GeorgeE. Mustoe and CynthiaA. Carlstad,'Geo ogy Department.Western Wash ngton Un vers ty, Be ngharn Washngton 98225

A Late PleistoceneBrown Bear(Ursus arctos) lrom Northwest Washington

Abstract

Thei]i"corcno|aprrria|shllindele\enl.l'Lhoftivlialllarn]g]&ielselljnlrrs|romthjdllcr kno\L,r n\ud of bro$f h, ars h rhe Pu.ifi. \onhiresr. AlLhoLrghrhe eurh dispersal of Li. akl,r ;r \orh r\nrfrn ! Iemains enis- mlli.. the lse and o1 \,sr. North Arrrn,a lrom the Al:rska/lukon tsi,)r at the close ol Lh| lce -\ge.

lntroduction $,ho noticed l$'o ja$ fragnlents containing'1, large teethexposed b,v coaslal erosion along lhe contact The discor,erv of a bcar skuil liorn Late Pleisto- zonc bel\leen fllaciomarine gravcl and overlving krn sr-,uthol l)oinL ccn(' stratain a coastalblull2 dune s:rnd. A more extensivc search in 1993 (Figure I) Partridgc. Whidbev lslanrJ.S'ashington yielded a large ser:tionof skull and 7 aclditional rrei!' contr-ibullon to our providcs a signilicant teeLh(! igurc 2. i:i). lcrowlcdgeof lcc Agc lile in thc PLrgetSound rt- gion, an areathat containsextcnsive exposurcs of Description glacial and glacionarine scdimenls.hut lc1r'!er' of a large fragnrent oi cranirrrn. t-l'r';rt-lo--i1.. lrrterl'retrli,,rr.,,l I'nlri.t,,ric rni Thc hrssilsconsisl (l'.1'o. mnl lifc in this region largel,vdepcnd orr inferenccs rdth the lbllowing teeth: U'. \{']. and alvcoli made fr-omothcl region-r$,hcrc Lrnusualcnvrron- fbr vestigial premolars Pr and 1". Three detachcd mental condilions produccd great accumulatiorls uppcr incisorsrlerc found in the acljacentsediment. (Table in ofboncs. such as thc tar seepsofsotrthern (ialifor'- The dentition l) cornparesvcrl closelv size and form ltr modern Lhrrr drchs funibili: lgriz- nir. liuano.lepositsoIthe southlc,.ternll.S.. ancl in rhe W WfT rnammal collec- the pcat bogs and olher sedirncnlsof Alaska anrl zlv bc:rr) specimens tion. thc lcclh beinplmarkcclly- larger than Lhoseoi Yukon Territory. Tn contrast. skeletal remains llom 'l the modcrn black hear. hr-'-vare the Pacific Northwestlrc usualll limitcclto disar- U, anteicoruu. clifferent from thc tuo other bcar spe- ticulalr:d fragrnents lhat were disper-seclprior lrr cliscreln:rbll cics that inhabittxl the \testern f,.S. cluring the [,atc burial bv strcanrsor -ecat'cnginganimi s. l)lcisLocene.ArclodLs siarus Copc (sho|tfacecl bcar) Icc Age ,'efiebratc fossils h:l\'e Pr-cliousiv becn and Lia|l.! opiirtLr Sc'hLrltz.The httcr species $,as reportcd 0.clmthe Pacific Nortlllcst bv Scotl (1967). originallv bclicved to be an ancestral black bear. Harington (1975), Nel'comb arrd Repennirrg Latcr clescrilled as a variant of Li omrncanu.r lKurtin (19?O).Peterson cr al. (1983). arrdRensbcrgcr et and Anderson I9B0). However. Craham (199I) a]. (198,1).but this publishuJ lecord is lar lrom considers Li r),lritru^sto Icpresenl a smdll-slalured complctc. For exarnplc. a ground sbLh skeleton face of Ll. arrlr)\. Based on cnamel t'ear. thc Whid- \ililegalout sp.) discoveretl in l96l during con- bev Islanclbcar wasprobably at full maturitl. and strudion of tbc Seattlell acorna{irporl is a feaLured pcrhapsol advancedage.'fhc inc;sorshavc bcen exhibit at the llniversitv of Washington Buriie abradcd to about halfoftheir original cloun height. l'luseum. arrrl the \{cstcrn Vashington l. niversity :rnclupper lcft molar #l conlainsa verl ltrge de- geolog-"-clisplal- collection includesa large rnammoth ca-ycalily that nppears Lo havc dereloped follol- tusk from the Skagit River r.allc,vfound in 1979, ing frilclrre of a cusp. The upper lcft canine is anrl severd rnarnmoth boncs cxcavated {ronl cen broken ut the tip. but lhc absenceoi. lrcar-marks trrl U.r-hirglnrrih lq.r l: ronc ul th,-- -1,ecim'rr' on the adjacentenamcl srrggeststhis damageoc' have been dcscribed in thc scientific litcralrrre. curled follorvingthe dcath ol the anirnal. The S hidbe_,-lslnnd bcar ibssil r,;as lound in Geologic setting 199I bv D. J. Easterbrookand C. A. Carlstad" Thc geolopl ofl'lcistoceneand ITolocenedepo-cils 'Presorudrlress: l.l8l5 l01thAre \li. BoL|cll.X.{ 9801I of ccntral S'.hidbcvTslancl has bccn discu-"sedin

