SuzanneM.Wilson and Andrew B.

Carey,lPacific Northwest Research on USDAForest Stat Servce 362593','AvenueSW O ymp a, Washngton985T 2

in SmallMammals OakWoodlands the Puget in Trough, Washington
We surveyedlhe 22 largcst sitcs dominaledb] Oregon white oaks on the Fofi Le\\'is Military Resen ation. Washingbn. to derefmine small-mammal community sruclurc and population abundances. Study areas{'ere in the Puget Trough physiographic province and westcm hcmlock !egetation zone. Most oak communities rlere ecotonal belwccn prairie and Douglas-fif fbrest. Small mammals wcrc sampled at each site using paired lines of live traps for four nighls, July and Augusr 1999. In ordef of decreasjng abundance. deer lnouse.vagrantshrew.Trolvbridge s shre\\',and crecpingvole were the nost abundantand lvidethe spreadspccics.Thc dusky shrew and $e southemred-backedvole were infrequentlycapturcdin oakecolonesbut wefe abundani in nearby sccond growth Douglas lir lbr€st. The relative influencesof prairie versus Douglas ijr fores! on oak ecotonesdetermined understoryplan! compositionand occurrences ofsmall mamm: species. The combinationof abundantv agrantsbrewsand few dusky shrewsin oak ecotones suggestthatroil lbod webs and organicmatter accumulationdifferedbetweenoakccolonesand Douglas-fir fbrest.

Oneof thegreatest concentrations Orcgonwhite of oak (Quercus gertj^and) within westem Washington is found on the 34,,100-ha Lewis MiliFoft tary RcseNation near Tacoma.There, oaks are usually ecotonalbetweenprairic and Douglas-fir (Pseudotsugo menzlesii)forest or betweenpmi(Stein 1990;Ryan and Carey des and wetlands l995a,b;ThysellandCarey2001). The abilityof Oregon white oak to c(tonize both dry and sea sonally wet sitesresultsin a variety of oak communities. Where oaks border wet sites,they occur with Oregonash(Fraxitruslatifolia), bigleaf maple (Acer macrophyllum), and Douglas-fir. Where oaks are ecotonalwith coniferoustbrest, quick-grcwingDouglas-tirdevelop underoaksand eventuallyexcludethem (Stein 1990).Oak stands that extendinto prairie developa sllvanna-likestruc ture and have few competingtree specles. Historicalevidence people suggests indigenous maintainedPugetTroughprairiesand oak woodlandsby buming (Nonon 1979.Kruckeberg1991. Agee 1996). Burning ceasedwith settlementof the Puget Trough by irnmigrants from the eastc m U n i t e d t a t e .S i n c e h e n . r b a n i z a r i oa g r i S . t u n. culture, and natural succassionhave destroyed many oak woodlands, Douglas-fir forests have
I Author to whom corespondence shouldbe addressed. Email:

replacedoaks in ecotones, and the extentofoaks hasbeenmuch reduced(Leighton 1918,Franklin andDymessl9ll8.Kruckeberg l99l,Agee 1996). Today.even whereconifer encroachment not has significantly reducedthe extentofoak ecotones, weedy exotics,especiallyScot's broom (Cltisir,s (c.g., perennial grasses ;cv2arlris)andsod-lbrming Agrostis capillaris), are a widespreadproblem (Toneyet al. 1998,Usseryand Krannitz1998). Ongoingurbanization, fire exclusion,and in v a r i o nb 1 e r o t i cs p e c i et.h r e a t e t h e r e m a i n i n g n o r l u n d p r a i r i eb i o t i cc o m m u n i t i ei' n w e . t e r n (RyanandCarey 1995b. Washington Thomasand Carey 1996).The expectationof additional loss of the oak-prairie-wetlandmosaic (Thysell and Carey 2001) suggests least two mammalsare at at fisk: the westerngray squirel (SclLrirs grl.reas) listcd by the Stateof Washingtonas Threatened anda Federal Species Concem(RyanandCarey of 1995a, Bayrakqi al. 2001)andthewestem pocket et (Thomom):s nra:ana), a FederalSpecies Eopher of Concem (Ryan and Carey 19950, Steinberg 1999).Concem about loss of biodiversity in the Puget Trough has prompted scientistsand land managersto seekmore infbrmation on all flora (Ryat andCarey andfaunafound in oak ecotones 19950).To date,thereis little infbrmatior on the disribution and abundanceof cryptozoic small mammals(mice,voles.shrews)in oak-dominated communities the PacificNorthwest. in


NorthuestScience, Vol. 75, No.4,2001
C 20ll b! dic Nodhs€r ScicnrifirAi$!i1rio.

