the aspen mortality summit
TRANSCRIPT
The Aspen Mortality Summit
December 18 and 19, 2006Salt Lake City, Utah
United States Department of Agriculture / Forest Service
Rocky Mountain Research Station
Proceedings RMRS-P-60WWW
April 2010
Bartos, Dale L.; Shepperd, Wayne D., comps. 2010. The aspen mortality summit; December 18 and 19, 2006; Salt Lake City, UT. Proc. RMRS-P-60WWW. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 21 p.
Abstract
The USDA Forest Service Rocky Mountain Research Station sponsored an aspen sum-mit meeting in Salt Lake City, Utah, on December 18 and19, 2006, to discuss the rapidly increasing mortality of aspen (Populus tremuloides) throughout the western United States. Selected scientists, university faculty, and managers from Federal, State, and non-profit agencies with experience working with aspen were invited. Participants were first asked to share information on recent aspen mortality. Subject matter working groups were then asked to determine factors associated with recent aspen mortality, recommend research needs, and organize those needs into testable questions and hypotheses. This report documents their findings, and will serve as a platform for Resource Managers to address the Sudden Aspen Decline issue.
Keywords: aspen, Populus tremuloides, Sudden Aspen Decline, aspen mortality, aspen diseases, aspen ecology
The Compilers
Dale L. Bartos is a Research Ecologist, with the U.S. Forest Service Rocky Mountain Research Station, and an Adjunct Assistant Professor in the Department of Wildland Resources at Utah State University, Logan, UT.
Wayne D. Shepperd (retired) is a recognized authority on aspen. Wayne published numerous scientific papers on aspen ecology and management during his 37-year career with the U.S. Forest Service.
Contents
Introduction ................................................................................................... 1
Issue ............................................................................................................. 2
Aspen Summit Meeting ................................................................................ 4
Forest Health Working Group Report ........................................................... 4
Forest Ecology Working Group Report ......................................................... 6
Wildlife Ecology Working Group Report ....................................................... 9
Genetics Working Group Report ................................................................ 10
Silviculture/Management Working Group Report ....................................... 12
Terminology ................................................................................................ 14
Overall Needs ............................................................................................. 16
Definition of Terms ........................................................................... 16
Role of Herbivores ........................................................................... 17
Role of Aspen Roots ........................................................................ 17
Genetics........................................................................................... 17
Rapid Assessment ........................................................................... 17
Recommended Actions............................................................................... 18
References ................................................................................................. 20
Appendix..................................................................................................... 21
1USDA Forest Service Proc. RMRS-P-60WWW. 2010
Introduction
Quaking aspen (Populus tremuloides) are the most widespread tree speciesthroughoutNorthAmericaand is found in theRockyMountains fromCanadathroughtheUnitedStatesandintonorthernMexico.InthewesternUnitedStates,aspenaremostabundantinColoradoandUtah.Acrossitswesternrange,aspenareamid-elevation,shade-intolerantspeciesthatoccupiesawidevarietyofsites. Aspenareoneofthefewbroad-leavedhardwoodtreesfoundinmanywesternforests.Itisavaluableecologicalcomponentofmanylandscapes,occurringinpureforestsaswellasgrowinginassociationwithmanyconifersandotherhardwoodspecies.Whileaspenprovidesdesirablescenicvalue,thediversityofunderstoryplantsthatoccurinthefilteredlightundertheaspencanopysupplycriticalwild-lifehabitat,valuablegrazingresources,andprotectionforsoilandwater.Aspenprovidescriticalbiodiversitywhereitoccurs. Aspenthrivewhereregularandfrequentdisturbance(typicallyfireinmanywesternlandscapes)promotesvegetativeregenerationthroughrootsuckersthatrisefromlateralrootsthatliewithin6inchesofthesoilsurface(DeByleandWinokur1985).Aspensproutprofusely(upto500,000stemsperacre)followingadisturbancethatkillsorremovesoverstorytrees.Thesehighnumbersofaspensuckerstypicallygrowveryrapidlyandself-thin followinganegativeexponentialdecaymodel.Mostlossesoccurinthefirstfewyears(Shepperd1993).ManagementactivitiesrelyonthisprocesstoregenerateandrestoreaspenforestsinthewesternUnitedStates.ThemostcurrentliteraturepertainingtoaspenecologyandmanagementwassummarizedbyShepperdandothers(2006)fortheSierraNevadaarea,butitwouldpertaintomostwesternaspen. Comparedtoconifers,aspenarerelativelyshort-lived.Theoldestknownaspenstemsareonlyabout300yearsofage(Shepperdandothers2006).Mostaspenwereunlikelytolivemuchbeyond100yearsunderthenaturalfireregimesthatexistedpriortosettlementoftheWest.However,duetotheabsenceoffireoverthepastcentury,aspeninmanyareasareoldertoday,havesucceededtomixedconifer/aspenforests,andaregraduallylosingmaturestemsasthestandsage.Additionally,heavygrazingfrombothnativeandwildungulateshasdamagedmanyaspenforestsbyeliminatingordamagingaspensuckersthatareproducedtoreplaceoverstorytreesastheyageanddie(DeByleandWinokur1985).Techniquestoidentifyandrestoreaspenintheseconditionshavebeendeveloped(Shepperdandothers2006)andarebeingputintopracticethroughouttheWest.
The Aspen Mortality Summit
December 18 And 19, 2006Salt Lake City, Utah
2USDA Forest Service Proc. RMRS-P-60WWW. 2010
Issue
Recentreportsofdyingaspenhavecausedconcernaboutthefutureofsomeas-penforests.Thismortalitydiffersfromnormalaspenvegetativesuccession(fig.1)orage-relatedreductioninstocking.Maturetreesaredyingatanacceleratedratewithlittleornonewsproutsoccurring,indicatingthatthelateralrootsmayalsobeaffected(fig.2).Ifthisisthecase,thenaffectedaspenwillnotbeabletoproducenewsuckersandaspengrovesthathaveexistedforhundredsandperhapsthousandsofyearscoulddisappear(fig.3). Thisrapiddeathofaspentreesseemstobegininepicentersandspreadradiallythroughanaffectedaspenstand.Standsonalltopographicpositions,moisturere-gimes,andsoiltypesareaffectedandthephenomenonhasbeenreportedthroughouttheWest,fromArizonaintoAlberta,Canada.However,specificcausesofaspenmortalitymaydifferfromareatoareaandwedonotassume,atthispoint,thatauniformsyndromeisinvolvedeverywhere.Generally,therapidmortalitycanaf-fectoneaspengrove,leavingothersnearbyuntouched.Youngerageclassesandpre-existingsproutsareoftennotaffectedtothesameextentasmatureoverstorytrees.Cytosporacankers,poplarborers,andotherdamageorstressagentsareof-tenassociatedwithdie-offepicenters;however,thepossibilityofayet-unknowninvasivediseaseorinsectcausestillexists.
Figure 1. An aspen stand in southeast Idaho where mature aspen trees died but sufficient aspen regeneration exists to restore the stand.
