Greg L. Schfoer,'Olyr.picNationa Park,600 E. ParkAvenLre Port Angees, Washington 98362 KurtJ. Jenkins,rNatona ParkServceCooperatve ParkStudes Unt Co egeof ForestryOregon State lJnrversty CorvalisOregon 97337 Bruce B. Moorhead,Oympc Natona Park 600 E ParkAvenue PortAngees Washngton98362
RooseveltElk Selectionof TemperateRain ForestSeral Stages in WesternWashington
Abstract tr'f srudied hlbitur selecrion br Rooserelt elk (CcrrLj.ldplM rdos.r.ltt) D a remperare rain fi'est ir the 1,,$er Que.ts Ri{ir ld!l,i! ol the vestern Ol-rrnpic Peninsula, N ashington from JuDe 1986 June 1987. ELL annual hone ranges in'hded prcdonrnranLh ur- loggecl forests protected rdthin Olrn4ic National Park ard logged, regeneratnrg lorests arljacent r, thc park. Rarlirnrlharl rlk selected rallev {loors during all se.sons e\.eplrinter, shen elk frequentl ustd an adjoiirins plar,aL, riO n abor: thr' lloor\,lulrr. In idnter. rodio collared elk seLected 6 I5 rea old clearcuts. rhich were arailable or rhe plateau. Elk sdrtcd natrrrc rlcrnluors lnrfsrs ol rh. lallf! floof durnrs sprnrs. sumrrer. and outLrnu. md generallv tliel selected old age Sitla spruce l,rests rlurnrg au- r,,m,! an,l sinr.r. lirng clcarcuts (1 5veufsodl ! even rged. regenerutingsr.nds i16 150ledsold) generallr rere aro d dutinga|lsrlasons-llalugtn|.nLI)ral|i(l.sthaIn]tainl)l]li]nr.{l stands. and olil rge c0nife stands'Silricu]nrra]a]ternatireslolrpi.ale!en'agedsuldmalragcnnrL.su[aslrrcrcn.agri nng. should also be considered for improring ard nailtaiiirg inrerspersid oi frragc anrl r ot.
Introduction forest succcssionalstagcs l'ould be uselul lbr long range planning ol loresthar,resting in thc Pacific Logging has altered habitaLsofRoose!elt elk Ccn&.\ North$,est. elephusr,.roseteki throughout the Pacfic Northwest. -qincethe Lurrrof lhe ccnlLrrv.mosl lot-clcvalion Preliou-qsLtrdies o[ habitat Lrseinclicated that ol.l-agefofesls ( ) li>0 years)within the range of old age coniferousforests. deciduorrs forests. and Rooseleltelk in Olegonand [.ashingtonhare been some young age-classesoflogged coni[er-ousforesls cllt lbr lrood producls and conler1ed to even-aged {r'e impo ant habitats lbr Rooselelt elk (v.irmer stands harvested in 60-90 _"_earclltting cvcles and dcCulcsta1983, Jcnkins and Starkel-I984, (Judav 1977. Nleslr* er ol. l98 l ). For cxampic, Schroer 19f17.Brunt eI al. l9B9). Thosc snrdics. lon' elevation Dougl:rs'fir' \It.setLdotsngante nz iesi iJ however. did not include a broad arrav of sur:c.es- fofesls grciltcr thdn 200 vcars olcl covcr about l0 sional ages of forcst r,hich pclmit an asscssment percenl ol their original dislribution (Franklin et of all stagesof forestsuccess;on. Thc purposc o{ a1. l91.tl. Franklirr and Spics 19821,). this stucl,v,therefore. ras to deLerrninehabitat usc Flxlensircconvcrsion ofolcl-agc forcsts to cvcn- patternsof RooselelLelk \!'hosehome rangesirr- ageclstands in shortLerm harvesl rol lrons ma\' cluded Lrnloggedforests protectecl within Olvrnpit: have sippificant long-term effects on elk popula- \ational Park (ONP) and a valiety of successional tions. Thcsc cffccts includc rcduccd ar.ailabilitl of foresLsresulting lrorn loggingadjoining thc park. preierred elk habitats and forage (Ilarris er nl. 1982. \{ itmer and deCalesta19llil. Schroerei