e e d ..we estimated percentcovels ofoak andDouglas-firin the overstoryandgrass. (3) compare mammal among oak ecotoneswith different abundances (i. Here we rcpoltthe resultsofa sur"ey of small mammal communities in oak ecotoneson Fort e r L e w i s O u r o b j e c t ie \ \ c r e :{ I l d e t e r m i nm a m . such as prairie.Sec ond-growth Douglas-fir forest is the most widespreadcommunity type and providesrhe context in which more restdcted communities. . Scot s and the coverofunderstoryshrubs.wetland.8 ha with a sufficient density of oaks to have been inhabitedby the oak-dependent westerngray squirel in the recentpast(Ryanand sites Because most oak-dominated Carey 1995&).Eachtransect at 20-m inteNals and were used both as small mammaltrapJinesand as plot centersfbr habitat On assessment. at each station along the for transects.5 cm). commonly snowberry (.whole oats.Symphoricarpos (3) oak-conifer. salal. Methods Study Design and Habitat Analysis 22 We selected sitesfrom 573 sitesoccupiedby Oregon white oaks on Fo Lewis (sec Ryan and Carey 1995bor BayrakEiet al. Deparlment a o l A g r i c u l t u r e T.1). Within the broader 1990:65.dominated by oak of with understories various shrubs.andmolNLsses bait to Small Mammals in Oak Woodlands 343 . Seasons distinctwith are nualtemperatures most precipitationfalling as rain betweenOctober and May and summe$ aremild andrelatively dry (Pringle 1990). elevation generally Proximity ofthe Pacific Oceanand PugetSound ancreatesa maritime climate with mild average (l2oC)..1 cm x 6.J n dv e g e t a l i o n : . conthe were ecotonal. con\ u m p t i o n f i n \ e n e b r a l e s. \lmcture Jnd mal rpecie'uhundancei cofiununit) in oak-dominatedbiotic communities:(2) relate to mammal abundances the relative abundancas of oak and Douglas-fir in the overstory. u e assigned each site to one of four types: (1) oakdominatedby oak canopieswith grassy savanna. wetland.1)mixed deciduous.g. canopies ttlbus).9cm) andone smal)(5.and carnivores.9 cm x 22.and to grasses. and oak types exist (Franklinand Dymess1988)..S. lvlammal Sarnpling We placedtwo Sheman live traps.prairie. Area Study Fort Lewis lies south of Tacoma. We used as peanutbutter. broom in the understory. riparian.pamllel to the long axis of the oak-domi nated biotic community.Washingtonin the PugetTroughphysiogmphicprovinceandthe westem hemlock vegetationzone (Franklin and Dymess 1988). Soils are derived from glacial till and glacial outwash(Pringle 1990).and Scot'sbroom cover in the understory to the nearestlclc. bigleaf large(7.e.consumpWoltr 1996) tion.we determined landscape text of each site with Fort Lewis GeographicIn and conformation Systemmaps and databases b l i r m e dl h e s e f i t e v i s i t . a total of 60 traps/site(useof trade and firm namesis for information only and does not imply endo$ement by the U.and aerating and other seeds by and enrichingthe soil seed-bed burrowing and defecating in the leaf litter below oaks (Stein small )andscape.reptiles. At each site we placed two transects'+0 m apart. 2001 fbr maps).We sampled10 plots per site. o .Gaubheria shallon): and (. Selectedsiteswere either the largestarcasoccupied more or lesscontinuouslyby oaks (8-44 ha) or were )4. shmbs. lO-m-radiusplots centeredat evthe ery third station.Topographicrelief is gentle and is below 160m (Pringle1990). andburyingacoms(Ashby 1967. conitcontexts landscape erousforest): and (4) comparemammal communities in oak ecotonesto those in second-growth Douglas-fir fbrests. we adjustedtransect orientationto lit within the areagenerallyoccuspaced had 15 stations pied by oaks. mammalsresidentin oak ecotonesalso scrve as prey lbr raptors.D u r i n gt h e r i s i l .significant representation oak of and Douglas-fir in the canopy with understones of forest shrubs(e. girdling and consumptionof the cambium layer of young oaks and encroaching conifers (Kruckeberg 1991): chewingon twigsandbranches containinggalls (Stein 1990).+cm x 16. (McPherson1997). understories.removal.oak canopywith additypically Oregonashor tional deciduousspecies.6 cm x 8.h e u s eo [ l u o l r a p s t a s t a l i o n t trap interference among speciesand decreases trap availability in the eventoftlap disincreases turbance (Calhoun and Casby 1958). for a total of 220 plots.(2) oak-shrub.The roles small marnmalsplay in oak ecosystems include disseminationof spores of ectomycorhizal fungi to the roots of oaks.

