Ornithological Monographs No 54 PopulationDynamicsof the California SpottedOwl (Strix occidentalis occidentalis):A Meta-Analysis ALAN I3.FRANKLIN,R J GUTII•RREZ, JAMESD NICHOLS,MARKE $EAMANS, GARYC WHITE,GUTHRIE$ ZIMMERMAN,JAMES E HINES, THOMASE MUNTON,WILLIAM $ LAHAYE,JENNIFER A 13LAKESLEY, GEORGE N $TEGER, B^RR¾ R NOON,DANIELW H SH^W,JOHNJ.KEANE, TRENT L MCDONALD, AND SUSANBRITTING PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION POPULATION CALIFORNIA DYNAMICS SPOTTED OF THE OWL (STRIX OCCIDENTALIS OCCIDENTALIS)' A META-ANALYSIS ORNITHOLOGICAL MONOGRAPHS Editor:JohnFaaborg 224 Tucker Hall Divisionof BiologicalSciences Universityof Missouri Columbia, MO 65211 ManagingEditor:BradleyR Plummer Proof Editors: Mark C Penrose,Richard D Earles AOU Publications Office 622 ScienceEngineering Departmentof BiologicalSciences Universityof Arkansas Fayetteville, Arkansas72701 Ornithological Monographs, publishedby the American Ornithologists'Union, has beenestablished for majorpaperstoo longfor inclusionin the Union'sjournal,TheAuk Publicationhasbeenmadepossiblethroughthe generosityof the late Mrs Carll Tucker and Marcia BradyTuckerFoundation,Inc Copiesof Ornithological Monographs may be orderedfrom ButeoBooks,3130 Laurel Road,Shipman,VA 22971.Priceof OrnithologicalMonographs54:$10.00($9.00for AOU members) Add $4.00for handlingand shippingchargesin U.S.,and $5.00for all other countries.Make checkspayableto ButeoBooks Corresponding authorof thisissue,R J.Gutierrez The costof thisOrnithological Monographwasdefrayedby theU.S.Departmentof Agriculture, Forest Service Library of CongressControlNumber 2004100002 Printedby CadmusCommunications, Ephrata,PA 17522 Issued30 January2004 Ornithological Monographs, No 54 vi + 54 pp Copyright¸ by the AmericanOrnithologists'Union, 2004 ISBN: 0-943610-00-1 Cover:SpottedOwl (Strixoccidentalis), ink sketchby Viktor Bakhtin POPULATION DYNAMICS CALIFORNIA SPOTTED OF THE OWL (STRIX OCCIDENTALIS OCCIDENTALIS): A META-ANALYSIS BY: ALAN B FRANKLIN, 1'2R J GUTII•RREZ, 3JAMESD NICHOLS, MARKE SEAMANS, GARYC WHITE,2 GUTHRIES ZIMMERMAN, 3JAMESE HINES,4 THOMASE MUNTON,5 WILLIAM S LAHAYE,3JENNIFER A BLAKESLEY• 2GEORGE N STEGER, BARRYR NOON,2DANIELW H SHAW, 5JOHNJ KEANE,6TRENTL MCDONALD, AND SUSAN BRITTING •Colorado Cooperative FishandWildlifeUnit, Colorado StateUniversity, FortCollins,Colorado 80523,USA; 2Department ofFishery andWildlifeBiology, Colorado StateUniversity, FortCollins,Colorado 80523,USA; 3Department ofFisheries, Wildlife,andConservation Biology, University ofMinnesota, St.Paul,Minnesota 55108,USA; 4U.S.Geological Survey, Patuxent WildlifeResearch Center, 11510American HollyDrive,Laurel,Maryland20708,USA, sU.S.Department ofAgriculture,ForestService, PacificSouthwest Research Station,2081EastSierraAvenue, Fresno,California93710,USA; 6U.S.Department ofAgriculture,ForestService,SierraNevadaResearch Center,PacificSouthwest Research Station, 2121Second Street,SuiteA101,Davis,California95616,USA; 7WEST,Inc.,2003CentralAvenue,Cheyenne, Wyoming82001,USA;and 8P.O.Box377,Coloma, California 95613,USA ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 2004 NO 54 D.C UNION TABLE ABSTRACT STUDY OF CONTENTS AREAS LASSEN STUDY AREA ELDORADO STUDY AREA SIERRA STUDY AREA SEQUOIAAND KINGS CANYON NATIONAL PARKSSTUDYAREA SAN BERNARDINO STUDY AREA METHODS FIELD METHODS 10 10 I0 Surveys 11 Estimation ofreproductive effort I1 Capture, banding, sexandageidentification, andresighting ofowls 11 Pre-analysis datascreening 12 Meta-analysis workshop format DATA ANALYSIS 12 12 Changes in analytical methodology fromprevious Spotted Owlstudies 13 