3USDA Forest Service Proc. RMRS-P-60WWW. 2010
Figure 2. Dying aspen stand in the summer of 1990 on Cedar Mountain in southern Utah (photo by James Bowns, see Ohms 2003).
Figure 3. Same stand as shown in figure 2 during the summer of 2002. Few living trees remain and no regeneration is present (photo by Seth Ohms, see Ohms 2003).
AcceleratedratesofaspenmortalityhavebeenreportedforseveralyearsinUtahandArizona,butonlyrecentlyhavetheseratesbecomeapparentinColorado.Aerialsurveys,conductedbyForestHealthManagement,USFSRegion2,indi-catedthatnearly140,000acreswereaffectedbyaspendeclineinColoradoin2006;andby2009,thatareawasinexcessof500,000acres(seehttp://www.aspensite.org/SAD/sad_faqs.pdf).Theapparentlackofasuckeringresponseinsomecasesisdisturbing,asaspenmustsproutbackifitistopersist.Mortalityofthismagni-tuderaisesconcernsaboutthefutureofaspenforestsinsomeareasandjustifiesacomprehensiveinvestigationintothephenomenon.
4USDA Forest Service Proc. RMRS-P-60WWW. 2010
Aspen Summit Meeting
TheUSDAForestServiceRockyMountainResearchStationinvitedselectedscientists, university faculty, andmanagers fromFederal,State, andnon-profitagencieswithexperienceworkingonaspentoanaspensummitmeetinginSaltLakeCity,Utah,onDecember18and19,2006.Participantswereaskedtoshareinformationaboutrecentacceleratedaspenmortality.OveradozenparticipantsfromthroughouttheWestpresenteddescriptionsandexamplesofdyingaspen.Participantswerethenassignedtothefollowingsubjectmattergroups:
• ForestHealth(pathologists,entomologists) • ForestEcology • WildlifeEcology • Genetics • Silviculture/Management
Eachgroupwasaskedtodiscussthefollowingtopicoutlineandmakerecom-mendationstoaddresstheaspenmortalityphenomenon:
a.Determinefactorsassociatedwithaspendie-off. b.Determineresearchneeds—factorsandquestionsthatneedtobestudied. c.Organizeresearchneedsintotestablequestions/hypotheseswithbudgets.
Broadguidelinesweregiventothegroups;however,nouniversalprotocolwassuggestedastohowthevariousgroupsshouldreporttheirefforts.Eachsubjectmattergroupproducedoutlinenotesfromitsdiscussionsandpresentedthemtoallparticipants.Transcribednoteswerethenusedtoproducethenarrativereportspresentedbelow.
Forest Health Working Group Report
Clarification of Terminology
TheForestHealthgrouphadconcernsabouttheterminologyusedindiscuss-ingtheaspenmortalityproblem.Thefourtermsofmajorconcernwere:decline,die-back,die-off,andsuccession.Thesewordsmeandifferentthingstodifferentpeople.Thefollowingdefinitionsexistfromaforesthealthperspective:
• Declineistheresultofaforestdiseasethatinvolvesmultiplespecificallyordered,interchangeableandinteractingfactors(Manion1981).Theclassicdefinitionofaforestdeclinetypicallyinvolvespredisposing,inciting,andcontributingfactors(table1).
• Die-backisasymptomcharacterizedbydeathofpartsofaplant.Itisoftenpresentaspartofforestdecline.Thetermmaybeappropriatetodescribedeathofstemsinaclonalplant,suchasaspen,whenrootsremainaliveandareabletosucker.
• Die-offischaracterizedbymortalityofaspentreeswithnoregenerationtorestorethestand.
5USDA Forest Service Proc. RMRS-P-60WWW. 2010
• Succession(asequenceofchangesinaplantcommunity)isaseparateprocessfromdecline.Forestsuccessioncouldplayaroleincausingtreestodecline,butforestdeclinecanoccurwithorwithoutsuccession.
Commonlyrecognizeddefinitionsareneededsothatparticipantscaneffectivelycommunicate(forexample,theSilviculture/ManagementWorkingGroupusedthetermsdie,die-off,anddie-backintheirsummarylaterinthissection).TheForestHealthWorkingGroupconstructedtable1toillustratehowfactorsaffectingaspenmightfitintothepathologicaldefinitionofdecline.
Research Needs
Thereisaneedtoidentifycausalagentsintheaspenmortalityphenomenonthatwearecurrentlyobserving.Tounderstandthisphenomenonandmakerecommen-dations,weneedtoknowwhatagentsandstressesareinvolved,(biotic,abiotic,andsitefactors)andtodistinguishvariousinteractionsthatareoccurring. Biotic Factors—Tofullyunderstandthismortality,specificbioticagentsthatare involvedneed to be identified.Thismay includevarious diseases, insects,andtheimpactsofungulates.Someofthesebioticfactorswillplaymajorrolesinaspenmortalitywhileotherswillbesuperfluous.The roleofungulateswillundoubtedlybeamajorfactorincertainareasoftheWest.Whatisthemaximumpopulationlevelofwildungulatesthatwouldallowustostillgrowanaspentreefromasucker?Whendodomesticlivestocknegativelyimpactaspenregenerationandcanwedevelopguidelinesforthemanagertoaddressthisproblem? Abiotic Factors—Thesefactorsareprobablythemostsignificantcontributorstotheaspenmortalityproblem.Themostprevalentfactorcouldbedroughtanditseffectsontheaspenclone.Howdoesdroughtimpacttheaspenoverstoryaswellastheregeneration?Doesdroughtactasastressortostimulateregenerationinadditiontokillingmaturetrees? Site Factors—Additionally, therolesofvarioussitefactorsneedtobebetterunderstood.Factorssuchaselevation,topography,andsoilsareallanintegralpartoftheaspenecosystem.Whatpartdothesefactorsplayandhowdotheyinteractinthedemiseoftheaspen?Worrallandothers(2008),inaquantitativestudyofnumerousaffectedaspensitesinwesternColorado,stated:“Ourdataareconsis-tentwithahypothesisthat(a)predisposingfactorsincludestandmaturation,lowdensity,southernaspectsandlowelevations;(b)amajorincitingfactorwastherecent, acutedrought accompaniedbyhigh temperatures, and; (c) contributingfactorsandproximateagentsofmortalityarethecommonbioticagentsobserved.”
Table 1—Factors potentially involved in aspen decline.
Predisposing Inciting Contributing
Marginal sites Frost Animal damageOld age Acute drought Canker fungiWater tables Sucker mortality from browsing BorersSuccession Defoliation event BeetlesGenetic mutationIncreased temperatures
6USDA Forest Service Proc. RMRS-P-60WWW. 2010
Testable Questions
• WhatisthemagnitudeandextentofaspenmortalityintheRockyMountains? • Whatsiteandbioticfactorsareassociatedwithaspenmortalityinthewestern
UnitedStates? • Whatmeteorologicalfactorsareassociatedwithaspenmortality? • What are the long-term effects of diseases and insects on the aspen
ecosystem?