The deer mouse (Perrlmy.u. WilsonandCarey 2000.r'ival.p = 0.15 individual mammals of l2 species (Table 1). 61. inl. Carey and Wilson 200l).m o f l e m r a m i n a l i o n .1.Shon duratiol] trappingsessions reducedthe likelihood of immigrationtion populations outsidethe study area(Myllymaki et al.79. and (3) our sarnplingdiffercd in intensity totuleffon(Hrlnsson and 1975.).We thenrankedrelativeabun ( dances f-6) on a log scale(2. we comparedthe structure of small mtrmmalcomrnunities found in oak communities those to foundin thinned andunthinned 60-70-year-old second-growth Douglas-lirforesrs on Fort Lewis by Wilson and Carey (2000). and reprocluctive statusfor each animal and noted all spmng traps. 1.02). Our procedure was basedon the designofthe NorthAmerican Srnall Mrrrunrl Cen. Oakscomposed grcaterponion of canopycoyer a in ecotones thim did Douglas-tir (Table2).01) .hut u irhr 25. e StatlsticalAnalyses Wc reporttotal captures (includesrecaptures) and the total numberofindividuals capturedtbr each small mammal species. Bail wll.shmb. Howevet we haveonly oneyearofdata from oak communities and we have not demonstrated oak that r n a m m r c o m m u n i t j el \ r ea \ r l a b l ei n . 357o).We re corded species. rodentswere markedwith a uniquelynumbered Monel eaftag(NationalBand andTag Company)ivirtually all insectivores died duringlive trappingandwerereturned ourlaboto r u t o q f o rp o .improverlap efficiency(Stickel 19.. Polyester battingwas placedin each trap for bedding nraterial.rcrs mqni..02)and grass coverwas negatively corre(r latedwith shrubs = -0.Douglas-fircoverwasnegatively correlated with coverofoak (r= -0. r n d l o l t e d a m m i l l c r f t u r e sl \ l J t i o n \ s p m a as an aid to interpretingrcsults. Kruskal-Wallis H tests were used to comparevegetativecovers arnong oak community typesand Mann-Whitney U tests werc usedto determinewhich typesdifferedfrom one another We prepareda cover type map of e a c h i l c .Microtu. Results We caught . Preston 198l.Prior to release. Other species were relatively low io abundance and frequencyof occurrence. 16. 32. CareyandJohnson 1995)to indicatethe relatiyeimponance ofeach species within theoak andDouglas-firforestcommunities. t . was a measure ofbiotic integritythat is robustin space andtime in coniferforests across PacificNorththe west: their resultswere suppofiedby subsequent studies coniferlorests in (e. Caley and Harrirgton 2001. (2) probabilities of capturemay varl between dillerent environmenlsand within and 344 Wilson and Carcy among years. The upper limit of oak andDouglas-fucoverwassjmilar (ca.r) t were the most abundantand widespreadmam mals in oak communities: each was caught on 96q. although sitelackedDouglas-firAlmost one one-half (457.) the siteswere in the oak-conifer cornmunity type.P < 0. P = defincdasoctavescalerelative abundances basedon ICpUE. on of respectively.49. We exploredreas lationshipsbetweenhabitat elementsand abundances mammal species of with Spearman's rank colrelations. forest.Trowbddge's shrew (Sorer trowbridgii) and the creeping \o\e (.Sites were continuously trapped1br four consecutive days during July andAugust 1999(240 trap nighrs/sire). Douglas-fir cover was negatively correlated with grass0. Thus.= -0.18a).\ oregoal)were captured 777oand 68olo sites. age. 8. weight. We calculated relativeabundance thc ofeach species as its ICPUE as a percentage rhe total ICPUE of for the community.g.. used numbersof inWe dividuals capturedper unit elIorl (ICPUE) as estimates ofspecies abundances: effoft wascorrected for closed traps (Nelson and Clarke 1973) and standardized l00 trap nights. or prairie + fbrest.Kruskal-WallisH testswere usedto compare mammal abundanceamong fbur oak community types and among three landscape contexts (prairie. and Scot'sbroomcover(Table2).snot expectedto alter unduly the nolmal patternsot movcnrentby the small mammals (Stickel19. t r u c t u r e l l as conifer-nammal communities. ulatus) and\ agran shrew(Sorcr lag rar.rel.. Direct comparison of small mammal abundances bcLween oar and Douglas-fir communities was not possible (l) because the communities were sampledin different years and there is substantial annual variation in population densitiesof snall mammals. for all corrparisons. Among oak cornmunitytypesthere were significant differencesin Douglas-fir.51.ein surn pling effon (Calhounand Casby 1958).grass. trapswcrechecked fbranimalseachmoming.18a) added and carrotsfor moisture to incrcaseanimll sur.of the sites. ButCarey and Johnson(1995) demonsrrated how small I]rammalcommunitystructure. 197i ).N=22).