Estimating adultsurvival 14 Estimating fecundity 16 Estimating ratesofpopulation change Comparison ofSierraandSequoia andKingsCanyon national parksstudyareas RESULTS 19 19 ADULT SURVIVAL 19 FECUNDITY 21 RATES OF POPULATION CHANGE 23 27 Meta-analysis across studyareas COMPARISON OFSIERRAAND SEQUOIAAND KINGSCANYONNATIONAL PARKSSTUDYAREAS DISCUSSION 30 30 GENERAL INFERENCES 31 31 Apparentsurvival 32 Fecundity 33 Population trend STUDY-AREA-SPECIFIC INFERENCES 35 35 Lassen studyarea 36 Eldorado studyarea 37 SierraandSequoia andKingsCanyonnationalparksstudyareas 39 SanBernardino studyarea CONCLUSION AND RECOMMENDATIONS ACKNOWLEDGMENTS LITERATURE APPENDICES CITED 40 41 42 48 From the Editor With Ornithological Monographs #54,theAmericanOrnithologists' Union implementsa new philosophyin the productionof its monographseries.Sincethe seriesbeganin 1964,Ornithological Monographs havebeenpublishedsporadically,primarily to presentarticlesthat were too large to appearin TheAuk.Someof thosemonographs havebeenenormous(over1,000pages),although manywerein the50-100pagerange.Theyweresoldasseparateissues,with pressrunsof at most a few thousandcopies Thisandsubsequent monographs will be providedto all AOU memberson a regularbasis,packagedwith TheAuk.Thereare many ornithologicalresearcheffortsthat theAOU and I feel deserve to be publishedin one setting,without beingseparatedinto two or threemanuscriptsthat appear in differentjournals.If youhavea dissertation, majorresearchproject,or evena smallsymposium longerthanthe50pagesallowedby TheAuk,we hopeyouwill considerpublishingin Ornithological Monographs Ornithological Monographs is opento all aspectsof ornithology.All thatwe askis that theresearch involvegoodscience,havereasonablybroadornithologicalinterest,and cantruly justify the need for monographic treatment.Financialsupportfor publicationis not a requirement,althoughit can certainlyhelp the AOU and may be necessary for largervolumes We begin the "new" Ornithological Monographs with an analysisof the demographyof the CaliforniaSpottedOwl Althoughthe SpottedOwl hasbecomethe focalspeciesfor both sidesin argumentsaboutforestrypractices in thewesternUnitedStates,mostof thenationalpublicityhas involvedthe NorthernSpottedOwl of northwestern California,Oregon,andWashington Similar controversynow surroundsthe CaliforniaSpottedOwl An attemptto haveit listedas an endangeredspeciesendedup in the courts,which forcedthe U.S Fishand Wildlife Serviceto conducta statusreview.The U.S Fishand Wildlife Servicechosenot to list the owl, in part becausethe U.S ForestServicehad developeda management plan (theSierraFramework)designedto protectthe owl and many other resourcesof the SierraNevada.However,on the day that the U.S Fish and Wildlife Service announced that it would not list the owl, the U.S Forest Service announced its desireto "modify"the SierraFramework.The modifiedframeworkwill be completedin early2004, and it is unclearat thistime whetherthiswill be potentiallyharmful to the owl We will undoubtedly hear more aboutthis situationin the future As with mostthreatenedor endangeredspecies, we need soliddata on demographicpatterns acrossthe species'range.This monographprovidessuchdata,combiningmodernmethodssuch as meta-analysis with sophisticated capture-recapture modelsacrossa varietyof Californiasites The topicis criticalfor conservationpurposes,and the approachwill introducereadersto the stateof-the-artin the conductingdemographicstudies.With an interestingand importantbird, 16 wellqualifiedauthors,and pioneeringmethodsof analysis,we believethisstudysetsa high standard for the new