106 Nolthnest Science.Vol. 69. No. 2. 1995 l.l 1995 l,r Lhe \r.th\rFl 5( r" lil' A-!,.iillon A "gr'ls .ci.nrd 1250 1230 121"

Figure l. l.(!,ation nr!t,. cletailbv E.rstcrbrool(1992) and Carlstad(1992). ginning to be filled b,vdune sand. l'hc thickness 'lhc fos-"ilsitc occurs in an arca of kettle topogra- of the sandunit variesfrom 20 ro 70 m, plor iding phv that lbrmcd llhen the rctrcaling ice and ac- a prominent stratigraphicindicator ol thc abrtrpt companvingisostxtic rebound c{)mbined to expose transition lrom glaciomarine deposition to aeolian a 20-lJ0 krn uide zonc of unforestcd louland sedimentation lollordng the final retrcat of the ice scparatinEithc$aters ofl'ugct Soundliom tlre ad sheet (l'igulc 4). Although no other large fossils jacent loothills of thc \orth CascacleRange The havc been found at this site. the sand beds con- lossil appears to lcpr-esent disarticulated skclctal tain rvirlely scattercclfauna.l remains, Senerallvcon- rnaterial that i,rccumuhtcd in a kettic lhat $as bc- sisting of a single bird or rodent bone aligntd

PleistotrrreBroun Br.rr 107 ''': ' i

,'I

.t il IOCM r*r*

Figure 2 Dorsalriew of pari.td and frontll porrion ofb(iar skull fron Whidbey Island- The anterbr rcgion hm bern destroied bv $,,lLheringi bor(js ol lhe zygonari( rrch appea. Lo hdve been losl during sedin,"ntun rra,o1o,r prior t,, ti,,rt burial.

'..: 5CM ee I M2 I ;Ps ta1 ul P4

|isure 3 \.ntr!lliew of ul,rrr.ja$. sho$i,,!ic.oln and 1rofile vieos. I : incisors-C : canines,Pl dnd p3 = lesric;al premolars. I'1 : prcrnolar. l\l I aftl lI2 = m,lars. Skerchol sku is redu,,{ 0.5 X r.t!ri!c ro phorogfalhs.

108 I'lustoc and Carlstad = 3

!

l 1i

. .s I 1 3

': a = ^l l

L=

aa = *: t - E

= -li ;, .- -.E *':, : :"i a -i = a

ie - '':

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i-

':o r:j :: :E

2e :

Plei:toceneBrorn Bear 109 m Recentbeach sedimenl lt1

Recentsoil

ll:-:: I t.r..:-l Holocenedune sand

Eversonglaciomarine drift l-----1 appproximateage - l-- -:l 12,350- 11,900 BP

X fossilsite

Fisure1. Srrlrisraph!ol lossiLsne.

