI' explaining jhf

{k"chnt' of n{·!.dx~ur s<-';~I~t t'PhtKtf d(uliltn'~i U~ j '~db1L( h:hUHi. jOUf­

fINt vr7.(~oh~'{,r 26 g; 1 SS·- J6.L lloyd, 1.1.. F,1\I:L P"l' FH1<1UCtillH imd lh"l.rHmtion of hr('('(Hn~

AnC_utH\' fur ~\,;-lb (Al't·lt:Ct'jflulu:{ Hazell'(1) ,:H S.o~lU1 G\~t~Jg~(t

illIimUicSc,((,'IIU"ilr: U ;n. 7.0(11], s;i\k"dqwm)('rl\ [('limt)' in pi!wiF",.h: c.onl!J\Ung

c~§FH~d ~\l:'ld jn<YH1W bn;('d{,"rs, Fmvf;ml;;/} i'\'u/.oSV 1~1: 623-.-630.

B-uv\.t LL, ,LP.Y. ~\fa~fuldi T. 1'L~ftonJ ;lnd f.F, ('.!HA,dL 11;t)4

F~)r~~glng n~,.:b~'hiuur uf /~Ht~H-'L:ti{' ftar ~;{'iJb {hwing fwrklfh Ot

C~'rntraf;:nng pn~y {lhHn(hnn,'~ )UUUNr} 1.1/ .~HiJliaI Feolasy 6'k :.:-o::\·~;; Li.

Boyd, 1.1." J,J: t,,(n:aiL N .. I. L1JllU, and IC ltdd. j1j!)5 l'opulilt!<.lt! df'!1~f~g),i'jpky (Jl /~HHHi_ rk' §ur ~,\.fdh~ rhe 'CO\l:) of l'cr,toducthJJ)

,<htd H1)L~Vl1-~tL:)Ht _tnT Bk~l]),sl(nit'$, fowi/o! u{ Al:iwal J:~~(jhwr

,,'J: 505·) W. H(>yd, Lt., N.J, tunn .. (Hid"L B,HtUfL fl)9~. tHl{l !nr~

i.\gil)l~, d",Ul(H:lhlic."oi \a,:\,\\lIll; AntM(1K fur wah. jummd or ;'iW'iTIUi J~~'(}lns)' (-AJ: )!?-·~92_

Boyd. f. ~ .. ~~tHJ A. \-V.A, ;vha-ray, 2ntJ'§ \'kH~If"ndng.il nM{l'ol' l';.~OSyS.

km \l~illg n:',jlou,es Ilf Upp('[ trl1pl\k l{{vd pK'dators. jll/Wld

"i!lllimni j.,'oioSi' i(): )47 .. '760.

[,,'yd. L I., TK W"I](~t, elml J. Pone{;\'. 1'1911. S!aL", "r ~(>t1~h\'m ,.'~epi::anL se~ij·; al 50u~h \i'ft).rg!iL !tir£ari.'tic Si'h'HC~'" g: 2:i/c·,~,~,~~t.

HtHll~1} ),j. 19i~~, ?\i£~'.ai'j<' m'~lww_~1 ?i"'1;wt PH~a-E~~~~Hnhl.'_"U .. ~~_w.\ Pr13·

ie~ t Repud \y. 11 ;~J \;\:~ J /-,A~ ,'t~1d \\",',1 j·S. JVH(,;:g;.: Ab:;{·;.::i

I k!"IT\owfli. of Frs!! ;:m1 Gal1w.

H~!f:tOf!J lLiL, r ,J\rnbotD; f,L, Ftoyd) :kL UiJ5h~r< D. \/{:tfUH~L ,HH'f

L VV1H~~n'l{)n, ~9i}:", S'lp;\~HiG~}H difk.'10ncc) in \'~<~;~Jl~nJt 1f§l':!\~1 uE

~oHrh(~rn <Jepharn 5(;;:;islrcuH j'P:f' ,,:uh./\.nLnl rk hbHd'\ in ft\l.d'-"

tbJ!1 to pnpu L.ttton I...kdin'l;;);, H~ /~n~an:lh' -t;D.~~Hnti't1ifi<;,';: -lip,;,; if.'t?