Ornithological Monographs Any scientific editorwill admitthatoutsidereviewis criticalto thescientific publishingprocess Findingreviewersfor the long manuscripts that are potentialOrnithological Monographs will per- hapsbe a challenge, but we hopereaderswill be asexcitedabouttheconceptaswe areandwill be willing to volunteertimewhennecessary Forthismonograph, JeffreyR Walters,EvanCooch,and KennethH Pollockof the AOU Conservation Committeedid an exceptionally detailedreviewof an earlydraft.KatieDugger,JeffreyR Walters,and a reviewerwho wishesto remainanonymous madecommentson what becamethe final product.We thankthesereviewersfor the considerable time theycontributedtowardmakingthisnew monographa high-quality,interesting,andimportant pieceof science.We alsowant to thank KimberlySmith,Brad Plummer,Mark Penrose,and RichardEarlesof theAOU Publications Officefor helpingtrainthisnew editorin theart of producing scientificpublications JohnFaaborg Ornithological Monographs Volume (2004),No 54, 1-54 POPULATION DYNAMICS OF THE CALIFORNIA SPOTTED OWL (STRIX OCCIDENTALISOCCIDENTALIS):A META-ANALYSIS ALANB FRANKLIN, 1'2'9 R J GUTI12RREZ, 3'1ø JAMES D NICHOLS, 4MARKE SEAMANS, GARYC WHITE,2 GUTHRIES ZIMMERMAN, 3JAMES E HINES,4THOMAS E MUNTON, WILLIAMS LAHAYE, 3'nJENNIFER g BLAKESLEY• 2GEORGE N STEGER• BARRY R NOON,2DANIELW H SHAW, 5JOHNJ KEANE, 6TRENTL MCDONALD, AND SUSAN BRITTING •Colorado Cooperative FishandWildlife Unit,Colorado StateUniversity, FortCollins, Colorado 80523,USA; 2Department ofFishery andWildlife Biology, Colorado StateUniversity, FortCollins, Colorado 80523,USA; 3Department ofFisheries, Wildlife, andConservation Biology, University ofMinnesota, St.Paul,Minnesota 55108,USA, 4U.S.Geological Survey, Patuxent Wildlife Research Center, 11510American HollyDrive,Laurel, Maryland 20708,USA, sU.S.Department ofAgriculture, Forest Service, Pacific Southwest Research Station, 2081EastSierraAvenue, Fresno,California93710,USA; 6U.S.Department ofAgriculture, Forest Service, SierraNevada Research Center, Pacific Southwest Research Station, 2121Second Street,SuiteA101,Davis,California95616,USA; 7WEST, Inc.,2003CentralAvenue,Cheyenne, Wyoming 82001,USA;and 8P.O.Box377,Coloma,California95613,USA ABSTRACT We conducted a meta-anaiysis to providea currentassessment of thepopulation characteristics ofCalifornia Spotted Owls(Strixoccidentalis occidentalis) resident onfourstudyareasin theSierraNevadaandonestudyareain southern California Ourmeta-analysis followed rigorousa priorianalysis protocols, whichwe derivedthroughextensive discussion duringa week-long analysis workshop Because thereis greatinterestin theowl'spopulation status,we usedstate-of-the-art analytical methods to obtainresultsasprecise aspossible Our meta-analysis includeddatafrom five Californiastudyareaslocatedon the Lassen NationalForest(1990-2000),EldoradoNationalForest(1986-2000),SierraNationalForest (1990-2000),Sequoiaand Kings Canyonnationalparks (1990-2000),and San Bernardino NationalForest(1987-1998) Four of the five studyareasspannedthe lengthof the Sierra Nevada,whereasthefifthstudyareaencompassed theSanBernardino Mountainsin southern California.Studyareasrangedin sizefrom343km2(Sequoia andKingsCanyon)to 2,200km2 (Lassen) All studiesweredesigned to usecapture-recapture methodsandanalysis We used survivalin a meta-analysis because fieldmethodswerevery similaramongstudies.However, we did not usereproduction in a meta-analysis because it wasnot clearif variationamong individualstudy-area protocols usedto assess reproductive outputof owlswouldconfound results.Thus,we analyzedfecundityonlyby