parallelto the sedincntarv lavering.Shells of tcr- alone.'l'hescrre all prescntlvconsidcred to ct)n- restrialgasLropods occasionallv occur in thc same stilutc a single spccies, {-Inilrsdr.to.,r. which also in- cludesr.arious European and Asian brown bears (Erdblink Thc ageoi the Shidbev Islandspecrrncn rs es- 1953, Rausch l96lJ). Tr,".osubspccies timatcd at 12,000-11.500 BP. based on strari- are forrnd in North America. L. arctos horrihilit graphicevidcnce. Olar.iomarine clrift underlling thc (grizzlv) and U. antos middentlorf. (kodiak bear). fossil sitc conrains mollusk,.hclls that yicld,.C Kodiak trcarsoccur onlv on the coastalislands of datesranging from 12.350 + 400 ro 11.900 + Alaska. probablv having evolved in rhe Late 1700 lll' (Easterbrook1992). and rerrcstrialhab- Pleistocene aftcr glar:iers isolaterJ a local grizziv itat Nolrld not have existcd during this phase of population. (jrsus arctos subspecies can not relia- depc,siLion.Hor,".er.er. thc sedimr:ntcnclosing the blv he distinguished bv skeletal ol denral charac fc-,ssildcr.cloped drrring the earlv phases ol coaslal tcristics. but thc radiation of Li a. Ao,n 6i1isacross uplift. at the onsct ofthe ertcnsive aeolian and flu- much ofNorth America at the closeofrhe Plcisto- rial clcpositionthat had begun ro dominatc'there- cene suggcststhat the Whidbev Tslandspecimen gior h;- about 11.000 BP. The age of the uppcr reprcsents a grizzly: [,t,a. h,onibiLishas prcviousl,v podion oi the dune sand probablv cxtends rlcll into been dcscribed lrom Late Plcistocene asJrhalt lhc Holocenc. but no dates have been obtainecl cleposirsar 1'lcKiftfick. Calilbrnia (Schultz l93B), Ilom thcsc strata. and skcletalremains of glizzlicshar.e been found at five archacologicalsites in easternWashington Evolutionaryhistory that range in age from 9.000 ro fl50 B[, (Lvman r968). Thc taxonoml of brown bear-shas undergonccon- siderablc revision sincc Merriam (l 9 t B) desr:ribed Thc discorerv of the Whidbcy lsland lossii t'16species and subspecicsfrom North Amcrica ollers ralrrable insighls into the dispcrsal of