"tmelun: ,wi! .\w\'lml. 1i',.y,H, \1/0lUH\, ed. C:i1mhTi;!~;", \lIe

CuntlliLlg!c Vnll'n:;ill' !'i'c". pp. 3:\5·.:ng l(qUUW{I;Ut, 1'1.5., A.E. runt, W,IL ()m~)HH~f'n, Ml(\ J',A Wi<'["

f.~n~, P}!,};) , Htv pH.ens of J~HHh' i;!)n'.;lHHpHon. by thJ..; Cape hu

~nl on (;li:dw~ ;'Iv.! (;lh:11 r;,fb of !IN Cqw lnl:~s, 3, Model .. Hni~ tl~(' dyfHHn~.:.:~~ 1.)£ tiE! C;~p\' fur ~;('at Afc~o;~epJkdus pr!-;fnb'.~

f';J'hUffi.~·. :-iUltihAlth:aujouln/.f(j{!dJuiJ!eScio?ii" l6: It)l itU. Ci~ppO~!,:C\ l. L L. 2H{r~, ~()H~h Ankrlc~HI -;(71 Eon, in Fnn<·doV~1(H:l of

nhrritk' ~nmtuna{s, Vi P('~i in, n, VVtir~l}~< ~Hld J (;:.\1. 1 ht'v:.:b .s'dl~ t~(h_ S.:~n Olt.:~g(): ;\c~d~c.~nk PiT~;~~) l1P, ,~lIK<{S 1,

{>H·U\l.l~11 I.1/" J. hddo~v, K .. /\y fu~a~~y, &'L-~\1, (vrHf:01 flU':! f. Hak"t.

ZeOt, U.S, f\'Kifii' nUiJ;ur ~)WnHnaJ )t{)c).;. (!S5-e:;~nh"nt'J~ 20tH.

NOI\:\ T<.!,;Jllii~;11 ;\.1<nI10ritm!ulll N~;JFS·S"\,1"SC-317_ 1.;, lolk

C'\: :-i0i11'hwv,\ h,h':'rics SCihIH' C(:.Hjl~r.

Ca~ n:H.1, J. \<, rd,:VL h .. jtJto,,J. Ha:dol\;',.L Hak<~r~ !CA. Ff.HHf:YJ ~~nd "h.t ~.()1f.V(y( 2002. V,S. Padtic nwrine Nium.tf1d.l.s$m.'k: '{f5.1\'S,Hn~YHSt

2UO'J. N{l/I,', ]i;:d,nk;ll ~,kmnrMHhml N:Vll·\·:--WI· . .,C:j'th. L~l ,k.na,- ('A~ ):(}uthw~'9 H-<-.!:Pfic·\- :-idt'nrF CvntvL

CIlO[W1, L(, ;W,c[ H,I). SkW,Ul. l(fth PHW)gf~'ph~' 01 H<.nli:<:'m

dcph~Ul~ ~~(~:1b iJh~~mlX!.~ <'fN);u'":,rhn",f!J\ 'j9 ~ 1 , .. '~ 9H? l·;?.:tnw~· :.~'~ {,i;

1.f69 9/l.

(ti)~V';-, ft,P l!}~jl(L Hi'pfodndl;"i:~~Hd forngJHg tra·rgZ'lk:)fJ.t IlJ r;h

1ntH~,H'h" pc,ngu1ns j BH)lnf,,)S~~t"~9J ~~nd pH~Hipt:ch~ 1H1f,Et \),U~Hl~ f.tn

iiL' hbtory p"tklm. ,\I!I;',ilriJJ lUi/loS;>! 31! l! lUD, ....... ~ 19~-H~,t, Rt'VfOdlH nv\' dad ~u:j(!gldt;" (·n('f~t·OC5 ()f p1n,

H~rw"h: 1;H}pHcaHcJn'; fo~ lift- h~~iiory patt~~np;) in n~~Jhiduur (4 {J;!1nip,,"~i;L D, i~>i.}HDtd. ed .. Loudon: CikiPfthH! dud H{;/U! pp. :ltl(},:ili·!.

, 199:3. Th/' wlminm,hlp heiW"'!i\ rq,wi!ilClivc, and fOr" Hf(inJ; ,'"}~«rg.Ft"ks and flhl (;voh~tkn~ Df Ok PhnHpt,;dj~L in \htti'fu- 11lanunals: adwJ}n:es i~r bcfwviuHraJ ,wHi PVPif/ilij.{}jj

!Jial'~~r' LL Boyd. red, OX,10t>/: 0);101<1 Un,vN:\Hy l'rt'~,j, pp, Z9:S:Q,j

CO~L} ~XP . .( J-F Cft)X~)n! and c. DtKk. 19H9. r()n1H~Hg t~th:tJ;t)tj~~ (tj Ailt,ucik: fur seeds, !lrct(J;:~I'!ltilw Sf/lcdl", in .rd,11i(11} \0

(h~1n~e~ in PX('Y dV~ij1abH1tr, EnAo7Y TO; 596··-6U6.