individualstudyarea.Weexamined population trendusingthereparameterized Jolly-Seber capture-recapture estimator(At) We did not estimatejuvenilesurvivalratesbecause of estimation problemsandpotential biasbecause of juvenileemigrationfrom studyareas.We usedmark-recapture estimators underaninformation theoretic frameworkto assess apparentsurvivalratesof adultowls.The pooled estimate foradultapparent survival forthefivestudyareas was0.833,whichwaslower thanpooled adultsurvival rates(0.850) from15Northern ^Spotted Owl(S.o.caurina) studies Estimatesof survival from the best model on the Lassen( •b= 0.829,95% confidenceintervals [CI]= 0.798to 0.857),Eldorado( •b= 0.8•15, 95%CI = 0.772to 0.851),Sierra( •b= 0.818,95%CI = 0.781to 0.850),and SanBernardino( •b= 0.813,95%C! = 0.782to 0.841)were not different However,theSequoia andKingsCanyonpopulation hada highersurvivalrate( •b= 0.877,95% CI = 0.842to 0.905)than the otherstudyareas.Managementhistoryand foreststructure(e.g presence of giantsequoia[Sequoiadendron giganteum]) ontheSequoia andKingsCanyonstudy areadifferedfromall otherstudyareas.Thereappearsto belittleor no evidencefor temporal variationin adultapparentsurvivalon anyof the studyareas Althoughwe didnotdirectlycomparefecundity, estimates werehighlyvariableamongyears withinall studyareas(CV of temporalprocess variation= 0.672-0.817) Estimates forfecundity 9E-mail: alanf@cnr colostate.edu høE-mail: gutie012@tc.umn.edu nPresentaddress:P.O.Box523,BigBearCity,California92314,USA ORNITHOLOGICAL MONOGRAPHS NO 54 amongthestudypopulations wereLassen (• = 0.336,SE= 0.083),Eldorado (• = 0.409,SE= 0.087),Sierra( • = 0.284,SE= 0.073),Sequoia andKingsCanyon( • = 0.289,SE= 0.074),andSan Bernardino ( • = 0.362,SE= 0.038).Duringmostyears,theSierraNevadapopulations showed eithermoderateor poor fecrmdity.However,1992appearedto be an exceptionalreproductive year for owls in the SierraNevada In contrast,the San Bernardinopopulationhad less variablereproduction(CV of temporalprocessvariation= 0.217),but experienced neitherthe exceptionalreproductionof 1992nor the extremelypoor yearsthat characterizedall of the SierraNevadas•udyareas.Becausefecrmditymay be influencedby weatherpatterns,it was possiblethat the differentweatherpatternsbetweensouthernCaliforniaand the SierraNevada accotinted for that difference Except forEldorado, allestimates for_A twereS2,but truncateJ, S1, and S2 histories) across fivestudyareasusingQAICc modelselection criteriain programMARK {½(g* t * s)p(g* t * s)}globalmodel The following ½ models are to be estimatedagainstthe nine p modelsin step (4) below: ẵ(g* t * s) ẵ(*t+s) ẵ(g* t) 27 Models Includea sexeffect,if necessary, from the above27modelswith thebestp modelof the 27 and run the followingadditional models on ½(X+ t) ½(x* 13 ½(g* TT) Total models = 43 Goodness of fit determined with RELEASE iii Age-specificfecundity (b) by study area analyzedin PROCMIXED in SAS Fixed effects a Female age (S1, S2, A) as a fixed effect b Fixed TT model c Fixed Tmodel d Fixedintercept-only model(dot) e Fixedeven-odd model (denotedEO) f Fixed even-odd model with a linear trend (13 Random effects with ALL of the above fixed effects: a Territories as a random effect b Year as a random effect Structurevarianceasproportionalerror distribution, LOCAL= EXP(femaleage, year, or even-odd) for the on-diagonal elements Candidate variance structures for off-diagonalelementsare compound symmetric(CS) or autoregressive with lag of (AR1).Choicebetweenthosetwo modelswill be madeusingAIC model selection model with REML Models run will be: Age + TT Age * TT Age + T Age * T Age Age + EO Age * EO Age + EO + T in an AGE + T POPULATION DYNAMICS OF THE CALIFORNIA SPOTTED OWL Age * T + EO Interceptonly Model selectionusingAIC Tenmodelsto run for eachstudyarea iv )* from Pradel model Truncatedata sets to first year when "density" study area adequately surveyed Selecteither{)*(t)•(t) p(t)} or {)*(s* t) •(s * t) p(s* t)} basedonAICc Eliminate)*• (confounded),)*2 (may be biased), and )*• (confounded)from analysis Do variancecomponentson bestmodel abovewith the followingstructures,and 51 i Sierra study areas should have higher temporal variation in survival and reproductionthan SAB becauseof weather patterns ii Two northern Sierra study areas (LAS and ELD) should have different temporal variationin survivaland reproductionthan the two southernSierra study areas (SIE and SKC) Not sure about the SAB b Temporal process variation (•2temporal) i Within eachstudy area ^2 Estimatec•temporal for adult apparent survival(•^) fromrandomeffectsmeans selectthebestof thosethreewithAICc: model in MARK using estimatesfrom model {•(a * s * t) p(a * s * t)} if there is a sexeffectand {•(a * t) p(a * t)} if there is a no sex effect T ^2 forjuvenile apparent Estimatec• temporal b TT c dot survival (•i) fromrandomeffects means Total of five modelsrun for eachstudy area Goodness of fit determined with no sex effect RELEASE ^2 v )* from Leslie matrix Estimate c•temporal for)*R•S fromrandom Use4-ageclassprojection matricesfor all studyareas Estimate age-specific• and b (and their standard errors) from best model resulting from age-specific survival modeling in MARK and from best model in fecundityanalysisfrom PROC MIXED, respectively,as inputs for the Calculatejuvenile(Ej•x=•)and adult (E^•x=•) emigrationrates necessaryto achievea stationarypopulation()*= 1.0) Calculatejuvenile(•j•x•) and adult (c•^•b•)survival rates necessaryto achievea stationarypopulation()*= 1.0) EstimateSE(Musingdeltamethod If unableto estimatejuvenilesurvivalfor a givenstudyarea,thenwon't attemptto )* b Not to be estimated: ii Number of owls detectedper unit effort (assumingN worksabove); iii Juvenileemigration; iv Year-specificN, from Jolly-Sebermodel have used same data used to estimate )*Rid v Changein age of new recruitsthrough time to evaluatechangein agestructureof floaterpopulation Estimate temporal and spatial variation in demographicparametersfor California Spotted Owls acrossfive studyareas a Predictions: Estimate •2temporal foradultfecundity (b^)fromintercepts-only modelin PROC MIXED in SAS Donotestimate •2temporal forC•s•, C•s2, bs•,or bs2 because of knownsamplesize limitations Within eachstudy area a Estimate •2spatial for b^basedon territories(owl sites) Amongstudyareas a Estimate •2spatial for adultapparent survival (•A) from mean adult apparent survival (•A)computed from each study area (see 4.a.v.(2)) and using method-of-momentsvariance componentsoutlinedin Burnhamet al (1987) i Percentterritory occupancy; Would effects means model in MARK using estimatesfrom model {)*(t),•(t), p(t)} c Spatial process variation (•2spatial) matrix estimate model in MARK using estimatesfrom model{•(a * s * t) p(a* s * t)} if thereis a sexeffectand {•(a * t) p(a * t)} if there is i Estimate temporal covariance from modelg + t in meta-analysis of apparentsurvival b Estimate •2spatial forjuvenile apparent survival (•) from mean juvenile apparent survival(•l) computed from eachstudy area(see4.a.v.2)and using method-of-momentsvariance componentsoutlinedin Burnhamet al (1987) c Estimate •2spatial for)*Ris from•IS computedfrom eachstudy area (see 4.a.iv)andusingmethod-of-moments variance components outlined