a!. StudyArea 19BB): reduceclplant specie-srichness and diver- The sLudvuas rcnducrecl ahng l5 krn of the lor,;cr sitl'. particularlv in pole sized stands (I'ranklin and QueetsRirer Vallel in and adjacenLto LheQLrccls I'cchancc 1968. Alaback I9B2): and increased Corridor of Ol,vmpic National PalL. The corlidor distrrrbarrce caused bv roads (S'itmcr ancl is a strip of national park land (l 3 km l'ide) that de(lalestr 1985. Schroel I987). (ireater under- extendsalong both sides of the Quccts Rivcr for stancling of clk habitat pleferences relati!e to approximately 20 krn. liom the inlerior core of ONP rvcst to thc Inclian llesen'ation. 'Presentaddress: Resources \oih{esr. In... lll00 ll2th.{re Quinault nue \E. Ste 100. Be erue.tr,L 98004. The Queerscor.ridor l{as addcd ro ONI'in 1945 ']Presentaddress: Xrangell St. Elias \ational I'arL ard I\e- 1,,prnlFi I ril'arian','mmrrrriti.s rrr,l r,lj,'inirrg rrc.b sene. P.O. Bor 29. Clennaler.-{K 99s8[] on the broad floor of Queets River Vallcy.
NorthwestScience. Vol. 67, \o. 1. 1993 23 The principal landfonn in the study area is a lock. Vcgetation of thc valle;- floor also includes broad. glacial out*ash plain thal is prcdominanr 55 abandonedhomcstcad clearings that range ;n along the soulht'csteln edge ofthe Olvmpic lloun sizc liom 2 to I 6 ha and arc composed primarilv tain Range (Tabor 1975). The Queets River ha-scur of non-natir.egrasscs, into this plain and created a l-2.5 Lm rddc r.allev Vegelation o1 the acljoining glac;al plateau ;s fl,,ur.lhe rall.r Il,,ori- alnrrtt,0 m in .l-r.rrion characteristic ofthe I'sugo heteropl4lLa fott'st zone $ilh at least 4 ler.cls of alluvial terraces (Fonda ol $estern ['ashinpJton (Franklin and Dyrness 1974). Valle,v walls cxtend approximatel_v60 m 1973). t,nhan'ested old-age stands of westcm Iromrirhe r.ide of rh, r.rller11,rnr rrp ro rhehnrrrl hemlrck ( ) 150 vears old) cover approrimatclv glacial outlash terrace (platcau). A majo tv of the 1l percentoftheir originaldistriburion on the gla- valley floor ancl vallcv t'alls arc r,".ithinLhe ONP cial plateau. Thc remaining frrrcstshave bccn con- Queets corridor, whercas the platcaus are within verled h I 150 year-old, even-agcdlbresls bv S ashington l)epafimenl of Natura] Rcsources clear-cuttingand planLingof nurserysccdlings. or (DNR1 and Olyrnpic National Forest (ONF) lunds. bl t'ildfire. Onc to 15 rear-old clearcuts are domi- Climatcis maritimc tith mild. .\!ct$inters and natcd bv non-native grasses and forbs. as $'ett as cooi. dry summers.Annual rainiall averdgesap- shrubs, inclrrclinghucLleberrics and Rudar spp.. ploximatelv 300 cm, wirh thc majority falling be, and rcgenemtingconifers. The majoriqrof t:lear-cuts (Jctober trrecn and Mav. Snorvfall occurs infrc- r,".erepiantcd with Douglas-lir lPseu.dotsugarnen- quenLl)-and accumulalionrarely ercecds 30 cm. zierii) or testeln hcmlocL seedlings,titir somc :u.ea.s Abundant rainfall, moderalc temperaturcs.and conta;ning dense stands oi nalurall,v seeded wersF 'l'he r:oastal fog har'e crcated a tempcrnte rain forcst ern hemlock. per
