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Interaction of Fungal Sporocarp Production with Small Mammal Efren Cazaresand DanielL. Luoma, Departmentof ForestScence OregonState Unvefsty, Corva is Oregon 973317501 MichaelP. Amaranthus, USDA Forest Serv ce Paclfc NorthwestBesearch Stat on Box4'10 GrantsPass Oregon 97526 CarolL. Chambersr,Department of ForestScience Oregon State U n versty. Corva s Oregon 97331-7501 John F.Lehmkuhl. USDA Forest Serv ce PaclflcNorthwest Research Station, Wenatchee Wash ngton 98801 Interactionof FungalSporocarp Production with Small Mammal Abundanceand Diet in Douglas-firStands of the Southern CascadeRange Abstract Snlall nanmal populariondensitie! arc highl! \'.rfi.rbleacross fofest stundsand landscapes.Thc speciescomposilion and abun- rlarcc of ecromy:conhizatfungi (EN{F) nraf influencc the abilit} of for.sls to pfovide suitablchabitrt for snlall mammals. Iden tilica(ion and interprctationof chang.s in the rbunduncc of the\e organisms.or in their inrcr rclationshrpsduc lo cxperimentrl har!cn. requife thar \c ilrsr idenrii_v(hc pattefnsllnd porcn!ial causesof nriLrra!r'afiabilily in drc pre-h.rNestcommunjtie\' Pre- treahlcnt data $ere gathcredtiolll the Warson!-alls block of a green-treerelcnlion experinent lo cslablishblL\eline conditions. The sir crpcrimentai lrcal cnts that comprise ihi,i block lic in r$o sprtiall) di\ljncl areasthat dillcr in en|ironmenl and ibresl composirion. the initial !ariabilit) in ENIF. snrall maDrnals.and lheif rel.rtionshjpswas documenlcd. Three prilnary qucsllons xre addrcsscdin this papcri (l) Afe the abundanceand specic! compositionof ENiF sporocarpssinilar bet$een the two arca\ ol (3)Forcommon the Vatson l-alls block l ( I ) Hox doessporoc arp consumplion\ ary rmong small n1annnalspecic\ and h) e.Ll rru11]cgcnera. is sporocarpbiomrss conclated$'ith the sporciiequency oftho\c generaiD smNllmlmmxldiel\? TheWalson Fall\ bloct iras founrtlo have spatial rnd lcmporrl larirrioli in ENIF production. small mammal n,vcophrgy..rnd srrall mrnnnal lbundance. Ho\\e!cr. truiTles$ere con\istenlly the primar,vfood item in thc diet of.rlL three lmall mamnal sPecicsin thrs study Snull rna'nmrls rre polcniall,\'illrpo(ant agentsofrruIl. dispeA.Llinlo dislurbedafers \NhercEMF afe locally c:.liryated. This studyfu hcrskno$ledgcoflheroleofsntll manmal nycophigy inthe funcaonjngof fofestccosvsGms. lntroduction tion. varyilg trom a lew scatteredtiuitbodies to conceDtratedclustcls of numerousfruitbodies (EMF) lbtm symbiotrc Ectomycorrhizalfurgi (Nonhet al. 1997,States and Gaud 1997, Whters with the rootsoftrecs andothet vcg- relationslips et al. 1997).In plannedexperinrenls, docunen- ctiltion. Trees supply carbon fuom phok)synthe tationof eristing fruiling patternsis critical to sis to the fungi. ir turn. EMF absorbminetals the irterpretationofchanges that nay resultfron] and nutrientslronr the soil and transfer them to rnanipulationof treatmentunits. hcc roots(Smith and Read 1997). Mycorrhizae ale essentialfor survivaland gro$'th ofnost con- Small nammals are integralcomponcnls ot iferoLrsforest trces alld olher shntbs anclherba man_"-forcst ecosystemsincluding the Douglas ceousvcgetation (Fogcl and Trappe 1978). Ir (Pseudotstrl.iunerate.sli) forests that dominate nuch ofthe rvesternCrscade landscape. Numerous Ectomlconhizalllngal specicsvary in thcir speciesol forestdwelling, srnall natrmals rcly rbundanceand phcnology oft'ruiting (Fogell98l; on the fruiting bodiesof EMF asa primary source Huntand Trappe 1987: Luoma 1988, 1991 I Luoma oflixrl (Fogeland Trappe 197E, Mascr et al. 1986, et al. 1991.1997r Amaranthus et al. 199.1:Nofth and Maser 1988.Hall 1991,\,\'alers and et al. 1997:States and Caud 1997)and in the Mrser 1995).A fbocl-wcblinkage of pa icular nutritional value of thcir fruit bodics (Fogel and Zabel thecldangered nodhcm spotted Trappe 19781.Various abiotic and biotic factors interestis between (Strir ..rrri?.r) and lhc nofihelt intlucncethe lruiting of EMF (Villeneuveet al. owl ot:t:itlentall.r (Glaricotnts The norlhem 19911.Sporocrrps are non-uniform ill distribu flying squinel sabrinus). spottedowl fecdsprimarily on flying squirrels (Forsman Cuncrt addres\:School of Fofe\tf).\orlhern Arizor| Unr over most of their range et al. 198.1. \cr\it!. POBox 15018,trhg\t.rll. AriTona Sfi)ll 50ltl Thomasctal.1990, Carcl' l99l). Nodhcmflying o+ NorlhwestScience, Vol.73. Speciallssue, 1999 (!r lt!! l.\ rhr \oirhre\L S.,.dr'ni1\nnrrrr.n \lln!h6r.