in 52 ORNITHOLOGICAL MONOGRAPHS NO 54 Burnham et al (1987) d.kpMrepresents the asymptotic changein the i Estimate temporal covariance from modelg + t in meta-analysis of apparentsurvival d Donotestimate •2spatial for•s•,•s2'bs•, or bs2becauseof known samplesize limitations and not estimate for b^becauseof differences in protocols among study areas However, femalepopulationsizegivena specificsetof apparentsurvivalandfecundityrates i Doesnot includeimmigration ii Represents asymptoticconditionsfor fixed valuesof apparentsurvivaland fecundity (i.e emigration[part of apparentsurvival] is a functionof studyareasizeand edgeto arearatio,sothatkpMis a functionof study area characteristics) comparetrends in b^ acrossstudy areas Interpretationand reportingof results, a Apparent survival estimates: i, Changesin • overtimerepresentchanges in emigration,death,or both ii The emigration component in juvenile apparentsurvivalis greaterthan for other age-classes, iii Bias from resighting heterogeneity in estimatesof apparent survival for S1, S2, and A will be small iv Inferencesaboutapparentsurvivalestimates apply only to the markedpopulation b Fecundityestimates: i May be positively biased because of methodology used to determine reproductiveoutput (number of fledged young) ii Pointestimatesare not comparableamong study areas because of differencesin protocolused on differentstudy areas.An exceptionis comparisons betweenthe SIE and SKC study areas iii Differencesin fecundity estimateswithin each study area are comparablebecause bias due to protocol should be similar amongyearswithina particularstudyarea Thus, trendsin fecunditywill be examined within study areas c kRls'represents change in number of territorial owls i • providesinformationon probabilityof changeover a specifiedtime period, given • underconditions of study ii Change basedon)•Rls canbe dueto local birth, immigration, death, emigration,or both iii Assume50:50sexratio of juveniles e General inferences, i Inferences are confined to: Within the study areas Within the study period The territorialpopulationof owls?This interpretationneedsto be reconsidered carefully Amongstudyareasfor meta-analyses f Reportingresults i Lead responsibilityfor compiling final reportwill beR.J.Gutierrez ii Draft reportwill be written (not submitted) by end of October2001 All participantswill be authors, All participantswill review final report before submission a Deadline for participant reviews; report will be submittedregardless if participant reviews not received within deadlineto be establishedby R J Gutierrez Final report will be reviewed (prior to submission)by two outside reviewers familiar with analyticalmethodsused in report (may be vetoedby U.S Forest Service) a Outside reviewers will be paid to ensuretimely review Internal editor will arbitrate reviews and contentof finalreport a Gary White selected iii Finalreportwill eventuallybe publishedin a peer-reviewedoutlet Wildlife Monographssuggested outlet Order of authorshipand inclusionof authorswill be determinedby group POPULATION DYNAMICS OF THE CALIFORNIA SPOTTED OWL APPENDIX 53 believethat SpottedOwl populationsare similar to many populationsof passerinebirds (DeSante1995), FURTHERDISCUSSIONOFPROJECTIONMATRIX insects(Connor et al 1983), small mammals (Nichols POPULATION MODELS and Pollock1990),and marine fish (Roughgardenet al 1985,Armsworth 2002) Some of the recommendations to use 3.pMsugDuringthe workshop,the groupdebatedthe utility of developingprojectionmatricesand computing gestthat that metricmay provideinsightsaboutthe asymptotic 3.PM Subsequent reviewsof