24 Schroel. Jcnkins. and Nloorhead habitnt use could not be 20 June 1987). An adult female of the Strealer avoided. Vclietation lo held lvas not radio-collared until earlv summer; stdtisticallv determincd lbr calving season due (12) ol thercfofe.it n'asnot included in the summer data. the small sample size and largc nrrmbcr habitat tvpcs (Koehlcr and T,arntzl9B0) S'e relocated clk b-'-lirst obtaining a gcneral rrsing radio-tclcmetry arrd thcn by ap- location b,v Results proachingto confirm a visual sightingor aural lo- cation. Locations$'erc not rccorded if elk Icre Radio-r'ollarccl clk selccted tirc valle,vfloor during disturbed prior to detcction. Tr 7 percent of the all seasonscxcept 1{inler (Table I}. During Iin- rel()alions. r!e uscd remotc triangulation pr(x'e- ter, elk genelally used vallcv floor and plalcan dures to locatc clk from distanccsof 150-1.000 regionsin prup,,rtionlu .rraihbrlit.' derivcd liom field tt:sts m. Alerage location clrors During summer all radio-collaredelk seLected \!ere ll0 meters.All locations at lhose dislances maturc deciduousforests () 10 years old. Tablt to 100 m coordinates.l1 ,n.r".""erfl6d accnrding 1). Additionally. oollared elk tended to sclect Mercalor SYStcm.Datd the Unilersal Transversc coniferous/deciduous fbrcsts and olcl homcstead r'"n u.'d in u_'e'sint from r, rnntelriJn{:lllJliurr fields. although those trends t'erc significant ibr patlerns onlY ifthe error polY- \'egetation selcction only one of thrce collared elk All collared elk !!ilhin a single vcgetation class. Eion\ras completelv avoided l3I- 150 year-old conifcr stands on the val- Vegetation selcctic-rnplltterrrs 1t'credetermined lcv floor. and also I -5 and 16-150 ,vear-oldconi- by locating ctch elk once or tlrice dailY on ran- fer stancls on the plateau. Old-age ( ) I 50 ,vears dom davs and at random tilncs betltecn daltn and old) Sitka sprLrceforcsts on the vallev lloor t'cre dusk 3-5 davs per week.To help ensurethat loca- us(l in proportion to availabilitr-.but old-age Nesr tions from the sarne day \\'cre indcPendent, suc_ ern hemlock forests on the plateau *er-e avoided cessi!elocalions lrere sepdfatedbv a minimum ol during summer. A rninimumof thrcc h()urswas a sulli- threehouls. Each of the radio-collared elk conLinucd to se period for the clk Lochange activity or vcgc- cienl lect mature deciduous lorests ()]0 years olcl) class ill greater lhan 50 perccnt ol trtion throughout autumn (Tablc l). ddditionall,v, 3 o1 locationsuithin a day. consccutive ,1, elk selected coniler'/deciduous and old-trge lndiridual patterns o1 !egetation selt'ollon 1{cr-e sprucc forests on the valley-'floor. All scral stages eriaminedby comparingproportional usc of vege- available to clk on the platcau werc generallY tation classcs to proportional availabilities o{ vcge- avoidcd during autunn. tdtion classes $ithin anntral home rangcs ol During rtinter. all radio-collared clL selectecl indiviclual elk. Annual homc ranges rvere 6-15 I'car-old conifer stands on thc plateau (Table delineatedusing 100 percentc()ntoul intervals ol l). Tncreaseduse of 6-15 vcar-old stands cor- the harmonic mean Lrtilizat;ondistribution (Dixon rcspondcd nith dccreasecluse of malure dccidu- and Chaprnanl9{30) using the cornputelProgr;lm ous lbrests (> l0 ]ears olcl) and coniler/deciduous I\L:PAAI- (Sturvc and Bloharviak 1985) Vegeta- forests on the lallev 1loor. Two of four clk con- tion classes ircre mappcd t'ithin home r-angeson linued to selectold-agc spruce loresls on the