I10 llustoc and Carlstad N. rurctos.fhe oldest knon'n representalile ol the speciesmust have occurred earlier than prerio sh spccricsis from China" rlatcrl at approximatclv bclicr.cd. pcrhaps r.i:ran icc-fi-cc zonc bctr,".ccnthe 500.000 tsl'.'fhese earl,v bror,".nbears appear to Laurentide ice sheet and the eastern llank oi the have descended iiom U. etru^scns.a common earlv Rc.,ck;-Nloultains (Hoplins 1967). l,porr rear:lring Ple'istrrccncspr-'cics r-ccordcrJ frorrr liuropc and lhc soulhcrn tcrrnini of thc contincntal glacicrs. Asia. Li. arclos probably er.olved iiom an Asian blown bears t'ere relativelv unrestricted in their racc of {.r. ert)'c&.\ lhilt bccanrc isolalcd $hcn abilitv to travcl. T,atc Plcistoccnc LI. arctos fossils glaciers :rdlanced -routhward Irorn Scandanavia. occuf in Ontado (Petelson.I965). Ohio (Miller anclulong thc Ural llountains. Thc datr of origin lB99). Kentucky{Cuildav l968). Wyoming (Kur- of the nerr species is nol Lnown, but Li. erast:us t6n and -{.nderson1974). and Calilolnia (KurLrin Le(ilme e\Linct sorner,'herebetrreen 1.5 and 0.5 r 960). ma iKurt6n J968. I 976). AlLhoughalpine glaciers occrrpiecl high vallevs {i atrtos clispcrscclrapicllv throughout Asia and r' ithirrthc \urth C.r-..r,1"-,m'r"l' ul the morrrtrrirr Errrope.eventuallv emigrating to NorthAmerica arearemained ice-lree during the Late Pleislocrlnc" r',hen lorrered sea level createtl a land blidge con- posing no lbrmidable obstacles lbr bear-" on the nrctingSibrria ancl Alaska. Kurti:n (l 963. 1976) rnor'e.The arrival o[ U. art:lorin the Pugct Sound has suggested this nli€lration occurred during the lol'lands is not surprising, since the region'-r tenl \l,isconsinglacialion: l$o ai .r'.1orspe.imens fronr perale dimate uould hare oflered farorable thc rlcslcrn YLrkon shot radiocarbon agcs of habiLat. 41.000 + 1.050 and 36,000 + 1.I50 tsl'.Al tcr Lhcirarriral. brown bears inhabited the tLrndra This wcsl$ard migrationis urnsistcntlith thc region ol southern dlaska and the l ulon. an area behavior ol rnodern grizzlies.t'hich rnav travel con- known as Rcringia. This rcfLrgiumpror idcrJ a sidcrablc distanccs from scason to scason (['lundv t.rrnrcLleLrrt g.oSrJfl,ic'rlh i-nl;rte,l enrirnnmerrt ancl!'loot I9713).Although the diet of theseorn- for rnoose.bro*rr bear. and othef aninlals that nivoresirrcludes elk anrJother largc mammals,as rcachr:rl Nolth Arncrica at a timc lhcn furthcr r,".ellas rodents. birds. and lish, much of their mrtri soutlllard dispersal las blocked b,v the (iordille- tion is.-,btaiDedfrom pl:rnt m:rlerial.parLicularlv ran Ice Sheet. In addition lo the newlv-arri,'cd r-rxrlsand bcrrics. l\laturc conilcrousforcsts pro- Asian emigrants. Beringia -supportedlarge popu- vide scant Iood. and grizzlies afe attracted to open lationsofhorscs. bison. clcphants. and rnanvothcr ifcds it'hcrc ample sunlight allorrs the clelelopmeDt speciesthat occupied both the tundn of the iir ,,i urrder-tolrrFlclati,,r. Thc-. , ',rditi,,r. ,,,1.r" rrorthand the grasslanrlsthat lav southof thr-'zonc in subalpincmcildo\rs. along thc marginsof ldrgc of contincntalglaciation (Hdrington 1989. I'iclou lakes. and near the coast line. and it is in the-.e 199r). enr,ironmentsr,ihere mocler-ngrizzlies are rnost 'l'he aLundanl.Plcisloccnc brown bcals displiwcclsirni- rnigration ol l,r. arctoslrorn Beringia to mid- l:rr geographicprel;fences: lbs-silsiom lJ4 Dan- contincnt locillions is not hllh. undcNtood. Onc in- ish sitcs indicrtc that from 11.500 to 4.500 BI'. Lert)reLation(KLrft6n 1976) is LhaLthe clerelopncnL {,'.aftlo.\ miilraLed from inland locationsto coastal of an icc-frcc conidcr along thc 1\'fstcm malEiin habitats as ibrcst clcnsitv increased due to plant oithe Rocl_,-N{oLLntains at 12.000-l1.000 BP al- succession(Noe-Nvg:rald I 9tli3). lolccl brou rr bcars lo rcach an ar-canr:ar-thc plcs- ent frlontana-Alberta border'. Humans mav have Paleoenvironment follortecl Lhe ice-free paLhwal during the same peliod. as su€{gestedbv (ilovis age sites scattered Environmental conditions at \t hidbev Islancl lele acrosslhc Lnitccl -(tatcs.inclucJing t$o locations probablr sirnilar to thr Nlanis l'lastodon sitc on thc in \\ a-shin€{on(Cu-stalion et al. 1979. Xlehlinger OlvrnpicPeninsula.30 Lm to the soulhnesL.Pol- l9ti{3. \'lehlingcr and Foit 1990). This hvpothc- lcrr and plant rnacrofossilsshow that thc I'lanis iuca sis is clouclcd b_"-thc discovcrl oi a iiagmcnt ol remaineclrelati!el_y treele-.s durin€i the peliod frorn humerus lrorn .r ]arge bear in Nliddle S'istonsin 12.000 Lo LI.0O0 BP. \|hile orher $esLern til1 aL S'oodbridge. Orrtario.Churt her arrcll\'|or- S'ashington sites containcd abundant urniftrs g.ur (1976) identified the specimen :rs {,i. dr.lor. (Hues-ser1977. Petersonet a1.198ll). This diller'- and cslirnati'thcagc at 50.000-.10.000 BP. Tf cncc l rs causcrl bv thc rain shadov of thc Ol_vm theseinterpretations ale correct. migrationoi the pic Peninsula.as rell as adclitiorralariclitl causr-'cl