thedraftwork- relevanceof movementto populationdynamicsthat shopreportalsoraisedthat issue,sowe presenthere cannotbe obtainedusing3.r We not believethat ourreasoning fornotestimating 3.PM morecompletely that is true Instead,we believethat the key issueis A central theme of our internal discussions and the re- not one of which to use, but of how to estimate rel- viewswasthat 3.testimatedusingcapture-recapture evant quantities.Inferencesaboutmovementrequire data reflectschangesin numbersof birds resulting the abilityto separategainsand losses in the estimafrom all sourcesof lossfrom andgainto the studyar- tion process.Indeed, at the workshop,we worked for decomposing recruitmentinto in eas.Althoughwebelievethat3.tis relevantto popula- on expressions tion changeat the scaleof the individualstudyareas, situ reproductionand immigration componentsuswe recognizethat for somequestions,it would be ing a multi-ageversionof the approachsuggested valuableto separatelossesfromdeathandemigration by Nicholsand Pollock(1990).But althoughthe apand gains from in situ recruitment and immigration, propriateestimators were developed,that work was because SpottedOwl populations at thescaleof study not completedat the workshop,sowe haveno such appearto be geographically open.Thus,dynamicsat estimates at this time Thus,parameters thatwerewell estimatedandthat thescaleof studyareasaredeterminedat leastin part by contributionsfrom other areas.Here, we discuss were suitedto incorporationinto projectionmatrix reproductive ratesandrates geographicopennessand the related source-sink modelswereage-specific dichotomyand ask whetherdevelopmentof projec- of lossthat includebothdeathandemigration.As nottion matrix modelsmight be usefulin increasingour ed hereandin otherSpottedOwl reports,that asymmetry in the treatment of movement would lead to understanding of thoseopensystems During both internal workshopdiscussionand projectionmatrixresultsthat wereof littleuse.Thus, externalreviews,it wasnotedthatcomputation of 3.t previouseffortshaveadjustedor correctedestimates was inadequateto determinewhetheran areawas a of juvenilesurvivalto removepermanentemigration "source"or a "sink."It is fairly clearthat the studied as a sourceof losson the basisof poor estimatesof SpottedOwl populationsdo not correspondto the juveniledispersal.The rationalewasthat removalof strict source-sinkmodel systemof Pulliam (1988) movementfromprojectionmatrixentrieswouldyield However, we understandthat most ecologistsno lon- inferences about3.thatwerebasedon only reproducger view thosetermsin a strictmanner,and that the tion and mortality,thusprovidinginferencesabout woulddeclineor increaseif term"source"hascometo meanan areathatsupplies whetherthe populations recruitsto other locations.Similarly,we believe that they were geographically closed.Projectionmatrices many ecologistsview a "sink" as an area in which usedpreviouslyfor SpottedOwl workwerebasedon modeling,assumingthat all surpopulationsizeis not maintainedstrictlyby recruit- closed-population mentof locallyproducedyoung.Webelievethateven viving individual'sproducedin the populationexhibthoserelaxed definitionsreflect a conceptualframe- ited thestage-specific survivalandreproductive rates workthatmaynotbeespecially usefulwhenconsider- of the area So even when we "correct" estimates of ingopensystems suchasthoseof SpottedOwIsat the apparentjuvenilesurvivalto removethe movement spatialscaleof study.Indeed,we believethat at least component,the projectionmatrix is assigningsurratesof thestudyareato fourof thefiveSpottedOwl studyareasaresources in