Partners
Long-termresearchisneededtoaddressvariousaspectsofaspenmortality.Thiswillallowforthedevelopmentofmonitoringprotocolsandmanagementtechniques. Numerouspartnershavebeenidentifiedtocontributetotheselong-termefforts:theForestRestorationInstituteatCSU,theproposedCenterofExcellence(sincemorphedintotheWesternAspenAlliance)atUSU,andtheU.S.ForestServiceFIA/ForestHealthMonitoring.
Forest Ecology Working Group Report
OurcurrentknowledgeofaspenecologyshouldbeusefulinunderstandingtheextensiveaspenmortalitythatisoccurringintheRockyMountains.However,thereismissinginformationconcerningtheecologyofaspenthatwouldhelpdecipherthismortality.Thisgroupapproachedtheirsubjectintwoparts:(1)suspectedfac-torscontributingtotheaspenmortality,and(2)significantquestionsthatneedtobeevaluated.
Mortality Factors
Arapidassessmentmethodologymustbedeveloped todefine themagnitudeandextentoftheaspenmortalityproblem.Thismethodmustbeusefultore-sourcemanagersandapplicablethroughoutthewesternUnitedStates.ThefirststepinachievingthisgoalmaybetoadaptthemodelspecifiedintheSilviculture/Managementbreakoutgroup. Inaddition,a remotesensing techniquecouldbecoupledwiththerapidassessmenttomorepreciselydeterminethismortalityandbetterdefinetheextentorpatchynatureoftheproblem. Droughthasprofoundimpactsonmostvegetationsystems.Doesdroughtcon-tributesignificantlytothecurrentaspenmortality?Howdoesthismoisturestressrelate toelevation,aspect,etc.?Recently,Worrallandothers(2008)foundthataspenmortality,orSuddenAspenDeath(SAD),wasmoreprevalentinsituationswheredroughtwasmorepronounced. Aspen are a sun-loving, disturbance-dependent pioneer species that quicklyre-colonizesasitefollowingfirebutwilleventuallybereplacedbymoreshadetolerantconifersifasuitableconiferseedsourceisavailable.Givenenoughtime,mostaspenwilleventuallysucceedtoaconiferdominatedsystem,butathirdoftheaspenintheWestcanbeconsidered“climax”orstableformanagementpur-poses(Mueggler1989),sincenoconifersarecurrentlypresent.SADiscurrentlyaffectingbothstableandsuccessionalaspen(fig.4)andwillundoubtedlyaffectthefuturecharacterofbothtypesofaspen.
7USDA Forest Service Proc. RMRS-P-60WWW. 2010
Successionisusuallyaffectedbyfire.Historically,manysitesburnedonaregularbasis,soconiferregenerationwasremovedandthestandsrevertedtobeingaspendominated.Becausefirehasdiminishedinthesesystems,aspentreesarebecomingmatureorover-mature.Astressorsuchasdroughtwouldgiverisetothepossibil-ityofmoreincidencesofdiseasesandinsects.Inadditiontofire,numerousotherfactorscancontributetotherejuvenationof“stable”aspen. Aspenusually regenerates in thewesternUnitedStatesbyvegetativemeans.Seedlingsdooccurbutnotonalevelthatwouldmeanwideexpansionofaspenseedlingsacrossthelandscape.However,withrecentdevelopmentsinthefieldofgenetics,thereismoreincidenceofregenerationbyseedsthanwasoriginallythought(seeGeneticsgroupreport). IncertainareasoftheWest,thereishighincidenceofherbivoryofaspenbybothdomesticlivestockandwildungulates.Theseherbivorescanhaveasevereimpactonaspen regenerationandonmature trees in somecases.Someaspenclones/standshavebeeneliminatedfromthelandscapebyexcessiveungulateuse.Whatroledoungulatesplayinthecurrentmortalitybeingobserved?Isexcessiveusebyungulatesremovingtheregenerationfromfadingmatureaspenstandscausingsomeofthemortalityweareexperiencing? Quakingaspenin thewesternUnitedStatesoccursoverawideamplitudeofconditions,fromshrub/woodlandecotoneonthelowerendofitsrangetotreelineattheupperlimits.Willtheprojectedclimatechangehaveaprofoundchangeinaspendistribution?Itisbelievedthatclimatechangemightcauseaspentorecedefromdriersitesatthelowerextremesofaspen’soccurrence.
Figure 4. Mortality of mature aspen in a mixed conifer/aspen stand on the Gunnison National Forest 2006.
8USDA Forest Service Proc. RMRS-P-60WWW. 2010
Aspen Ecology Questions
• Areaspenrootsdying?Ifso,why?Ifdiseaseorinsectsarepresent,aretheygirdlingtheparenttreeandcausingtherootsystemtodie?Aretherootsbeinggirdledandcausingtherootstodie?
• Does herbivory act as a vector for pathogens? Do herbivores transferpathogens?
• Terminologyneedstobeclarified.Howdoesdeclinedifferfromdie-off/die-back?Weshoulduselessconfusingtermsthatreferjusttoaspenmortality.Specificadditionalwordsneedtobedefined,suchasstableaspen,clone,stand(grove),succession,anddisturbancespecies.
• QuestionsattributedtoWayneShepperd(giveninhisopeningremarks): o Ismortality(die-off)anormalevent? o Aretherenewdiseasesorinsectspresent? o Canthismortalitybepredictedbystandage,growthrate,stocking,or
othermetrics? o Isclimateinvolved? o Howlongwillthismortalitycontinue? o Whatcan(orshould)bedoneaboutthismortality?
• Isthiscurrentphenomenonjustareturntohistoricconditions?Ifso,shouldwebeconcernedaboutit?
• Isthereameansto“age”clones(seeGeneticsgroupreport)?Doesdiseasebecomealargerplayerastrees(clones)age?
• Howcanwebuildpublicsupport(fortreatment)tostopthelossofaspenincriticalareas?Whataretheeconomicsassociatedwiththismortality?Whataretheimpactsonthepublicasaresultofdiminishedfallcolors?
• Shouldwemimicdisturbanceeffectsinordertopreventthelossofsomeaspenclones fromthe landscape?Should treatedsitesbeprotectedfromherbivores?What newand innovative techniques couldbeused to keepherbivoresfromeliminatingaspenregeneration(forexample,hinging,slashpiling,andharassinganimals)?
• Whathappensaftermortality?Whatarethelong-termimpactsofthismor-talityonaspenclones,understoryvegetation,soils,scenicandrecreationresources,andanimalhabitats?
• Whyaresomeclonesaffectedwhileadjacentonesarenot?Aresomeclonesgeneticallypredisposedtothismortalityorisitjustarandomoccurrence?
Specific Questions and Approximate Costs
• Quantifythemagnitudeandextentofthemortalityissue($200K). • Definetheproblemandterminology;reviewandsynthesizeliterature;com-
municate researchneeds; anddefinepertinentquestions tobe addressed($50K).
• Developadie-offriskassessmentandakey(decisiontree)toidentifystandsatriskandwhichwouldbemosteconomicaltotreat($50K).
• Determinecausesofdie-off($200K): o Areelevation,aspect,andinsectanddiseasestressesresponsible?