l)lcistoccncBrorvn Bcar' I I I bv the more -.oLrtherivpalh of $ inlcr-stor.rn tracks lsl:rndm:* hale beerrtonporaril_"_ crpo-.ed. nlter- irr rcsponsclo thc ploximitv ol the retreetingite natclv.thc 0.5 km channelal Decel)tionPass pfo- sheet. vides eirsvaccess Lo Lhe rnairrland bv r,;al of l'idalgo 'lhc ,17 cm thick lossiliferousholizon at the Islald. A final possibililv is that Ll. arctos ctossed Mani-.tr'lastodon site rielclsradiocarbon clatcs that frorrrthc mainlanclvia ice blocls cahecl fron thr: rangelrorrr 12.300 + 310ro11.000 + t5OBP. reherting ice -.heeL,whi.h was locatcd approxi pre-"urnecllo be contenrporancouslith thc Whid- rnatc\' 100 knr to the north. near the preserrtf..S.- bcv Tslanclbear fossil.ln both locations.the Latc (l:rnada borcler'. Pleistotene topographv consistcclof gravel floored Crizzliessrrrvilr-'cl thc rigors of the last glacia- porrdsdclclopecl on sta€trantice ten-ain.Pollc'n tion.anrl ilourished in the milclclimaLe (rf lhc IIolo- at the frlanis site indicaLe-cthe shallow ponds r,;crc cene. \\'itll :rn estimaledpolrulation of 50"000 in lith in aquatir'|1ants.$hilr: acljaccntuplancls con the lo$er 4,li staLrs\\'hcn T.cl,;is ancl (llark visited taintd alrunclantshrubs aDtl herbaceousplanls. in 1805. lluc to combinedimpact-c of unreslficlcd 'l he plant assemblageappetrs lo rcprscnt rln earlv hunting anrl ertcnsir'chabitat loss, bv the I930s stagcol successionas shrLrbr illorrs(Solir). nran- {i ol.tor $'asextinct in the northuesterrrLI.S.. cr- zanita \Arlostophtkr-r) and othcr sun-loling sle cepLfor scanl local population-.in isol:rteclmoun- cies colonizedthe ba-ren llelds of Lilllcft bv thc tain areasnlonpl the border xith British Columbja. Ietrellling i.re. In northxest ['ashington. ] 0- 15 grizzliesare esti- 'fhese enrironrnenLalrnnditiorrs loLrltl har.c maLc(llo ha\c sun'ivcd in the Nolth (la-scacles.*ith pror,idecl arr ecological rnagnct fol ncr,rlv nrrived pcrhaps 35 more in the Selkirk \'lounlains of nonh- blot'n bears. but the rnanner in vihicfi thev rcachul ern Srashinglon and acljacent ld:rho (O'Cala $.hidbev Tslandrcmains cni€lnatic. -{t presenL.the 1992J. Ol the coa-stalpopulations that dr:sccncled islnnd is separatedfrorn the nrainlandb_r r 3 km ir,,rrtlr, ShiJl,r T-l.rn,lb.,rr,rnJ it- ,,,rrt,:nr- uirlc channcl to the east and the Olvmpic Penin- polaries. none remains alir,elodav, sLrhlics 5 km to lhe souLh$esL.dist:rn(,es lhat arc' (Hea within Lhes*inrning abilitvof rrodcrn bears Acknowledgements ton ard Crtdv 1993). l'hc a€ieof the specimen doe-.not allot sullicienLtime fol sigrrificantplatc V e thank f)r. (i. Xl. Senger.S S U Riologr llcpart- leclonic adiritv to havc occurred. but fluctLrating menl. Iol his assistanccin identiliing and inter- see ler'elsnrav hare ternporarilvrcduccd channel preting specimensirorn Sihidbcv lsland. and l. tidths during thc LatcPleiskrene. X'lultiplestrancl F. l\lcCallum.D.D.S.. ibr he$ in ev:rluatingrJen- lines on the mainland easLol S'hirlbcl Tslrnclarc tai chu:rcteristics.Dr. C. R. Ilarington. Curator ll5 67 m aborr: prcscntsca le1'el.and date liom olZooLrg"-at thc Canadian\'luseurn of\atule (Ot- alrorrtl2.000 to I 1.500 tst'](E:rsterbrool 1992). tona) pror.idcd rnuv helplirl slrggesti.,ns for irn- lf this pattern ol rapid emergercc occurrcd at an proving the oliginal manuscript. tith later erralic ftrlc comprrccl to custatic se:rlelel rise. por- conlribulionsliom rcviclers f). L. Petersonand tions ol the shallowcoasl arca fringing sJhidbev Xl. H. Beeson.