(~lJqa f), e. i~nd .~.J, (~,lii"l), ?{h)f). fo.r\lr~!nFf ('n~;¥rH,;~t!cs ;lnct dlvii ttt twlmviolH of I.)[t;ltinj! N0W L.e;l!'H)'! ,wal Hum" P!wcmflo" iIImk'ii. J<wnml ,,{Exl'erimmt,ji 1l}{,I"XY 2(n: :\6'\5·- :1(;{,5.

, 2003, En'2'q~(:'th.~s oi' {\: b(.'.othk (UV0J: $(?~~s()J)td ~{H~)p;jn-g

ec;)jogr {l ~h(~ Aus1raHtln ~{Jj1 lhrfl~ N{'bpimnt duere-r}. J:({)lo~\.[INtl

.I.:joll{lgwph, n: :':;"'.'1''1. Costa, !J'p., (·i.J, Giiles, i!m! M,E, Ce.wbd, 200!' A;,l()bk <livt.:'

limit: How nflen doe, il flfTtI' in nii!mri COIU[ian1tiw

Bi(!cilmriYl!y and i'ily:;jHloSY hilt :1: M01i'Cu!m awl [illtgratiw

l'iW,i!)ioSY 1~'9i\: ;nj··71U,

Cost", PY" CF, Kuhn, )"LJ. \V('IS~, S.A. Slwff,'l, .aml J.!',\,.

/unonkL 2()04, \Vh\!H do!:, plry$i()lnt~Y limit tile, fmal.\lnA

hehaviuur oj fru:1y {Bving rJ)~lnHH;~h? tU /\n}ttw.h ~'lm.l ;"'fivirof{­

W-<.'fU:\,' p.f{/t<·i~./dhrs) ~~((j?r? 3:n! hth-nwNolhd Cj'}n!~~roh.'!.:. WI COtn{NU­

(iilv.':' rhY:ihi1oXF and Riot. he~NI\t~}~ J\Houlu{(.;/Nahtt, ,)'m1ifi .ittKa~ ? ·J3 A1(;;n,[ 2(10'i. S_ ;Vhlffb ;Hlll ii, Vi\,joo, j,'(K II1H',I',illionai

Co-ngj.'~$~\ S~~rk) 1:5/~, \-\n t~U~rdaH~~ hJ"x~vj;pr. Pl" :l~v --,}66.

Cfl~t;L D.!', i!ll(J 'UvL Williams. 'HtW, 1,iUl'in<: !Hammal f:HCl'gd·

Ie" illlliolc~\~' IIFm'llil1l' lliillWl/(il,. j, E, R~\'JJnkh lIml S.k ]lum· mel, <:d~. V\!i"lliHglO'L DC: Smilhsonl;m lp~HhHion ('Wi;;, PI", 176~),i},

J.JC!ncrc, 'fA. 1994, The family OdO!!C!lid0C A phylog,mctk nn;i1y,'l1'< of fo~:~H and HV~Hg LIXiL PJo(tJ~!dh15?; or the S'l.U J )iclfl

SDd<!iY o(\'aiHmi l1i'ft1Jfy 2,9: Y;f"l:l'" nt:mr:~c 'LL A. lkrt<l, ;l1ld 1'.,1. tHlam, :WO:t Plnnip(>dlmmph

rVt)lmionary blog;·()graphy. !JiI/leli/) i.{lhl' ;1INaitllt! MII)i'UiIl o! Na}~lh!J His.iurr 219: JZ, l6.,

E~te;.lAq M.T Tiak,'f, U\L \Villiil1m, imd D.L DO:ll" 19%., Kilh.:T wh-r;k JHedaUon on S(,;l. olh~f;'; hHk~ng ~jf".\,.;\~ni( ~IHd

n(~ill~.Jl1rH" e:..~o.-;y~H·~11"1~ :1::if;;;lice 2HZ: ~~!3-"·"1?0.

Ev(CritL JU.l, imd lU, Ht';"'-h. 19k':', ~1Mil.i(' ll),IIIHnal·!hhnic' lnteractHHL':> in Ort;gOll i'!~~tt \V:'l\hingtf1H: ·nn o\;crV~eWl In

}\<taIin(' ?ffanil?L~h;; C(u}tnd'i vdth fi<;Jkr[~;_>;~ tJtlu:r JiWlutj!:(FdCnlpmh-­

!ems, !lwi iI'."wcil II!!";;]';, !>,(", Ch~pllw,n iH1tl L L. E1.li~[hift(h.