9USDA Forest Service Proc. RMRS-P-60WWW. 2010
o Whatistheroleofstandage,drought,andherbivory? o Identifyprimaryversussecondaryagents,includinggenetics. o Isungulatesalivaavectorfordiseasetransmission?
• DeterminetheknownNaturalRangeofVariability(NRV)ofaspenandtestforaprecedenceofdie-off($50K).
• Determinetheregionaleffectsofaspenmortality($1.5M). o Inter-clonalinteractions o Successionaltrajectory o Erosion o Alleopathy o Pathogenaccumulation o Insectaccumulation o Wateravailability
• Identify and test techniques to protect regeneration and restore “fading”standsthataretreated($100K).
Wildlife Ecology Working Group Report
Key Questions
TheWildlifeEcologygroupidentifiedseveralkeyquestionstoaddresspotentialchangesinwildlifehabitatresultingfromthelossofaspen.First,thecurrentstatusandprojectedtrendsoftheaspenresourceneedtobedeterminedandhowthesechangesmightaffectwildlifeatdifferentscalesneedstobeestablished.
• Whatimplicationswill thesetrendshavefordifferentspeciesofwildlifeassociatedwithaspen?
• Whatspeciesmightbemostaffectedbylossofaspen? • Whateffectswouldthelossoflargeaspentreeshavecomparedtocomplete
lossofaspen? • Atwhatscaleswouldlossofaspenbecriticaltowildlifespecies? Understandingtheroleoffireinshapingtheaspenresourceofthefutureisalsocritical.Re-establishing thepredominanceof fireon the landscapemightofferopportunitiesforaspenre-colonizationthatcouldpotentiallycompensateforthecurrentlossofaspen,butitisunclearhowthesefuturescenariosmightplayoutforwildlifespecies.Changingthespatialandstructuraldistributionofaspenonlandscapeswillundoubtedlyaffectwildlifehabitat.Gainingspecificknowledgeofhowwildlifespeciesuseaspenhabitatwillhelpmanagersunderstandhowtominimizeconflicts,predicttheeffectsofcurrentaspenloss,andplanfutureman-agementactions. Determiningtheseriousnessofthecurrentlossofaspenandhowitwillaffectpopulationlevelsofelk(Cervus elaphus)andmoose(Alces alces)iscritical.Thesespeciesuseaspenforforageandcoverand,consequently,mayaffectthehealthandvigorofaspenpopulations.Gainingabetterunderstandingofthedynamicsofthisinteractionwillaidmanagersinunderstandinghowtheycanbuildprovisionsforsustainableaspenhealthintointegratedvegetationmanagementstrategies.
10USDA Forest Service Proc. RMRS-P-60WWW. 2010
Thegoalsforthesestrategiesshouldbedesignedtoachievedesiredfuturecondi-tionsforaspen,wildlife,andotherresourceobjectives. Anothercriticalneedislearninghowtotransferknowledgeaboutaspen/wildlifeinteractionsfromoneregiontoanother.Thisrequiresdetailedinformationonthefeaturesthatarebothcommonanddifferenttotheseareas.Wealsoneedtounder-standhoweasilytheresultsobtainedfromspecificsitesandscalescanbeparlayedintobroaderscalevegetationmanagementgoalsandstrategies.Similarly,weneedtodeterminehowtoassembleandsynthesizeelementsoftheaspenproblemfromdifferentdisciplinesandassesstheknownandunknownsinfuturescenarios.
Actions Needed
TheWildlifeEcologygroupsuggeststhatseveralimmediateactionsareneededtodealwiththecurrentaspenmortalityissuewithrespecttowildliferesources.First,areviewandsummaryofstatewide(wildlifeandfish)comprehensiveplansandTheNatureConservancyeco-regionalassessmentsshouldbeconductedtomorecompletelyascertainspecificquestionsandneedsrelatedtoaspenandwildlife.Inaddition,theUSFSprojectplanningandappealsdatabaseScheduleofProposedActions(SOPA)shouldbereviewedtoidentifytrendsandcriticalissuesassociatedwithplanningandimplementingvegetationmanagementprojectsinvolvingaspenandwildlife.Thiscomprehensivedatabasedescribesproposedvegetationmanage-mentprojectsonForestServicelandsandwouldprovideameansofidentifyingpotentialprojectsthatcouldbemonitoredorstudiedtolearnmoreaboutwildlifeandaspeninteraction(aswellasgainknowledgeaboutotheraspectsoftheeffectsofmanagementactivitiesontheaspenresource). Thegroupalso recommends thatamulti-partnerworkshopbeorganized thatwould o increaseunderstandingofaspen, o identifyresearchneedsforaspen/wildlifeinteractions,and o developanactionplanforaspenconservationandmanagementthatcould
beusedtoguideforestplanningandintegratedvegetationmanagement.
Genetics Working Group Report
Overarching Need
Acriticalneedtounderstandingthecurrentaspenmortalityphenomenonistoconsidertheclone-specificeffectswheninvestigatingdie-offsandtreatmenteffectsandwhenconductingtrendmonitoring.Theneedtoclearlydefinephenomenaandtermsinthisprocessshouldbeemphasized.Forexample,thefollowingphenomenamaybetheresultofverydifferentprocessesoccurringinaspenstands:
• overstorytreemortalitywithoutregeneration, • mortalityofallageclasseswithoutregeneration, • displacementbyconifers, • overstorymortalityfollowedbyephemeralregeneration,and • overstorymortalityfollowedbyregenerationofdifferentgenets(clones).
11USDA Forest Service Proc. RMRS-P-60WWW. 2010
It isalsocritical to identify thegenetic factorsand issuesassociatedwiththe current aspendie-off.Potential factorsat thegenet level includeageneticsusceptibilityand/orlowfitnessofparticulargenetsthatisnowbeingexpressedunderchangingclimates.Giventhesuspectedageofsomeaspenclones,anotherfactormightbetheaccumulationofdeleteriousmutationsinoldergenetsthathaveexistedon-siteformanyyears.Earlierresearchhasshowndifferencesingrowthanddevelopmentinaspenthatcanbeattributedtothegenderinthispredominatelydioeciousspecies.Gendercouldalsoplayaroleinthedistributionandoccurrenceofaspenmortality.Inbreedingdepressioncouldalsobeafactor,ascouldthelackofphenotypicplasticityinparticulargenets. Potentialfactorsthatmightbeoperatingateitherastandorlandscapescalemightincludelownumbersofgenetsinastand,lowgeneticdiversityamonggenetsinthestand,ahighdegreeofrelatednesswithinastand,andlowlevelsofsexualreproductioninsomegenotypesorunderspecificecologicconditions.