LiteratureCited Gruhlm. R. SJ. 1991. Voirbilit\ !r rb. siz. ol \,)frli {miifi crn l) r.[ L.dfs (airb an..i.a,!.9 \rirh thf drs{i.il! Carlstad.(1. A. 191)2.L!t(. Pl,.i"r,!r.nr.,l.gh, !rior hisroN el rn,n,,1 ! lossil hla, k b.rr irom Dill \effCare. Virgnri!. Itnir l'arrriilge. ( errr!l \\ li,lb( \ ls !,,(1.Uaslrinsrof. llli rii -st!l{' \1,,s,,,n, Sci. Pup. 2Jr 23i3 2U0. (hildr\'. (;rizzh ll.S. thesi:. tri.stear \\ ashir;1ton t ni\ff!irr. &rl J. E. 1968. lljaN lio,n Qsrcrr \odh lmerica. Aner. l\{nllarrl \at!.alisl ;9: 2:1.72;0. (1. (lhLrih|r. (1. Cusrafs,,n. E.- CillrN.1).. arrl Drughcrtr. R.Il. 1979. The S. anl \{orsar. \. \ . li)i6. \ sri,,l! berr from XlarrisMasrodor sitr: call\ mlf on rhf 0llmpic IJeDnl th. r,i,l(ll. $ is(lfsin ol tr oorlbrlrlge.Ontartu. Car. mla. (lar. J!1,r. Aftilfu,r,l. ll: lll; l6l. 1,,!r. llarLliS,i. ll]: l].1131;. Harnrgtor. C. R.. l9;;. l'1(iist(x1.n,V,,slor.n {.\jDroil honr l-iirt1b,ool. I). J. l.]92. {{l\rn,r. rn,l rt(et ol corililleror ,\lberta and British Cohurhir. (l!r. Jo!r. llaftl, S( i. 2ll: (i!,srat,lii. ice dreeG ;i \\ !sliirst,tr,. I .S. {. fthrslque 903-919. el Qudternaire 16: ;1 68. 19B9. Pl, i\r,!1.n(.!.nebfrte locdliriesin lhe \ u- 1]lllruk. D. P. l9;:J. A Ile\n\\ of lrxsil arl H,!r.nr ll,.rrs loD: ;r (l!rr.r. 1..li.- 1. I). Ilunilton.and J. P. (iallo ofthe Ol.l S o d. l'ul,1i,aLi,. \ aJ,U,.r \1in. Geologirh \a\..,1s.- l.ar.Cfn,,oi, Ilistorrol theIrternr Basnrs lnstn,,,,l Biiks t.,rircrsit,.itt lrtt. \.rrrr !fits. 1,|.Ah"ka akl Lli. \ ukon. 1..S.GeoL. Surr. (:n.ulal 1026: ()3-98.