val- 1:2,1,000 scale basc maps using scrial photo- ley floor during t'inter. Elk tended to aroid l-5 graphs. DNR and ONF timbcl rranagcrnent maps. and l6-150 vear-old conifelous slands on lhe val- ortilrphotoglaphs. and licld rerilication. Availabil- ley plateau, and they ElencrallyLrsed old agc $est ities of r,egetationclassts $'elc measurcd using a ern hcmlock forests in proporlion to availabilitl. digital planirreter. Duling spring.elk use of 6-l 5 year-oldlbrcsts We testcd null hvpothescsthat elk usedvcge- diminishedvhile use of maturc deciduoLrs{bresls tation classcsin prop{rrtion to a',ailability using Chi- () 10 old). coniler/dcciduousforests. and square tests and family conficlence intervals based -r''ears old homcstead {iclcls incrcased on the 'ialleY floor on Bonferroni Z-slatistics(Neu el al. 1974). A (Table 1). Elk used old-age spruce forests on the legetalion class t'as considercd selecled $'hcn !allcv Iloor in proportion to a"ailabilitv during proportional use cxceeded availabilitr (famil,vlevel spring. and avoided l-5 and l6-150 lcar-old P < 0.05). and wasconsidered aroided lihen usc coni{crs on thc was less than availability. Statisticaltests could not Plateau. evalu- be conductcd *'hen thete l'as no docurnented use Although sdection patterns could not be season. of an available habitat typc or topographic area' ated statisticallv cluring the shorL calving .vallevfloor although those t,vpes ol areas \!cre considered use of conifer/deciduous foresls on thc
Etk Habitat Sclectionin a Temperate Rain Forcst 25 1
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Suhr,,cr',Jerrl,irrs. .rnJ Moorhead and l6-30 vcar-oldconilcr standson the plateau frorn our dala. but uncloubtcdl,v the_vstem liom appearedto incrrascdLrring c:rlving (l able 1). Lse diflerenr,es in landscape pattcrns and fora€ie ol old homestead fieJclsancl olcl-agespruce fcrrcsts productivit,v in the 1$o phvsiographic zonrs. Prox- rr-'rnaincdhigh cluring cdling. r,".hereasuse of ma- imitv of old-agc Sitka spruce lbreststo preferred ture deciduous frrrests() l0 ,vcarsokl) clccrcrscd. deciduous ibrests in the,'allev floor nrav har.c cn- L sc of l'5 veu old. I6-L->0year-old. and olcl-age hanccd usc of those coniler stands in boltornlands. conifer stands on the piateau lenrained Llr dur- Additionallv. although we did not attempl to meas- ing calling. ure differenccs in ftrrage abundance among old- age fofestassociations in the QueetsVallev, cover Discussion of prelerred lorages ol elk differcd apprcciably among cJimaxlbrest associationsin the nearbv IIoh Vallcv bottoms and riparian areas :rlong the QLreets Vallev (Srnitband Ilcndcrson 1985). Other habi- Rirer *ere kev habitats sclcttccl by elk vear tat benefit-qallributed to olcl-age forcsts include: arouDd.Nlature deciduous foresls and crrnifcrous/ snon' interception and a-ssociateclenergetic bcncfits deciduousstands appeared to be importanl forag- during winter (Jones197,1. HaresLad el al. J982, ing.rr".r-',f "lk drrrirrgall ."a-"rr- e\, dl,t $intcr. Parkcr et al 1984. Zahn 1985); greater nLrtritional Tn gcncral, tlcciduous lbrests contain abundant for- clualitvcrfbrotsc appeet al.1990). and abun- age frrl elk in conrparison to conilerous ibrests of {H danl forage(Alaback 1982). Additionatresearctr