03). Mcans (SE) and mean ranks of percentco!ers for Oregon whjte oak (Qucr&\ gat'-dna).The creeping vole was also numer primafily ousat this site(n =10) but wastrapped with high coverofgrass.051 1 2( 1 .. mean ranks are wjth matching superscripts nol statislically significantly difterent Oak-shrub Habitat elemenl Mired O.2) 101' 3l (. \Yashinglor.1) 17t) 31 (3.10 0.001 <0. 0 t' .r-)01 i (sE) did of Abundances small manlmalspecies not ditl'er significantly amongoak types (Table3) or P landscapecontexts (a11 > 0.Only oneothcr at stations oak community had many creepingvoles.r) '70 9 (3.5) 129' 2 0( 1 . 6 ) i (sE) Crass x (SE) Shrub i (sE) 16" :15 (4.08 2l 21 11 15 E l 5 l 2 2 I I Emine Nofthem nying squirrel T0$nsend s chipmLrnk Southcm rcd-backed\olc Toral ) 581 (P.0.P = 0. whereas we Un313 deernice in othercommunities.10 0. 1 ) lt1' 38 (6./rrrgd TABLE 2.9) 109' 31 (2. Washingron.0.and 4 odk savanna(Savanna)in the Pugel Trough.001 8(1. s u m r r e f1 9 9 9 . r'/ (1.3) 80.59 0.3) 83. Spccies Dcermouse Tro$bridge's shrew Cfeeping vole Dusk] shrc$ Pacificjurnping mouse Totalcaplurcs lndividuals 2'.?.g.opdr"irli).46.t0 0.2) |]4" 5 (2. Trowbridge's shrcw was negativelycorrelatedwith grasscover (r = -0.12 2. and significanceol differencesamong types (Kruskalwallis P)l in rows.02 9.3) I 10! 9 (3.1 ll 5 6 2 2 I r5l 136 78 46 tl 5 5 5 2 2 I l '1'15 3.001 <0.0) <0.08).t 0. i P = U .r. Small Mamnals in Oak Woodlands 345 . caught derstoryvegetationat this site was lush. S a| mammal spccies.0) 118- 20 (3.gras!. l0 oak-conifer..3) 121" l3 (5.8) l09s l3 ( 15(2.1 0. 5 ) 0.5 mixed deciduous(Mixed).1 0.9) 100.sunmer 1999.indilidual s caughtper loli trap nighls (ICPLJEI TABLE 1.number of ind ividualscaplured.8) 114" 7 (3.5) 126b 18 (. containing severallife forms from procunlbentvines to tall shrubs.e. u h i \ h u a .ierlt).rk conifef All types i (sE) Douglas-fil 23 (2.Deer micewere negatively corelatedwith coverofScot's broom(r = -0. ryDcs:3 oak-shrub.e) 125i 24 (3.toralcaplures. in onc oak community we trapped 24 deer mice. Douglas-fir Scots'broon (C)rir&r 5..r) 79i 38 (3. jn the Pugc! Trough. and nalivc shrubs in 22 oak-dominatedbiotic coDnunitic! of'1 .There.7) <0.t"rl. p r e \ e n l n a l l b u t o n es i t e that Maps of capturesand cover types suggested abundance species local cover typesinfluenced at individual ffap stations.78 1. 2 3( 5 .76 138 78 5.02 0. and number of Oregon $hirc oak sires in which they werc captured (N = 22).5) 1308 (2.23 0.9.

5 2.).1 1.8 2. (2:l-48%). 5 LomntuntrY rvpe" Oregon whitc oak Thinned Douglas fir Unthinned Douglas fir Small nammal species Vagrant shrew Tro$. sunmer1999.rr gdr. Wilson and Carey2000). o l e Southemrcd backed vole Shrervmolc 5 1 l 4 t 5 1 1 1 3 3 2 5 2 1 4 3 'Differcnces in rank are considcred ecologicalty significant (Carey and Harringbn 2001).08 0.otr average) andwe did not find Keen'smousein the otk communities.5 0.43 0.c Sa!anna Specics D e e rm o u s e \hgran! shew Trowbridge's shrew Crccping \ole Duskv \'ole Pacilic. Creeping voles were crpturedon15 rlong lhc gr \s) prairieedge.6 2.).only threespecies each Rankcd relative abundanceof small rnammalsfrequentlv capruredin Oregon white o. influence in understory processes indicatedby the stlong as negadvecorrelationbetweengrasses and shrubs TABLE'l rn oakecotones. Negarive corelationsbetweenDouglas-fir andoak andbetween Douglas-1irand grasson our sites indicate successlonal tensions amongDouglas-firforests.56 0.and prairies. Smallmammalcommunitystruc turc is inJluenced unde6toryprocessesbut by more local conditionsalso detemine numbersof individualscaught(seealsoSoutheml965). washington.6 o.5 2.6-t2C.Canopies. ThysellandCarey2001).3 (.9 3.l ( I 2-21V.1 0.jumping mouse Shrcw mole Southemred-backcdvole Total ICPUE 5. turn. RaDksare basedon ocrale scalepcrcentages of total individuals qrpturedr 6 (>. Kmskalwallis terr p vatues statisricalti and for insisnifi cantdifferences among communi!y oak types.Forexample.*(eueftus garrrmd) biotic communities in 1999 and in thinned and un$inned Douglas-fir (Preudottugd men.8 2. of caDrures).2 13-6%).TABLE 3.1 0.'7 0.ietii) communiriesin 1992-199. oak communities.06 0.2 0. Oak conrnuntr tr r. Washington. Canopy composition lLndstructurc in oak ecotonesare govemed by adjacentcommunities(landscape context)andby intemal competition.include Keen's mouse (Peronlscus keeni')in low numbers(<17oof total individuals in the communities.bridgc'sshrerv Crccping vole D u s k y\ . 346 Wilson andCarey . however.8 3.1 (from \1'i]sonand Carey 2000) in the hge! Trough.ryrnd) communilics.and I (<37. Discussion Douglas-fir encroachment into oak communities confibutes to the net lossofincreasinglyrareoak communities thePugetTrough in (RyanandCarey 1995b.1 LI 0. Second-growth Douglas-fir communities.5 r. slnall mamnal abundance (individurlscaughr/100 nighrsr trap ICpuE) in oregonwhiteoak (o&"r. l\rlamma Community Structure Small mammal speciesrichnessin oak communities was similar to richnessin second-growth Douglas fir fbrestsin the PugetTrough (Wilson and Carey 2000). The distribution of abundance among specres moreevenin thinnedDouglas-firforest was than in oak communities (Table 4.08 ll tl onetrap line ran alonga prairieedgeand theother along a shrubby wetland. .1E%). Puget Trough.