Research Needs
Geneticresearchneedsincludeestablishingtheclonalboundarieswithinstandsbeingstudiedsothattheexistenceofdifferentgenotypesisknownwithrespecttotheoccurrenceofaspenmortality.Suchgenotypemappingshouldaccompanyresearchintoaspenregenerationtreatmentsandassessmentstodetermineormea-suretheroleofgeneticsinobservedresponses.Thisshouldalsobeacomponentofinventoryandmonitoringofaspen. Identifyinggenetictraitsthatmightbeassociatedwithdie-offisimportantaswell.Thiscouldinvolvelearningwhattradeoffsmightexistamongtraits(forexample,chemicaldefenseversusgrowth)thatmightbecontributingtoaspenmortalityandidentifyingwhetherthesetraitsareheritableorplastic.Potentialtraitsofinterestpossiblyrelatedtodie-offsusceptibilityincludedifferencesincavitationpotentialamong genotypes and physiologywith respect to above- versus below-groundviability.Geneticdifferencesinrootdeathrateandsecondarychemistry(herbivoryresistance)couldinfluencesuckeringrateandsuckersurvivabilityand,therefore,mightaffectsusceptibilitytoandtheultimateoutcomeofmortalityevents.Theroleofgeneticsintreestructure,pathogenresistance,droughtresistance,andfloodingresistancemightalsoinfluencetheoutcomeofaspenmortalityevents. Otherusefulinformationmightincludedeterminingtheextentthatrootgraftingoccursamongdifferentgenetsandquantifyingwhetherlossesinaspencoveragearecongruentwithlossesinthenumberofgenetsinaspenpopulations.Determininghowaspenstandstransformfrommonotypictodiverseandback(inotherwords,clonaldynamics)mayalsobeveryusefulinexplainingtheaspenmortalityphenom-enon.Althoughitmaynotbedirectlyrelatedtothecurrentaspenmortalityissue,determiningtowhatextentgenetdiversityinfluencesthediversityofunderstoryplants,insects,andsoforthinanaspenpopulationcouldbeenlightening.Thisistheconceptof“extendedphenotype”describedbyWhithamandothers(2003). Inadditiontogeneticinformation,knowingwhatenvironmentalconditionsareassociatedwithdie-offsisalsoimportant.Factorssuchassoils,slope,aspect,grazing/browsing,othertree/shrubpresence,climatehistory,anddiseaseoccur-renceshouldalsoberecordedinanyaspenassessment.
12USDA Forest Service Proc. RMRS-P-60WWW. 2010
Testable Hypotheses
Thefollowinghypothesesneedtobetestedwithrespecttoaspengeneticsandcurrentaspenmortality: Hypothesis 1: Thenumberofgenetsinwesternlandscapesisdeclining(aseparate
phenomenonfromdecreasesinspatialcoverage). Test by: Collectinggeneticsdatainconjunctionwithestablishingsampling
plotsforlong-termmonitoringinconjunctionwithFIAatabroadgeographicscale.
Hypothesis 2: Aspenmortality is genet-specificwithin stands (HA:Die-off isacrossgenets).
Test by: Establishingsamplingplotsandcollectinggeneticsdataforlong-termmonitoringatafine-scaleresolution(fewerplotsbutmoredensesamplingthanH1),focusingonareasexperiencingdie-off.
Hypothesis 3: Lossofaspenarerelatedtomaladaptationinthefaceofclimatechange.
Test by: Usingacommon“garden”approachtoassessfitnessofgenotypesfromdifferentenvironments(clone-specificeffects).
Additionalbasicresearchneedsthatdon’tfitneatlyintotheabovehypothesestestingincludedeterminingtheextentofsexualreproductionamongaspengenotypesandstudyingmutationaccumulationandfitnessamongaspen.Developingmoreefficientwaysofsexingandagingaspenclonesandquantifyingtheextentofrootgraftingwouldbeuseful,aswouldinvestigating“extendedphenotype”effectsortheecologicaleffectsofaspengeneticdiversitywithinlandscapes.
Silviculture/Management Working Group Report
Define and Clarify Terminology
There isacriticalneed todefineandclarify terminologyassociatedwith therapidmortalityofaspen.Afirststepistoidentifywhetherthecurrentmortalityistrulyacrisisorjustanormaloracceleratedmortalityofoldtrees.Identifyinganyrelationshipsbetweenrametageandoccurrenceofmortalityiscrucial.Second,thereisaneedtoidentifyandquantifyanydifferencesinmortalitybetweenaf-fectedstandswithconiferencroachmentandthosewhereconiferencroachmentisnotoccurring.Ifmortalityisindeedstressrelated,thenthepresenceofconifershouldcontributetoeithertherateorincidenceofmortality.Finally,anycommonfactorsthatmightberelatedtocausalityneedtobeidentified. TheSilviculture/Managementgroupidentifiedthreedistinctconditionsthatcanbeassociatedwithaspenmortality:
• Overstorymortalitywhereregenerationofaspenareoccurring • Overstorymortalitywithoutsuccessfulregenerationofaspen • Overstorymortalityofaspenwhereconiferregenerationispresent,but
aspenregenerationisnotorisinsufficienttomaintainthepresenceofaspenonthesite
13USDA Forest Service Proc. RMRS-P-60WWW. 2010
TheSilviculturegroupconcludedthattherewasnocauseforconcern,fromanecologicstandpoint,ifsufficientaspenregenerationisoccurring,oralreadypresent,todevelopintoafullystockedaspenstand.Whilethelossofamatureoverstorymightberegrettablefromanaestheticorwildlifehabitatstandpoint,itiswithintheexpected rangeofnaturalconditions foradisturbance-dependent relativelyshort-livedtreespeciesthatvegetativelyregenerates.However,ifregenerationisnotoccurringthesituationbecomesmorecritical.Ifsproutingisoccurringbutnotsuccessful,thereasonsneedtobeidentifiedandcorrected.Ifsproutingisnotoc-curring,thereasonsneedtobeidentified.Ifrootshavedied,wehavetoacceptthelossofaspenonthesiteandconcentrateourmanagementeffortsonareaswherethepotentialtoregenerateaspenstillexists.
Critical Research Questions
Severalquestionsmustbeansweredinsituationswherematureaspentreeshavediedbutsuccessfulregenerationisoccurring: • Whenregeneration(aspen)isoccurringunderadeadoverstory,howoldis
theunderstory? • Doestheregenerationpre-datetheoverstorymortality(e.g.,wasthestand
regeneratingpriortotheonsetofoverstorymortality)? • Ifso,whatfactorstriggeredtheregenerationresponse? • Ifnot,whywerethesestandsabletoinitiateregenerationwhenotherswere
not? • Couldpressurefrombrowsinganimalsbeafactor? Severalquestionsmustbeansweredwherematureaspenstandshavediedandregenerationisabsent: • Didthestandsattempttoregeneratebutsomethingkilledthesprouts? • Wastheoverstorytooweaktosprout? • Hadtoomuchtimeelapsedsincethelastdisturbancetoinitiatesprouting? • Didlateralrootsdiewhenoverstorystemsdied(orconversely,didrootsdie
firstandcausetheoverstorymortality)?
Whereconifersarepresent inaffectedaspenstands, theircontribution to theconditionsleadingtoaspenmortalityneedstobeidentified: • Didtheyout-competetheaspenformoisture,nutrients,orlightandthus
initiateaspenmortality? • Didtheirpresencecontributetothelackofaspensproutingbyshadingthe
soils?