I 12 l'lustoc and Carlstad Hcaton.T. H. andGrarlr. l'. 19913.Fossil srizzlr beor 1a-irslr N'tundr, K. H. r). anrl Froot. D. H. t97it. Bactgnurl f0r a..ro-()lrornl,.in.i,,,1 S'ales Islard. \ !ska.oller neir managnrg grizz \ berfs ni the ratnrnr prrks ofCareda. i,rsighrsiDro aDir!!l distx.,.s.l. inlerspe.ill..orrt)..tirion. Can. trildliki S(.1. R.fr. Seies 22:;-:t1. (ilrurr and aseol deglaciaron. Researh h rh, P.isro ]l.N{r,mb. R. C. aDd R.txJ,nins. C. {. 1970. o,r'!*:nce oi gt) .ene l0: 100. mannnh fossilsin rhe linxilid Beds. ioLrrbrr.nrral Ilo$$rr. C. J. l9;;. Qudtcrnanpah rolosy- ol rhr I,ucfic -.1ope \l'aslrirgLof. \onh{est S.;- l:r:16 lil. o{ tr'ashnrgt,,r.Quare.nrr! R'sfari.h 8r 282 306. Noe \lsaard. \. r 983. { nei! fiDd of Bfusn Belr {aisL\ dr.r,.\) Hopklns.D. \1. l96i. lhr Cenozoichiston ,)l llu,x,sh ! s\!- lron Star Crrr unil other finrls ir the Lare Clarial anrl rhesis.t, I). NrHoluns. --d., Ih,. U.rifg I-!nd Brids,i- Ibst Glacir ol Britah and l)r:r,r,ark.Jour. An hq,l. Sranfordt n;rcrsilrPresr: l;l-:[8.1. -qr:i.l0:31 I 325. K,rflln. B. 1960.A skul of thegrizzl blar(li otro;L.r frorr O Cor. C. 1992. { srizzlr'spla.e. -qi,irrr7?i5):5; 61. B081. Pit ltl, Ilanrho la Ilree: t.os Ansrli.s O,rnrr X'luseum Pelersen.K. L.. U.l,.i.ser. I'}.J.. Jr.. !,,d cusrafsoD.C. ll. Conrrib Sci :J9:l-;. li)ii3 Late Cla|ial rcgehrion and .linatc er rhe ll. r . l9(,3. \otes on srrrc Plcisrocene mamrialrrigra ftstn,lon site.0l\nui( tl ninsul,. tr'aihjrgr,n. Qi,arer ti.,hs iron, rhe Pilaeer.ri. r,) tlic \err.ti.. Eiszairalt.r nar\ RfseNrch 20:21;-231. u. a;ag,i,Nrrrr1,1:!16 103. Perer-.o!.H. 1.. l !165.\ $el1prrscncrl gril,lr bear sk,,ll nao . 196B. Pl(.isloceneN'lannn:ils ol lirrope. {ldnre tl,b- rercrl lrcm r Late Clacial rl,lnsir near Lakr Sinr or:. Ishn,s Conrpun!. ahr.ago. Ontario. NorLue20lJ: 1233 123,1. 19i6. Thr Care Bear -qLorv.CoLrunbia tlnirtslLr Pielou, E. C. I99 L. After th. l(1. Agc. t,nirersir\ ,,1(lh (!so I'hiii. \c$ Yofk. K,rrtin. B. and And|rson. L. l9;1. Asso,iarnn ol Lirrs ./i1or Renlberger.J. V. Ilrroskr..{. D.- arrl,sp.Dcer.Ir. 1981- (;(l und 1rullr.r sirzzs (I{ommalia: t.ris,l!.1 in rhe Lare ologl ard glaciologl ofa Pleirt(xr.n,'r,, Ilolocene l,xiss f'lcisroceneof \{,orrirrg. Rreroria 126:1-6_ $r.cession. B.rt,tr, Cdrnt!. tr ashirgt,)r. Archleolosi ixril. L\Dra!. H. 1.. I On the Holo, ,.n. hisrorl of ahL' in .usl ra ud Histtrical Sc^i(s Repot 10i139. Flasrern enr \\'r.hington. \ortli$.st .. S.i. 60:6;71. l\.-l n"o, Li,-,- | "n.,. \1-t,i Lr, , llehrnrgfi- P..1.. Jr. l98ll. \\.rrx,rs ol rncieDt tn!:r(drs. RaL,v.h. R. 1.. 1963. C.oi{.a|hnr Qriatnn nr sizc in \orth \at. Gographic l7l i1l:500 5011. Amcfifan broir! b|an. Livr-r aicar L _ r' inrlicared llehrnrse.. P..1.. Jr. and Foir. l.- [.. Jf. 1990. \ol.ann.!'h l,i condrlbrsal lergrh. Crn. Jour. 2,o1. ,rl:3J l;. dd ,s.l rhe CLons .a(h. at 116r Senatchee. \\'dslins -s.hLrltz.l-R. l93il A Lar. Q!ur.DraN Drarndrl a,,n! trom ton. \ar. (;..gf!phic Ites. 6(1):195 503. th tur ser:|s of\lcKttrn,l. (lalilornh. Carn.gi, ln*r. \lcrrirm. C. H. 191B. Rcrieiv ofthe grvzli anrl big brown ltars tr'ash. t}ub. ,lijl. IJrrt 1:llB-l(,1 + 17 plar.s. ol \orth.\n,rica. \orrh {Drer. ldu,u 1l:l 136. S,r,rr.$. F. 1967. A rnamrnothfron thr Toucher Beds rrer (;. \'lille.. S.. Jr. lfl99. A nor hrsil bear fron Ohio. Proc. \{ell, \lalla. \tr'ashingrona.bsth.ll. \odh$esr sci. Bi,)1.Soc. tr'aih. l3:53-56. ,11:6061.

Rece ecL28 llay 199ttr Acceptedfor pubLicntionla Decen$er1992tr

PleistocencBrol,n Bear I 13