ith ground cover consistingof short grassesmay be the most attmctive habitats tbr voles.Creeping locally with grassyareas voles were associated adjacentto heavy shrub cover Some sites with ho\ grass)underslories.+). Carey and Wilson 2001.The southernred-backed achievedhigh ranks in Douglas-flr communities andthe Oregoncreepingvole achievedhighranks in oak communities(Table4). In this rcgard. thinned Douglas-fir forest. and Verts 1998).+80). and3 species accounted tbr <36lo the individual mammalscapturedin of oak corlmunities (Table 4). foundthatdusky We vagrant shrews shrewsavoidedoak habitat.of specialnotei s the negative correlation we observedbetween Scot's broom and the deer mouse. important in Douglas fir forest communities. whereasin tlinned for Douglas-fiq four speciesaccounted >127cof i d c J p l u r ea n Ji n u n t h i n n eD o u g l a s . especially during the (Pe)es Banett and ofthe growing season early pa. was not captured in oak sain vannaandwasin low abundance otheroaktypes (Table3).ft (1970)andCareyandJohnson 1996). were rarely captured in oak ecotones(Tables1 and . oak-shmb. Differencesin soil organic-layeraccu mulations have been shown to influence the local distribution and inte$persion of dusky and (Hawcs1977).a n d . wilh rankunderslotl rbundanl siles at to possiblyasa response fruit andseedabundance H . Carey and Johnson 1995). Although found in Douglaslir fbrest(West1991. Cashwiler voleslimited by herbaceous (1995)foundcreeping vegetation. hrd [eu Lrecping voles.decidumore strongly assocjated (Newman1976. (C ey andJohnson 1995.important in oak ecotones. The effectsof Scot'sbloom on native prairie plantshasbeensevere(Usseryand Krannitz 1998)but its effect on prairie and oakwildlife is little known. Deernice.local vegetationgradientscan be panicularly sharp (Thysell and Carey 2001) Local conditions appearedto influence which namrnals were taken at a trap station. creeping of hadlow numbers smallmamOak savannas high shrubcover in mixed decidumals whereas.The dusky shrew.Thus.and that the vagrant shrew was markedly more important in oak communi3 ties than in Douglas-firforest(Tables and'1). common in westcm Washingtonconifer forests Wilsonand Lee 1995. may though edible (Car:raway not provide complete nourishmentor sufficient cover fbr creeping voles. 1 T a b l e . Halvo$on 1982) were the most abundantsmall mammals in oak ecotonesand were markedly \eFelrlion. associated a lmpllcations nd Llmliations Without managementintervention. in Trowbridge'sshrew. Harrington 2001). important in oak ecotonesbut the dusky shrew vole (Sore-\ monticolus)^nd southemred-backed (Clethrionomysgapperi). ous.which may signal a general decline in habitat quality for seed-and liuit-dependentspeciesin oak and prairie areasinvaded by Scot's btuom.especiallysiteswith grassyunderstories.f nro r p e c i e s s accountedfor <3% of captures. Carey and Carey 2000.e\er. l : V u n o m el g E 2 ) M i c e m r l e r e n 2 at have been underestimated thesesitesbecause home range size can be smaller in habitatswith more food.the dominantspecies each relatively remained r Douglas-fi forestcomrnunity. S p e c i e sL o c a lA b u n d a n c e s betweentwo are oak ecotones squeezed Because ditfering habitats.Grasses. the vagranl shrew is with meadow. Burning may reducevole populationsin the short al term (Gashwiler1970. providesprotectionfrom harshweatherand predation alsomay haveoontributedto the high density of mice. ofthe vagnnt shrewln oak coln The prominence munities is consistcntwith its ecological distribution in otherregions(VensandCanaway 1998).Terfy 1981).Wilson and Carey (2000) tbund numerous creeping voles where understoryherbs in and shrubswere abundant thinnedDouglas fir of stands. making individuals lessproneto capDensevegetativecover that ture (Stickel untbinnedDouglas-fir forest (Table4) The vagrant shre$'. which andcreeping increase with increasing production of acorns (Wolff 1996) and conifer seed(Gashwiler 1979.eachaccounted for>127. Canaway and Vens (1985) suggestedthat young coniler lbrests u. mamrnal community structurediffered markedly between oak and Douglas-fir communities(Table4). ranked last in Douglas-fir forests (Table 4). including two sites that had been burned. Douglas-tlr tbrests are poised to overwhelm a significant Small Manrmalsin Oak Woodlands 3'+7 . Hawes vegetation ous.and low abundances creepingvoles in unthinnedforestwith mossyforestfloors. The in deer mouse decreased ranked relative abunto dancefrom oak ecotones. and thinned Douglas-fir forest with greater numcommunities vas associated bers of small mammals.Fala 1975).andriparirur vole I 977. pafiicularly deer mice 2 voles(Tables 4).