similar ages(Franklin ancl Petienac l968). ['la- clcar\' is needed to determine influences of sitc lurc dcciduousfolests in our studv areacontained qualit_,_.forage prrxludion, dnd for'lrst stnlctural )9i> percent ground covcr (C. 1,. Schroer. Nat. characterislicson elk habitatattributcs dnd selec- I'ark Serv. unpubi. data) of imp{)rtantclk forages tion among old-ageforest associations. in the repfon(Leslie I9B2). Researcho[ othcr clk popuhtions in westem S'ashington and Oregon Managementlmplications and Vancouverlsland. B.C. also documentedelk --le, ti,,rr,'f 'l-, i,l'r,,rrl,r, -ts .rntlriplfiJn llrcri- (lon,'ersion of old-age conifcrorrs forcsts in manl (Brunt er al. I989. Coopel l9{}{t. Schrocr 1987. aleas of the Pacilic Northwest has created cxten- Jcnkinsand Starkcv1984. Witmer and deCalesta sile even-agcd stands less than 90 vears old that t 9lt3). are eroiJed l'! 'lk. Timh.r mJnr€r.mcnt proje.- Evcn-aged conifelous stands less than 150 tions indicate that by the vear 2030, more than rc s old rccri\.cd little use by elk. exceptduling 65 percent of the total area oI commercialforest nintcr. r'hcn 6- 15 vcar.old clearcuts were selected lands in western Washington and Oregon will be b1"all elk. Stands that u,elc 6-15 ycus old con betrveenthc agcs of 30 and 90 years old (Harris tained the most abundanl folage (780i ground I9B4), and olcl-ager:onifcrous forcsts l'ill cover rrrvcr) ofanr cven-agcd coni{crous standsless th:rn iess than l0 percent o{ their origirral arca (Frank- 150 vears old ((1.L. Schroer.Nat. Park Scr-r.un- lin et cl. l98l, Htrris 198,1). publ. data),r'hich mighLha,'e contlibutecl to thcir' The increasingpreralenr.e oI l6-90 ycar-old sclection.Conifelous stands approxirnatelv 10-25 cvcn-a€iedstands poses a significantproblem for vcars oltl in southeastAlaska also had the greetesl managcment oi some elk langes in the Pacific: abunrJancc of foragc cor.er of all -seralstages (Ala- North$'esl. Extcnsivc cven-aged conilerous lbrests back l9{32). The habitatbencfits of6-15 _vcar-old generallvdo not qrnlain lhe intcrspcrsionoi for- coniferousstands. ho$e!er. are shor't-livecl.Onlv agc and colel aleas recommenaled bv elk man- 10-15 percentof a 60-90 ycar cuttingcycle would agels (S'itmer et 01. l9B5). Wc recommend that include that age class. Succcssionofvoung conifer- comprehensivemanagement plans bc der.eloped ous stands to pole-sizeclancl mittLrrcstands rcduccs on a watershed or sub-watelshcd basis, includinpl foragerluantit,v. and typicall;-thosc slands pcr-sist state. plilate. and federal lands, kr cnsure long lor approrimatelv 75 vears (Bij0/o)of r 90-vear cut- tcrm ar.ailabfih ofprelerred ell habiratsr,irhin the ting cvdr-'. managedforcst. Thc majoritv of radio-collaled elk selecLedolcl- Forcst management in vallev lloor ancl ripar- age spnrce for-esLson lhe r.allc,vfloor during au ian areas should be crnductcd in a manner that tumn and winter. and generallyaroided olci-agt: prulc.l- rnd er'hance- pr.f"rr'.d !i e' lirliun l,".cstcrnhcmlock {orests on the plateau except dur- habitats. cspecially mature deciduous lorestswhich ing n'inter'.Reasons 1br the disclepan Elk Habitat Selectionin a Temperate Rain Forest 27 protected on cdtical autumn and n'inter rrnges. of trecs removcd (Hungcrford 1969, Bunncll