\enr(l squin.els,in tum, require truffles, bclow-ground diversemosaic ofhost species,habitats. and struc fiuiting bodiesof EMF. asthcir primary food source turespronrote EMF diversitvat thescale oIland- (Maseret al. 1985.Carey 1995). Other mvco- scapcs.Variation in communitiesol EMF u'ill phagous(fungus eating.) small mammals. such as retlectvariation in fbresttype. successional stagc, chipmunks(TZurlas spp.) are prey lbr raptors(e.g.. and the distinctiveassemblages of plantsand goshawks)tud mammaliancamivores (e.g., martcn microhabitatswithin eachof these.Silviculturrl andlisher) (Foge1 and Trappe 1978. Mclntire 198.1, practlcescan bc usedto createhabitat colditions Hayeset al. l986. Carey1991). that maintainor increaseEMF divcrsity. For ex- The presenceand abundance of EMF species rmple.1<ar ing lurge tree. l, rllou in! regencrrti,,l may chaDgeduring tbrest development(Trappe harvests("gleen-trcc' retention)mirintrins an 1977,Mehus 1986. Tennorshuizen l99l). Truffle energysourcc tirr cefiaiDEMF speciesand pro- fungi areprimarily dispersedby sntall mamrnals vides tirr firtule recruitnent of coarsewoody de that eatthe sporocarysand subsequentlydisperse bds, an inrportant habitat componentfor some spore-packedfecal pellets (Fogel and Trappe l97lJ, EMF (Amaranthuset al. 1994). Masel and Maser 1988). The lcvel and pattem Until recently,lirnited consideration has becn of structuralretention in harvestunits may influ- givcn to the conservationof EMF. dcspite their encesmall mammal species dirccdy. thereby in- known ecologicalimportancc. The Northwcst directlyinfluencing spore dispersal of EMF. Grcen- ForestPlan (USDA and USDI 199.1a)norv di- treerctention may aisointluence thc composition. recl\ Inre5lntanrper\ lo incorporrlerirre \p(cie\. abundance,ard fruiting of EMF and thus indi- includingtungi. into futurelbrest managenent rectl)'inlluencethe abundanceof small mammals activities. The standardsand guidelines of the yla e1l-ectson their fbod soLlrces. Planrequire that t'ederalecologists and botanists Studiesof animalmycophagy arc mainly based create rnd naiDtain databasesof knotv sites of on stomachcontent or fecal pcllet analysis(Fogcl rrrrelunci. rnJ Ih t the)dcrelop .pel ie. or rrerr andTrappe 1978. Maser et al. 1978.Maser and managcmentphns forthcsetllxa(USDA andUSDI Maserl988, Carey 1995. Watcrs and Zabel 1995). 199,1b).The report lists 527 specicsoftungi thirr Theseanalyses can provide an accuralc record of afe thoughtto be closeh associatedwith late suc an animal'sprevious mcal. Fecalsample analv- cessionalforcsts; lipproximately 807r ofthesc rre sis provides a non-lethal mcthod fbr long term thought to be EMF Srudiesof the spatial and andintegrated studics ofdiet habits.Forcsfdwell- tempomldistributions of EMF ald of thcir con- lng smallmammal species that depend upon liuit- surnptionby smallmammals in unnranagedlbr- ing bodiesof EMF containa diversearray of truftle estsare a necessarvtirst stepif u'e are to under- generaiDtheir fccrl rnaterial(Mascret al. 1978). sland the coDsequcnceof flture managemenl Small mammalpopulation densities arc highly practiceslbr thesecritical clcmentsofforest ec()- variableacross stands and landscapes(Carey et systems. aJ.1992. Rosenberg andAnthony l992.Wirr 1992) ln this papcr we describesome of thesercla- anclthe speciescomposition and abundanceof lionshipsin lbreslsthat will be haNcstedas pa EMF fruirin-qbodies may inlluencethe abilirvof ofthe Demonstrationof EcosystemManagcnrent loru:l\l,r Irro\ idc \utlJblc hrhitill lol smlll r Jrn- Options(DEMO.) srucly (Aubry et al. 1999).The mals. Identification and interpretation of chalges strongcontrasts in ler,elsand patternsof carropy in thc abundalceof thcseorganisms. or of their covcr producedby thc six retention-haNesttr.eat inter relationshipsdue to forcstharvest. requirc mentsarc likely to influencehabitat features and that we filst idcntily the patternsand potential envirolrmentalconditions that in turn will alfcct causesof naturalvtriability in the prc-harvest populationsof EMF andsmall narrnrals. How- communltles. ever.initial difterencesin Lhedistribution and The high divcrsitv of EMF in lirrests of the abundanceof EMF anclsmlll mammalsmay have Pacific Northwest suggeslsthar habitatsand cn- equally important etlects on post-treannentre- vironmentalconditions are also diversc. While sponscs. some EMF may thive in intensively mtrnaged Thc six experimentalunits that we consider torests,most arelikely adversclvafiected by such rn the cuncnt study(the Watson Falls block) lie nanagement(Anraranthus and Luoma 1997).A in two spatiallydistinct areas that diller in envi- SnrallManmal Mycophagy 65 ronmentand forest composition. Thus. it is nec- dispersed)applied to the 15 and,109creteDtior essaryto quantil) the initial variability in EMF. treatnerts. The aggregatedpattem consistsof small mammals,and thcir interactionsto facili- residualtrees retained in I ha circularpatches; tate the sepantion of treatment-eflectsfiom an) the disper-sedpattern consists of lesidual tlees etlectsduc to prc-htrNcsldilTcrcnccs. Tothiscnd, unitbnnly dispersedthroughout the unit (seeAubrl we llose three fundanental questionsabout the et al.ll999l for detlils). .patialirrd tenrporul\
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