S. Mittutus arego . A 5 Clure). of small mam als. H. burningprairie(Agee1996.A.M. a n dC .. J. 2001. Rodcnt occurrerce. S. l C : r e t . S.sand rhc slxe\r ntole in habi.Thysell and Carey 2001). Journa0f Zoology151:189 13. R. dominant the posrtion ofthe vagrant shrewindicated groundthat level processes oak ecotones in were distinctfiom thosein adjacentDouglas-fir forest. Ecology 63:413. 1995. H a r i n g r o n .cradicnrs rn rnc composition.s. Lrni|ersit) of Washingron pr€ss. Kruckebcry. M a n r n a l i a nS p c c i e s3 l : t 6 . and C. M. and Mi( /rt!r zrgf. 3. ci\. Depanpa ment of Army and the USDA ForestSer. B.SmaI mammatsD A t i nranagcd. Waslingron. Forest Ecologv and Mrnagemenl: Carcy.1. . J. J. end M. A s h b r . 65i I Fala. N. and LeachandGivnish 1999. Edu cadon. . cuttingScot'sbroon (Usseryand Krannitz1998. and old growth tbrcsts. C.B u l e t i n 1 8 . EcologicalN{onographs 6 9 : 3 5 33 7 4 . 2001.tanagemenr I 01. mal popularionsin a mixcd oak clearaut.1E Wilson and Carcy .vice cific NorthwestResearch Station.L. The countN aboulCanrpLewis. C a s b v . 1970. \'erts.Burning could have a short-term negative elfecton creeping volcs(Fala1975)and couldpromoteScot'sbroom andexotic sod-forming grasses ovcr native grasses like Idaho t'escuc lfe. c o l o g y5 t : 1 0 1 8 -t 0 2 6 . 1996. H a $ e s . c n i r o r i a l i r y:.trrrr grije"rl popuiation in lhe Pugcr Trough. A .. Carc}. T. statusol the $'cslem gra. Currcnt R.arur. F r a n k N t .10. Nofth r e \ r S c i c n c e 5 r 3 3 3i .i. R . Joumal of Mamlnatogv 58:35.. ['ashingron. FArno 72 (cditors). Plan!andnummal changcs a c learcur on i n w c s r c e n t r a O r e g o n .D. Carfawa. 1 9 6 7 . Hard) and S.027.aphNo. improperl5 incon:isit or lcnllv applied (Agee 1996.. Hans\on. Washingt0n.. l E Gashrlilcr.Todd Wilson.?rand . 8 .'. l9I8.jr. Literature Cited Agee. So. t L 7 C d h o u n .Loss of oaks could also mcan the loss of soil food webs and invenebrate communities that support vagrant shrews(Hawes 1977).nish. Acknowledgements Fundingwas providedby the Environmentaliind NatrualResources Division.10. 995. B. A . 55. Thc impacrof Scot'sbroom on oak ecotonefauna requiresadditional study.Dee.U l a h .lclrtr 5ri. Induced helcrogeneiry in Dougla\ lif canopies:effccN on smalt mammals.Continuingencroachment ofScot s broom and non-nativegrasses seriousfactorsafl'ectare ing nativepraideplantsl ThomasandCarev 1996.We thankGreg Braden.Oikos 26i398-. F. Leach. C.A. K. l . I Leighton. M Witson.Oltlnpix. Cur\rl . J.sir\. J."ruk{i. Acta Theriologica:10:4 5 .DavidFord. B. l-amborn. Pages l3 r? t r o n .Thc Natural Hisro. 1 9 1 7 . Ofelon Gash$iler J.1. . Thc use of iirc in foren resroration.ttttLuitlalutu.natLrrall. R. l 5 tsa.S i u d i e !o n $ e e c o t o g ]o f f i e l d m i c e a n d \oles |Ap.Anerican Midland Natu_ falisl 102:95. DeparrmcDl ofHcatrh. . Thyselland Carey2001)r'irh possible implications tbr the snall nammals.S m a t lm a m n a l si n sccond-gro!|lh lbfestst implications for management lor suslarmbility.Joumal of Forcslry 73:5tt6 587. Scatrte. L . Althoughthe smallmammals thatinhabit oak ecotonesare common species.t. L. Oregon State Univcnitv Pre. n1ouse reproducrionand irs rclatronshrp rherrecseed llr crop. a n d . and T. U. Charley Knox.352.Comparisonofpopnlalion characlelsrrcsol hree specicsof shre$. Durham. tah with diiferenr anounts ol coarse woody dcbrjs. 