and Whcrc old-agc forcsls cannot bc retained or re- Eastman 1976). Lneven-aged managemenland established,uneven-aged management r,".ith lcngh,v commcrci:rl thinning har.c bccn uscd cffcctir.clv on cutting rotations of over 150 vears ma_vbe required the Ohmpie I'cnin-rl.rto , r,.rtc lLr.rgingrrec- to recreaLea favorableintcrspersion of co!er an.l within the er.en-agedforest (M. Stoll. Ol,ympic Nat. lora€te. For., pers.comm.. 1991). Additionalsilr.icultural presciptions that maintain partial openings in the NI:rn,vadditional silvicultural practices such as ovclstorv cdnopv in cvcn-agcd stands and producc non-commcr-cialand commcrcial thinning, shcltcr- actreptablequantities of slash debris need Lo be i{ood culs and small patch-cuts (i.e., I 5 acres) dcvclopccland implcmentcd. have also been proposedas means of improving habitat interspcrsion and foragc procludion in Acknowledgments second-groi{h. even-aged lbrests (Bunnell and Eastman 1976. llall et ol. 1985. Witmer et dl. S'e are gratelLrlLo W. D. Edge. P. J. Happe. D- lq8ll. Pr''cnrnmercialthinrring. hnrrerer. i- typi- B. Houston. li. E. Starkev, C. \{ itmer, and H. NL callv not effective in improving for:rgc availability Zahn for rcr.ic*ing thc manuscript, and many for elk. ber:auseresulLing sla-"h depLhs oiten inhibit olhers $,ho have pror.ided assistance including: undclstorv productionand clk movcmcnts(Carri- J. Aho, C. Butlcr, NI. Butler, C. Degc. S. Ferrel. son and Smith 1974. Bunnell and Eastman 1976). K. l'ostcr. and N. Smith. Rcprcscntativcsof the Additionally, precommercial thinninpl oi vounpler V ashinefon State Deparlment ol Natural Resources stanclst_"_pically opcns ovcNtorr canop;cs and in- (D\R) and Olympic National Forest (ONF). indud- creases l;rage cover fbr onl,v relativel,v short ing J. Cederholm and A. Vaughn, also provided 'r.aluablc periods {i.e.. 5-10 ,ve:us)depencling on thc numbcr contr;butions. LiteratureCited Irranklin.J. Ii..and T- A. Spn:s.1gBil. Charr(rrrisLnis ol ol,l- gro$th Iln,shs fi. i,r.sts. ,l \c$ lor.sts 1.,,.a Chrns- Alabacl. I'. B. 19U2. Dvnanics ofunderston biornus nr Sitkr nrgWo d. Proc.ofthe 1983 Soc.Amer. ForestersNat. ip ce \restern hemlock forestsof southeastdlaska. Conr..I'ordand, OR. Pp. t0 16. EcoloF 63:193219.18. Carisor. C. A., and J. C. Smith. 197.1.Habitat of grazing Brunt, K.. lJ. Q. Becker.and J..{. Youds. l9il9. Vancourer annrals. In O. Cramer (ed.) Environ,nentalefferts o{ Island Rooserelt elk/intensire forestry inreractions: forestesiduesin the I'acifir \orthiresr. l..S.D.A. l or. PhaseI {1981 l!186) Job cornpl.report. Res. IIin. ol Serr.Cen. Tech. Rep. PNV 24. Fp. I 10. Enrir. & Forests.IVTFR 33, No.B 5 l. \-icroia,Brit HaIl, C. F., C. ltcComb, rurdS. Ruediger.198s. Sihicultural ish Cirlumbla. options./D E. R. Dro$n (e,1.)NI{nasement of wildlife llunnr'1.|.. anrll)- }]asLrnan.1976- tillitts ol hr.st m!n!ge- ard fr-.h hal,itats ir iorests ol sesterr Oregon enrl ntt pracfir,sor r i1d]ih,h thefrrsts of BritishColun trrshington. L.S.D.A. For. Ser.. Pub. \o. bia. 1nI,ro.. xYt ItJFIr0 trorld Congre*. Div. r. Pp. R6-tEWt.-192,t9B5.lb_ 291-304,_ 631 689. Happe. I'. J.. K. J. Jenkins.E. E. -qtarker'.and S. H. Shanor. Orrpt. K. C. 1988. SeasoDalrnolcmerts ard habitat use of 1990. \ulniional qualit_r'andtonnin oslringen.! o{ nigratorr Roosereltelk ni \'lount Rahier \ational Pak. brossein clcar 2A S,h','-r. J-',kins,an,l l\{,"'r|.,,1 J, bki,rs.K. J.. rn,l Fl.Ii. Srlrker. 