Sinceliverootsarethekeytotheaspensuckeringprocess,itiscriticalto: • identifyhowaspenrootsareaffected; • findwaysofassessingaspenrootconditionpriorto,during,andafterover-
storymortalityhasoccurred;and • relaterootconditiontoanyfactorsthatmightbeidentifiedascontributing
tothisrapidmortalityphenomenon.
14USDA Forest Service Proc. RMRS-P-60WWW. 2010
Inadditiontoansweringtheabovequestions,itiscriticaltoidentifyIntermountainWestbiodiversityhotspotswheremaintenanceofaspeninlandscapesiscriticaltoecosystemsandtodevelopameanstoprioritizetheneedformanagementinterven-tion.Tofacilitatethisprocess,thefollowingterminologymightbeadoptedtohelpdifferentiatebetweenvariousaspectsofaspensuccessionandclonalturnover.
Terminology
Die:when≥80percentoftheoverstorystemsinaclonediewithin3to5years. Die-off:notenoughregeneration(suckering)toperpetuatestanddueto
a)aphysiologicalfailureoftheaspentorespond,or b)thevegetativereproductionsystemis“turnedoff.”
Die-back:regenerationissufficienttoperpetuatetheclone
a)withpriorregenerationpresent,or b)whenregenerationcoincideswithoverstorymortality,or c)whenthereisagradualoverstorydecreaseinstandswithorwithout
conifers.
Bothdie-offanddie-backcanbeapartofthepathologicdefinitionof“decline,”butbothoccurveryrapidly.Wepreferthattheterm“decline”beusedtorefertothegradualdeathofaspenoverstoryineitherpureaspenstands,orinconjunctionwithconifer invasioninmixedaspen/coniferstands,whichisdescriptiveofanecologicratherthanpathologicprocess1.
Additional Questions Associated With This Classification
• Is themechanismforoverstorymortality thesamewhetherornotaspenregenerationoccurs,orwhetherornotconifersarepresent?
• What are the patterns of mortality progression associated with rapidmortalityofaspen?
• Doesmortalityradiatefromacentralpointoroccursimultaneouslythrough-outthestand?
• Ismortalitylimitedtoonegenotypeataspecificsiteordoesitcrossclonalboundaries?
• Istheabovedefinitionofrapiddeathvalid(forexample,3to5yearsfromonsettocompleteoverstorymortality)?
Assessment Needs
Itisimportanttogatherconsistentinformationtoaccuratelyassesswhatishap-peningtoaspenthroughouttheWestandcompileconsistentdatathatcanbeused
1 Compilers Note: In the time since the meeting, the term “Sudden Aspen Decline” (SAD) has been used to describe conditions associated with the rapid mortality of aspen trees. Within this context, we feel the distinctions made by the “die-off” versus “die-back” definitions formulated by the Silviculture Working Group above are still valid and should be considered, regardless of the terminology used to describe the overall rapid aspen mortality process.
15USDA Forest Service Proc. RMRS-P-60WWW. 2010
toanswertheabovequestions.Assessmentprotocolsshouldbedevelopedtosurveyaspenmortalitythatfallsintothecategoriespreviouslydescribed. Categoriestosurvey:
1)die-off(noregenerationpresent) 2)die-back a. priorregeneration(existedbeforemortalitystarted) b.coincidentalregeneration(initiatedbytheoverstorymortalityevent)
Thesurveyshoulddefinehowmanyacresofeachexistandlistotherfactorsthatmaybeassociatedwitheachcategory,suchas:
• insectanddiseaseoccurrence,especiallyCytospora, • elevation, • aspect, • ungulatepressure, • drought, • currentvegetation(habitattypes), • currentvegetation(soils),and • treatments (if any) used to dealwith aspenmortality and their apparent
effectiveness.
A template for providing consistent treatment application and monitoringthroughouttheWestshouldbedeveloped.Coincidentally,amechanismshouldbedevelopedtofacilitatecollaborationandcommunicationamongallaspenmanagersintheWest
Suggestions for Initial Research and Inventory
1.Ineachofthethreedie-off/die-backcategories,determinethenumberofacresinlandscapescontainingaspenthatareaffectedbyrapidaspenmortality.ThistaskcouldbedonebytheForestService’sForestInventoryandAnalysis(FIA)andForestHealthManagement(FHM)usingtheFIAdatabaseandexistingFHMinsectanddiseasesurveytechniques.
2.Quantifybioticfactorsandabioticstandcharacteristicsassociatedwitheachofthethreedie-off/die-backcategories.Thiswouldmostlikelyinvolvecol-laborationamongFHM,university,andForestServiceresearchers.
3.Determine the causal agents behind rapid aspenmortality (for example,theresponsiblemechanismsorfactors).Again,datasharingandamulti-disciplinaryapproachwilllikelybeneeded.
4.Todealwiththephenomenon,testmanagementactionsanddetermineaneffectivemanagementtimeframeforaction.Thiscouldbestbeaccomplishedbymanagersworkingcollaborativelywithuniversityand/orForestServiceresearchers.
Intheshort-term,thedevelopmentofacommonprotocolfortreatmentexecutionandmonitoringisessentialinboththeearlyandlatestagesofaspenmortality.Thiswouldincludeapre-treatmentstandassessmentinventoryprotocoltoassessandquantifytheconditionofthestandsbeforetheyaretreated,aswellascommonmetricstoassesstheeffectivenessoftreatment.
16USDA Forest Service Proc. RMRS-P-60WWW. 2010
Anothercriticalshort-termneedistoestablishaclearinghouseoraleadorganiza-tiontoensurecollaborationandprovidefeedbackamongmanagersandresearchers.Thiswouldincludesharingofinventoryandmonitoringdataandpre-publicationsharingandreviewofresearchdata.Theclearinghouseshouldhaveanofficialdatastewardtobuildandmaintaindatasets,maintainanddisseminatewebsiteinfor-mation,andensuredatacompatibilityandsecurity.Theclearinghouseshouldalsohaveapointpersonorfacilitatorwhowouldinteractwithstakeholders(managers,universities,researchers)tosecureanearmarkforfundingtodealwiththeaspenmortalityissue.