1975. L.1. Franklin. TvetenandFonda 1999) in the appropriateseason.rndS.). Joumal of Wildlile N. ('l(tttriononls gtdrcotu. 1982. .. srructure. M. Firc iD festorltion of Orcgon white oak wo|)dlands. A.and di\ersiry of rcmnanr oak sa vannas southern rn Wisconsin."er r?grd. in Jim Kenagy. and David Wallin fbr reviews of an earliermanuscdpt. K .Comparisonbetueen smlll nrannlal sampling with small and targc removal quadrals. may be acceptable methodsforprairie andoak-sitcnraintenance and restoration(Ryan and Carey 19954.The negativerelationship bet$'eenScot's broom and the deer mouse may indicate that broom areashave diminished value as wildlit-e habitat. 1988 NatLrrat Vegetarion of Oregon and Warhington. Ec(rlogical pplicatjons :116.. Fon Lewis.. 1985.and seed tilll iD a larch fir tbrcst.m d e c o l o g i H l calscparationin symparricshre\\. public Healrh MoDog. Pnnrer. and Angela Lovato for assistance thefleld andJimAgee. I 975. 1999. 1991.7rr.M . .young.C. ard B.podionof theoakareas rcmaining westem in Washington. Dymess. K.vsquirrel (S. \'ashing lon. and T. D.)'r. habitat distul baDce. Wihol].rrtr) in Houghalt \bod.1 0 0 1 .1 9 5 8 C a l c u t a t i o n f h o m e V o range :md de. o m c r a n g e .Jricus. Johnson. Slowing Douglasfir regeneradon.).12.+33.1. and Wtlfare Public Health SeNice.Tveren ard Fonda 1999). 1 .O g d e n . l. Etects of prescribed bum ing on small man.1-366.. 8 . S. M. 1979. A.y ofpuget Sound Counrry.11 lnlennountainForestand Rrnge Elperiment Sta. Wash_ i n g t o n c e o l o g i c a l S u r v e \ .USDA fofest ScI!ice. ajencralTechnicalReport INT-GTR 3.

s ).rl0g} 77:857 869. \\rest.R e m o v a le x p e r i m e n t o n s m a l l q u a d r a l s s a s a m e a n s f r a p i d a s s e s s m e o f t h e a b u n d a n co f s m a l l o nt e m a m n l a l \ ./eftbft calculationsof trappirg re\ults.. 2000.]qaf. Fl Ruggiero. and F.rti in $'ashington. B. J. H. . .Porlland. B. Carcy.S. Wilson.1998.) Link.D. andA. L .l'aslington: distribulion. Stein..Thc associationbetweenanthropogenic prairies and important tbod plants in wcstcm wash ington. (hegon. )\lorthwcll Science 2:268-273. L. o p u l a t i o n s . P r e s t o n F .Populationdlnanics ol lhc wandcring shrelv51rr. and sensiriveplanrsofFoI1 Lervis. andA. Olympia. l995ir. Peles. B. $'. S . D. J U l u q o P o p u l l l . Il. E. 1 9 8 2 . W L 1990. Endangefed.B . B. Fonda. R. \. USDA Soil ConscrvadonScnice. Northwest Anthropological ResearchNoles ll:175-200.and M. Carey. 1996. F l 9 . Burns and. . Carrarvay. M. Joumal of Nlannalogy 5. Nc$man. Land Mammals of Oregon. Quercus gdrt)a a Doxgl. Unilersity of Califomia Prcss. J|.. 1 Terrl. P ]!1.J.r i o n .E c o l o g y6 3 : 9 9 2 ' 1 0 0 3 . and R. PoIlland.Ecology and Nlanagemenr North ol A m e r i c a n S a v a n n a sU n i \ e r s i t y o f A r i T o n a P r c s s . J. 1973. . Ameican N{idland ). and F.t -. A. Pringle.1 9 9 E .threatcncd.Effccl of bail in live trappirg Pr. and L. Rice.laturalist106:119. R.. Cl.C. Seaitle. \\'ash ingon. Noflon. Legacy retention !ersus thinning: inlluences on snall mamnals.D.N_orthwcst Scicnce 7 0 : 1 . Carey. Ex Hook (Oregon white oak). Cauey. K.v and managemen! of ihe western gray squirrel and Oregon white oak $oodlands:$'ith emfhasison the PugerTrough.Exotic plant r e c o r d sn t h en o f i h w e s U n i i e dS t a t e s 9 5 0 . B. Silvicsol North America: Hardwoods. em WashingtonCascadcRange. a n d P G . D e p a r l m e n t f A g r i c u l t u r e . B.y ragmnr.J. G . Stickcl. l 7 : 2 1 72 2 l .