C"];;. 'i:l'il!1s;"clinm of !ht, '17th )ionh AllH.'tk.lr) Wikllih- and

N,,!w:al R<;MHHn,;,:\ CcnIJ(:rN,,'(\ Vi'''hingmll, nC: W'ildllf,,',

M,ln'lf("!lH'n! fnstitut!:, 1'1" 2~3 2(;3, FNh,,("·v, C. ,\.1'.17 1_ OlstribUlioH ami pOjlllla(illn of seab at pup

and molting rooj("nk., in tJr.· Sea of OkbNSh, Hllill' Mlilli1it!1,

es!;,u,'iu :';u;di}mp.],'h'dumld'8!;,W" lil,til!£H Hyhj!i).B''' Klwl:)Jilir/:l/{T

i t}kdwmgmf!! :N: l'll .. <j'J (in Rus;,;an).

-~ 20ttO. PJ)jJ!dUtlWl l;ivh~~y urke-!'1s-.w~dNjc'd l<)ttns ()j su;!s 1ind

!h.-i.} ~ol/ fU Ih: N<jtfh~'ni r"lldt~{ i:(;)S~-<:<'rffn't, \~ins\,{}"i\·.., ( \:nt.,·f h lr

1-tH~:..,iJ~n f':_n\.'il{'H'ji))d1t~l! h}hcy.

Fddbnnp, S.H" R.L !Jt~l.,\Hlf> jJ!1d f~,A, '\ntutHd~!>.. 198ft. DiVlftg

l:itHV:fTn; of CdHi~Huia ${<1 IJon~ .. >:a!(~{}hn:1 ((~nf~):mi~nw_"

LONadjd~j jWN"Jd! (if It,/ufoS>' flh ~I):l~~,-~x.:~,

h.Lvnd~Jt ~.r, :H~;j l .-';l.til~JrW. J'}){>, {.Hnvrv;~Hon\ on b.tnb ,uId

{,(>ali-; :H H()HVH-{~!,'~L Xf~r.\'k P,()/arinM-iUUf Athal\ 1 ~)7-(): :167·<~71 ,.

h'lf(i1.(!;'·I" L P.S, li2tBHnnnd, ,n~d /t, r\guihu-. i~FJ1>_ SI~i\H'~ ot" lh~"

~,i!.>(Hli::.~rriHi('an ~nHnl:: \,<id .\-lw-Wdu,r\ UUUW(tw.\ in 1Iw' \h/~'\h.·rn 5~lh;w;~ ;}rHi tl"W In)p~j(';1jj-OJ1S {~f a Ina~:~ Bl{.8rL::dity (~\'t.::nL Afa;iH:!

t(()}OS.y-·F't\,).'U{;'.C3:nYh)~ ~H6: 2:!9· 2.(jt, Gales, N . .I. ,mil P.J. Pk;(d.)>.:r 1999, ,\t.mmLmc<c, db\rltmlkrH :111d

jL~itU$ '.>f th.:: \\c'(.v lcabnd 5<:<1 JkHL Ph{YtH~'}o.~ lwo.~~'t{ i-\/ildJif>

lk,~{11(/1 ?.6: J5-S? r;"kli, N,j .. ]"D. Sllauglme5y, <miJ 'l:h, Dell n i!>. 19)}<j, DistrH,\!

nOH, ~~h~i$~(h~rH_'e and tH'-('{;d~l]g r:yde of lh" )\(~'l~ ~.~!Han spa JkH1

dth't"I.'{j nvi~nnfrH~ll~:I: Pinnipp-d~:I), }munrr! u( /()Hf<iSY (J,(WdD;t) 2:i4: :t~3<~lO,

G('lltr)', It L ,mel J!'L JullllSOll, 1.otS!.. Prd,(\1"1l hr Wi> linm on nort"lkf!.l fur j.(,~td llCOH;:Hct,." J1iUnHJdlfa 45~' .J23 .. ,.-"J<":iH.

Gvcu, /I,e .. , G- (~u>:u, ,md H. ()J('f;. 20f)..1., lI;!llILlt u~,' ,lwi Pl'<> Ihl]itl;;, y (lC)HOgfi1phk {'vull!atlon of t be' Hilkdly ,:m!il!ll;;':It't!

Mi)il!WWltWilll H'I<mk "'iiI (Mmlrll'ims 1I!1!l1iKhlll'j ill lhe Cmd~H

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WHALING EFFECTS ON PINNIPED POPULATIONS 359