1982. Socialorganizatn,r Srtrror:r.C. I.. C. tritmer, and E. E. -starkri. 1988. I.lliecls of R,!,s.\i,lr.lk in an olrlgroivth forest. J. N{annal. ol und use practices on ltooseleh elk s,nt,J .!ns.s. 63:331-3:11. W(isr,if,, Ur.l,inglon ond Oregon. 1n Nf. Zahn, J. Pi(K - 19u1. Hal)jtrt,,!.h) Roos.r.lr flk l. unnranaged arrl R. Johnsor l,rls.l Proc. trestern States !nd lirests of the Hoh \ialler'. \\ ashingLon. J. Wikll. \lan Prodices E1k \\orkshqr. wasl,. Dcpr. of Sildl. I,p. tsr. 113.()12616. 68 7.1. Jo,i.s. C. I97,1.. Innuence ol lbrest derelopmert or blar:1,-tail,d Smirh. B. C.. and J. A. Henderson. I985. Basclinf \fselarion drr:r rurge on vancourer tstand. 1D H. c. Bt!.k i,n.) suner ofthe Hoh and Dos.wlllils (lrainlses. Ohm Wil,llile lnd foresr m&rgemenr in dr Pafin. \,,.1h- pic National l'ark, trashingt(ri. l inal Ri,pr1. Ohmpic lest. Srhool ol For.. Oregon Statt'tlnir-, (iorrallis-Pp. \atioral Park. Port Ang.l(is. $ ashinslo.. I :t,)- 1.18. Srwc. lt. and C. Blohailial. 1985. trln ro(r,tr,r,u1,. prosrams Judar. (1. t'. 19i7. Location.composition and sbucture olold- lor rhe anahlsjs of animal locatnns. Qnsi,.vltio,, !n{l groiLth itr.sLs oiLli. O(.g,rr Colst Roge. Oregon Slate H.s.n.h C.nlo. \d. Zool. P[k. Smithsoniar lr*tiLrrtc. (hr (irnl. Unir.. allis. I'l-l). Th.sis. Tabor. Il. I). 1975. |o th,, seologr o{ Olvmpic \atnDal Koehler. K. J., aDd K. Larhtz. 1980. {n cnrirical invesris! I'ark. i'a.,. N.U. NuL.Parks and For. Assoc. tion ofFoodness of fil :talisti.s ior stlrs(i m,,ltinomi!ls. Sitner, C. X., ald D. S. ,l.CdksLa. l9iJ3. Habitat use br J. lnier. -qtat. Asir. ;5:336'3.11. Roosevelt elk in LheOn,go. Coasi Ringe. J. Wildl. }{aD Leslje. D. lL L9il2. Nutritnxial .(r,l,,s\ ol rlnirls in okJ groiih age. 17:933-939. forests in Olunplc \atnnll t'ar[. Vashingron. OresoD . 1985. Ell.(it ,)l lorcsL roads on habitat use b\ State Lnir.. (hrallis. l'h.l). Ihrsis. H^.,-,,, ,1,. \..t1 k :f, rrrt22t2;. Nlcslrx. E. C.. C. N'laser. anil J. Verrer. l9Bl. Olrl-grorvLh Sirmer, C. I., lt. trisdoh. E. P. Hrrshmrn, R. J. Ander tuf(isrs as aildlif.r habirar. \. .{mer. S ildl. l,lat- Hr:s. son. C. C:uer. It. P. Kunel, L D. Lrd,an. J. A. Roclrelle. (i,i1. 16:329-325. R. W. S(l,a.d. r lD. Smithe_!.198s. Deer ard Elk. Neu. C. tr .. C. It. Brers. and J- Nl. tt*k. l9?:r. I technique 1, E. H. ljr(,trn (,a1.) Nlanagement of rvildlife and fish for analrsis ofutilizatnra!ailahilit\ daL!. J. Wildl. Uln habitats in li)r.sts olststcrn Oreson ud WashiDgt,n. age.3il:5.11 515. LS.ll.A. fi,r.,q, r\.. Pub. No. R6 F&WL 192-1985. Plrker. K. L.. C. T. Robhins, ahd l. A. Hrtr cv. 198,[. Energl Itrlard. OR.t\,. 231-2s9. erpenditures for L,crnoLnn br dulc {lc(r und rlk. J. Zahn.H. I'1.lg8s. t,scoirh.mral co'.r br ek (Ccm,LreL4l,Lt Slldl. Nllanage. 1U:1 7'1-18B. oD a ir.st(Jn \lashinsron summer rage. Lniv. trash., Si hroer. G. L. 1987. Semonal distibutnn ard rnovcmcnLsol Seattle. I'h.ti. Thesis- nisflron Rooserelt elk in the Olympnr UouoLlins. \\ $h;nstor. of.son Sr^te Lnir., Conalis. OR. U.S. Thesis. Rer:eitedI February1992 At:ceptedfor publication 22trJuly 1992 Elk HabitatSelection in a T,.mpercteRain Fore.t 29