Overall Needs
Definition of Terms
Severalcriticalneedsandthemeswereevidentacrosstheworkinggroups.Fore-mostwastheneedforclarificationoftheterminologyusedtorefertothetypesofaspenmortalitythathavebeenobservedandreported.Allgroupsemphasizedtheneedforclearandsuccinctdefinitionsofthephenomenaandthetermsusedtodefinethecurrentmortalityofaspen.Subsequently,weendorsetheexplanationandterminologypresentedbytheForestHealthWorkingGrouptodescribedeclineinorganisms.However,thismodelisstillinadequatetodescribethesituationoc-curringinaspenpopulationsatlandscapescales.Termspreviouslyusedtorefertoaspenmortalityarealsoinadequateandcausedconsiderableconfusionduringmeetingdiscussions.Duringhissummaryremarks,DavidCleaves,DirectoroftheUSDAForestServiceRockyMountainResearchStation[currentlyAssociateDeputyChief,WO],suggestedtheacronym“SAM”forSuddenAspenMortalitybeusedtoavoidthevalue-ladenconnotationsassociatedwith“die-off”and“die-back”andtheconfusionwiththeclassicpathologicdefinitionof“decline.”2
However,SuddenAspenMortalitydoesnotcoveralloftheconditionsreportedanddiscussedatthemeeting.Anadditionalclassificationisneededtodifferenti-ateamongspecificmortalityeventsthathavebeenobservedandreported.Twoalternativelistsofconditionsweresuggestedbytheworkinggroups.WesuggestadoptingthefollowingwordmodelmodifiedfromtheSilvicultureandManage-mentgroup.Itprovidesaframeworkfordistinguishingamongmortalityeventsandallowsameansofinvestigatingspecificquestionsrelatedtotheevents:
1.Overstorymortalitywhereregenerationofaspenareoccurring 2.Stand (clone?) mortality—Overstory mortality without sufficient aspen
regenerationtomaintainaspenonthesite. o Coniferregenerationislargelyabsent o Coniferregenerationispresent
2 Compiler’s note: In the time since the meeting this term has morphed into “Sudden Aspen Decline,” which is consistent with the pathologic definition of decline presented earlier.
17USDA Forest Service Proc. RMRS-P-60WWW. 2010
ThismodelisconsistentwiththedefinitionsproposedbytheForestHealthgroup,as“die-back”couldrefertothedeathofapartofanaspengenotypeexistingonasite,andthemortalityofastandorclonecouldbeconsideredtobetheresultofa“decline.”Manycriticalknowledgeneedsandquestionsposedbytheworkinggroupscouldfitwithinthisclassification.Mostgroupscalledfortheneedtoidentifyandquantifysiteandbioticfactorsassociatedwithaspenmortality.Additionally,suggestionswereofferedtoquantifyclimaticandmeteorologicalfactorsassociatedwithmortality,includingsitefactorssuchassoil,physiographicposition,aspect,andassociatevegetation. Investigation isalsoneeded into therole thatclimatechangemightplayintheaspenmortalityphenomenon,specificrelationshipsbe-tweendroughtandmoisturestress,andlongtermdiseaseandinsecteffects.Betterknowledgeoftherelationshipsbetweenchangesinfireregimesandcurrentaspenmortalitywasidentifiedasyetanothercriticalneed.
The Role of Herbivores
Severalworkinggroupsdiscussedtheneedtolearnmoreabouttheroleofherbi-voresinaspenmortalityevents.Specificneedsincludeaspen/herbivoreinteractionsandinter-dependency,whetherherbivorescanbeavectorforpathogensthatcaninfectaspen,andidentificationofcircumstanceswhereaspenneedprotectionfromherbivores.Especiallyimportantistheneedtolearnatwhatscalesandunderwhatconditionsherbivoresbecomeproblematicinaspenforests.
The Role of Aspen Roots
Theneedtolearnmoreabouttherolethataspenrootsplayinthevariousmortal-ityscenarioswasmentionedrepeatedly.Areaspenrootsreallydeadinsituationswheresproutingdoesnotoccur?Ifso,aretheydyingbeforeoraftertheoverstorytreesdie?Doesrootgraftingplayaroleinavoidingorcontributingtocompletestandmortality?
Genetics
Alackofkeygeneticknowledgewasidentified.Whatroledoesgeneticsplayinthemortalityprocess?Cansusceptiblegenotypesthatarepredisposedtomortalitybeidentified?Iscloneage(theamountoftimeagenotypehasexistedonasite)relatedtomortalityoccurrence?Aremaleorfemaleclonesmoresusceptibletomortality?
Rapid Assessment
A theme common to allworking groups, and subsequent discussions by theentiregroup,istheneedforarapidassessmentofaspenthroughouttheWestto(1)determinethecurrentstatusandprojectedtrendsofaspenforestsand(2)moreaccuratelyquantifythemagnitudeandextentofmortality.Theneedtoestimatethedurationofmortalityandprojecttheeconomicandecologiceffectsassociatedwithitisespeciallycriticaltomanagers.Identifyingstockingmetricsassociatedwithmortalitywouldbeveryuseful,especiallyifaspenlosscouldbeassociatedwithage,growthrate,stockingconditions,orotherbioticfactors.
18USDA Forest Service Proc. RMRS-P-60WWW. 2010
Recommended Actions
Althoughworkshopparticipantsidentifiedanumberofresearchandmanagementneedsthatcouldimproveourunderstandingoftheprocessesinvolvedinthecur-rentaspenmortalityevents,wesuggestthatthemostcriticalneedistodevelopacommonrapidassessmentprotocolthatcouldprovidecomparabledatathatwouldmeetbothmanagementandresearchobjectives.Withthatinmind,werecommendthefollowingactions:
• Formamulti-agencytaskforcetodevelopacommonrapidassessmentpro-tocolthatcombinesremotesensingandgroundbaseddataaboutthecurrentconditionofaspenforestsand includesmethodologyanddataneeded tomonitorandassessmortalityevents.Thetaskforce,afterconsideringspe-cificdataneedsoutlinedintheworkinggroupreports,shouldrecommendaspecificprotocoltemplatethatwillallowconsistenttreatmentapplicationandmonitoringoftheaspenresourcethroughouttheWestandprovideinforma-tiontoplanfuturemanagementandresearchactivities.Subsequentresearchhasusedsimilarprotocols;howevernouniversalonehasbeenadapted.
• Holdafuturemulti-partnerworkshopafterthe2007fieldseasontore-assessthecurrentstateofaspenresources throughout theWest.Thisworkshopshouldbeavehicle to share informationcollectedduring the2007 fieldseason, identify additional research needs, and recommendmanagementactionsbasedonthenewinformation.[Update:ASuddenAspenDecline(SAD)meetingwasheldFebruary12-13,2008,inFort.Collins,CO.ThismeetingreportedeffortsconcerningSADforthe2007fieldseason.Awidevarietyof studies and issueswerepresented anddiscussed.Anuneditedsummaryofthismeeting,compiledbyPaulRogers,canbefoundathttp://www.western-aspen-alliance.org/.Thesummarycanbefoundbyactivating“Links”ontheleftsidebarandopening“SummaryoftheSuddenAspenDeath(SAD)…2/08.”]
• Reviewstate-widewildlife and fishcomprehensiveplans andTNCeco-regionalassessmentstoidentifyaspen-relatedissues,trends,andneeds.TheForestServiceSOPAdocumentsdatabaseshouldbereviewedtoidentifyspecificsiteswherevegetationmanagementprojectsareplannedorhavebeencompleted inaspenforests.This informationwillbeveryuseful intargetingresearchandmonitoringtoascertainhowparticularmanagementactivitiesaffecttheaspenresource.