< t r r n f . \\'. i c h e so f a d u l t a n d j u v e n i l e e e r n i c c N d (Peton)"scustna iculotur) in seral stagesofconifero u s l b r e s l .Quercus munidesinthe PugelTrough. 1 gan \and conlThysell.rlrr Sibb. F. Ct e]. 1965. Pa sikallio.. 1 515 E .125. Slril Sur!ey oi Thurnon Count).. I 948a.lournalof Biol ogy 3. \{ltl.1216.r and Ncrr"rri.)l". Brrett. Pankakoski. D.P s e u d o o g n o r n a l d i s t r i b u d o n sE c o l . R. 1 9 7 1 . 1 S t i c k e l . Journal of Wildlife Ntanagemert 12:153 61. The wavnann . Nol1hw e n S c i e n c e : l : 1 3 11 ' 1 5 . Aubrv.Nonhwest Science 5:219-235. S. and A. K .1 8 5 .Norlhwesr Scierce 69:20. Thomas. A.rlnd V Kaneno. o $/ashineton. Ryan. H.1. A. M.Uni lersiry of wishington. L r r' n r .rs. Iliolog.): the relatrvccon seNation merits ofpulling lersus cutting. u nl l r ( r L . andA.. Distribution and habitat ol the \restem gray squirrcl (Sc. Ph.. HuiI (technical coordinatort. Lewis. T u c \ o n . 7 V a n H o r n e . Oregon. i \ ' e r s i f i c a t i oo f g e n e s . o n t r o l o f S c o l \ J C broom (Crllsrs r. \\'. Pacific Nonhwcst RcsearchSlation. 72: Trclcn. 1991 Small mamnralcomrnunilics in fie sourh .IL Honkala(lcchnicalcoordina(ns). 1997. A n n a l eZ o o l o g i c iF e n n i c i8 : l l l . WashiDglon.1:295298. l . U .$/ashirgtur. C. 1976..Corection fbr sprung traps in catch."-nik. T. and G.\{cPhcrson. Effects of legetative covef on the population dlnamics of meadow voles. L.1:235'250. D n and species:evoluiionary genelicsofreal aDd!iftual pockel gophers1?ionomt r). 7 U s s e r y .U SDA FbrestSer\ ice Agricult u r e H a n d b o o kl . H. 1 9 9 9 . Notes on Brilish mammals: the trap linc indcx to snlall nammal populadons. l . ReceivedI0 Septernber 2000 AcceptedJorpublication I June 2001 Small Mammals in Oak Woodlands 3. Clarke.Pacilic Norlhwc\l Resealch Surion."rrr srirerr) oD Ft. R. K r a n n i t T . 1999. B. 1998.. Habilai ditl crcndadonamongthrce species of S. N. 1 9 8 1 . Carey. Pages650-6601" R. Bcrkeley. 7 \ \ o l l l .1 9 9 6 :n i t 1 a ecological assessment.USDA ForestServiceGeneralTechnicalRepon PNIV-GTR3. mapping.K. dcnts are corclared $ith producrionof acorns. R. 7 Tone). Ryan. Southem.. N o n h x e s t S c i e n c e 3 :1 . 1 8 -6 3 . F 1990.Fire effects on prarrles and oak \\'oodlandson For! I-c$i\.X9 . l 8 r .1995r.L...A r i z o n a . B.rrr)r. Joumal 01 N{amm. H. J. andA. and nanagemeDrreconmendations for sfecies conservation. Wildlife and vcgclalion of unmanaged Douglas-firforests.2O0l.lelson.washington. C. I 9 8 I .USDA ForestSer\iclr Ceneral Techrical Repoft PN$'-285. Washington.18.. p S t e i n b e f gE . Verls.D. \\'ashington. ogl62:355-36.r (L. J o u m a lo f W i l d l i i c N l a n a g c m c n t2 : 2 1 12 1 2 .Journ a l o f M a n m a l o g y7 7 i 8 5 0E 5 6 . I ' h e t r a p l i n e a s a n r e a s u r c o f a l lm a m sm mal populations.Joumal ol Z o o l o g y l . 1979. California. 1996. W . 8. NonhNestScience I 98-209. Disseflalion.Pages269 2831x L. Forcella.

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