• DevelopamechanismtofacilitatecollaborationandcommunicationamongallaspenmanagersandresearchersinthewesternUnitedStates.Thiscouldtaketheformofauniversityoragencyfundedinstituteoraninformalcol-laborativeworkinggroupthatwouldserveasaclearinghouseforinforma-tionanddiscussionconcerningaspen.Anintegralpartofthisorganizationwouldbetoserveasaconduitofinformationbetweeninterestedparties,providing assistance, consulting services, and disseminating informationthroughtheinternetorotherinformaloutlets.[Update:AjointeffortbetweenRockyMountainResearchStation(RMRS)andUtahStateUniversity(USU)hasresultedintheWesternAspenAlliance(WAA)beingformed.AformalMemorandumofUnderstanding(MOU)hasbeenestablishedbetweenRMRS
19USDA Forest Service Proc. RMRS-P-60WWW. 2010
andUSUtovalidatethisAlliance.TheWAAisaconsortiumofresearchersandmanagerswhowillcoordinateandfacilitateadvancesinaspenecologyinwesternNorthAmerica.TheWAAprospectus(fig.5)andwebsite(http://www.western-aspen-alliance.org/) contain additional information on theAlliance.]
Figure 5. Prospectus for the Western Aspen Alliance (WAA).
20USDA Forest Service Proc. RMRS-P-60WWW. 2010
References
DeByle,N.V.;Winokur,R.P.,eds.1985.Aspen:EcologyandmanagementinthewesternUnitedStates.Gen.Tech.Rep.RM-119.FortCollins,CO:U.S.DepartmentofAgriculture,ForestSer-vice,RockyMountainForestandRangeExperimentStation.283p.
Manion,P.D.1981.Treediseaseconcepts.Firstedition.Prentice-Hall,EnglewoodCliffs,NewJersey.402p.
Mueggler,W.F.1989.AgedistributionandreproductionofIntermountainaspenstands.WesternJournalofAppliedForestry.4(2):41-45.
Ohms,S.R.2003.RestorationofaspenindifferentstagesofmortalityinsouthernUtah.M.S.Thesis.UtahStateUniversity,Logan,UT.99p.
Shepperd,W.D.1993.Theeffectsofharvestingactivitiesonsoilcompaction,rootdamage,andsuckeringinColoradoaspen.WesternJournalofAppliedForestry.8(2):62-66.
Shepperd,W.D.;Rogers,P.C.;Burton,D;Bartos,D.L.2006.Ecology,biodiversity,management,andrestorationofaspenintheSierraNevada.Gen.Tech.Rep.RMRS-GTR-178.FortCollins,CO:U.S.DepartmentofAgriculture,ForestService,RockyMountainResearchStation.122p.
Whitham,T.G.;Young,W.P.;Martinsen,G.D.;Gehring,C.A.;Schweitzer, J.A.;Shuster,S.M.;Wimp,G.M.;Fischer,D.G.;Bailey,J.K.;Lindroth,R.L.;Woolbright,S.;Kuskeet,C.R.2003.Communityandecosystemgenetics:Aconsequenceoftheextendedphenotype.Ecology.84(3):559-573.
Worrall,J.J.;Egeland,L.;Eager,T.;Mask,R.A.;Johnson,E.W.;Kemp,P.A.;Shepperd,W.D.2008.RapidmortalityofPopulus tremuloidesinsouthwesternColorado,USA.ForestEcologyandManagement.255:686-696.
21USDA Forest Service Proc. RMRS-P-60WWW. 2010
Appendix
List of attendees at the Aspen Summit Meeting, Salt Lake City, Utah, December 18 and 19, 2006.
Name Organization
Bill Jacobi Colorado State UniversityBrian Kurzel Colorado State ParksCarl Edminster USFS - Rocky Mountain Research StationCindy Swanson USFS - Region 1Dale Bartos USFS - Rocky Mountain Research StationDarren McAvoy Utah State UniversityDave Cleaves USFS - Rocky Mountain Research StationDavid Burton Aspen Delineation ProjectHenry Lachowski USFS - Remote Sensing Applications CenterIngrid Aguayo Colorado State Forest ServiceJack Troyer USFS - Region 4James Hoffman USFS - Region 4James Worrall USFS - Region 2Janine Powell USFS - Rocky Mountain Research StationJim Long Utah State UniversityJohn Guyon USFS - Region 4John Shaw USFS - Rocky Mountain Research Station Julia Richardson USFS - Region 4 (Humboldt-Toiyabe NF)Karen Mock Utah State UniversityKen Hehr USFS - Region 2 (San Juan NF)Laura Moffitt USFS - Region 4 MaryLou Fairweather USFS - Region 3Melissa Jenkins USFS - Region 4 (Caribou-Targhee NF)Michael Wilson USFS - Rocky Mountain Research StationMike Duncan USFS - Region 4 (Dixie NF)Mike Kuhns Utah State UniversityPhillip Kemp USFS - Region 2 (San Juan NF) (Retired)Robert Campbell USFS - Region 4 (Fishlake NF)Ron Ryel Utah State UniversitySkip Smith Colorado State UniversitySteve Ambrose USFS - Rocky Mountain Research StationSteve Solem USFS - Rocky Mountain Research StationTim Garvey USFS - Region 2 (Grand Mesa, Uncompahgre & Gunnison NF)Tom Martin USFS - Region 4Valerie Hipkins USFS - Pacific Southwest Research StationVicki Berrett USFS - Rocky Mountain Research StationWayne Shepperd USFS - Rocky Mtn. Res. Station (Retired)
22USDA Forest Service Proc. RMRS-P-60WWW. 2010
Rocky Mountain Research Station
The Rocky Mountain Research Station develops scientific information and technology to improve management, protection, and use of the forests and rangelands. Research is designed to meet the needs of the National Forest managers, Federal and State agencies, public and private organizations, academic institutions, industry, and individuals. Studies accelerate solutions to problems involving ecosystems, range, forests, water, recreation, fire, resource inventory, land reclamation, community sustainability, forest engineering technology, multiple use economics, wildlife and fish habitat, and forest insects and diseases. Studies are conducted cooperatively, and applications may be found worldwide.
Station HeadquartersRocky Mountain Research Station
240 W Prospect RoadFort Collins, CO 80526
(970) 498-1100
Research Locations
Reno, NevadaAlbuquerque, New MexicoRapid City, South Dakota
Logan, UtahOgden, UtahProvo, Utah
The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of race, color, national origin, age, disability, and where applicable, sex, marital status, familial status, parental status, religion, sexual orientation, genetic information, political beliefs, reprisal, or because all or part of an individual’s income is derived from any public assistance program. (Not all prohibited bases apply to all programs.) Persons with disabilities who require alternative means for communication of program information (Braille, large print, audiotape, etc.) should contact USDA’s TARGET Center at (202) 720-2600 (voice and TDD). To file a complaint of discrimination, write to USDA, Director, Office of Civil Rights, 1400 Independence Avenue, S.W., Washington, DC 20250-9410, or call (800) 795-3272 (voice) or (202) 720-6382 (TDD). USDA is an equal opportunity provider and employer.
Flagstaff, ArizonaFort Collins, Colorado
Boise, IdahoMoscow, Idaho
Bozeman, MontanaMissoula, Montana