vivalratesandreproductive the sensethat manyjuvenileowls emigrateand likely recruitto thebreedingpopulationelsewhere.We also recognizethat thosefour studyareasrepresentsinks in the sensethat many if not mostbirds recruitedto thebreedingpopulationcomefromelsewhereandare the survivingjuveniIesin subsequent years.However, the reaIity of the modeledsystemis that juvenilesare thoughttomoveelsewhere andto experience thevital ratesof thepopulations intowhichtheyrecruit.Thus, we believethat closed-population projectionmatrix not producedon the study areas.Thus,the source- resultsaremuchmoreof an abstractionof the dynamsink dichotomydoesnot seemto be especiallyuseful icsof opensystemsthanis generallyrealized The above discussion is not intended as a critifor describingthat system.We suggestthat Spotted Owl populations,asdefinedby the scaleof study,are cismof the use of traditionalprojectionmatricesin betterviewedasopen-recruitment systemsin whicha generalbut simplyarguesthatwe not expectthem substantialfractionof the recruitmentto the breeding to yield useful inferencesfor geographically open One of the reportreviewersthussuggested populationis by birds that are not producedon the systems studyarea.In that senseof geographic openness, we that we modifythe projectionmatricesto incorporate 54 ORNITHOLOGICAL MONOGRAPHS NO 54 immigration (e.g seeCoochet al 2001).As noted, we In summary,we agree with reviewersthat more did not havereliableestimatesfor suchincorporation, detailed inferences about movement would be valuand matrix asymptotics(e.g.projected)•) would de- able In particular, the ability to decomposegains pend heavily on the magnitudeof the immigration.A and lossesto studypopulationshaspotentialto yield more satisfyingapproachwould be to use multi-site increasedunderstandingof thosesystems.However, projectionmatrices(e.g Schoen1988,Lebreton1996) we disagreewith reviewersthat that problem can to includenot only the dynamicsof the studypopu- be dealt with via modelingwith existingestimates, lationsbut alsoof the population(s)with which they in particularvia computationof )•PMas in previous are connected via movement Still another alternative SpottedOwl reports.If we did chooseto model the study systems, we would select different model structuresthat weremoreappropriatefor thoseopen systems.More importantly,we view that problemof insightsaboutmovementasfundamentallya problem in estimationrather than modeling We seeno reason to expectmodel-basedasymptoticsfrom projection now in the absence of estimates of the relevant movematricesof poorly estimated(even guessed)vital mentparameters andpossibly vital ratesof connected ratesto yield reliableinsights.Our focusshouldthus be on estimationof movement-relatedparameters populations is to use open-recruitmentmodels similar to those developedfor marine systems(e.g Roughgardenet al 1985).The centralpoint here is that we have consideredalternativemodeling approachesand have someideasabouthow to proceed,but we believethat it makeslittle senseto implementthoseapproaches ORNITHOLOGICAL MONOGRAPHS (Continued frombackcover) No 35 No 37 Ecogeographical Variationin Size and Proportionsof Song Sparrows(Melospiza melodia) J.W Aldrich.x + 134pp 1984.$15.00($12.00) AvianMonogamy P A Gowaty and D W Mock, Eds.vi + 121 pp 1985.$15.00 ($12.00) No 38 No 39 No 40 No 41 An Analysisof Physical, Physiological, andOpticalAspects of AvianColoration with Emphasis onWood-Warblers E H Burtt,Jr.x + 122pp 1986.$15.00($12.50) TheLingualApparatusof theAfricanGreyParrot,PsittacuserithacusLinne (Aves: Psittacidae): Description andTheoretical Mechanical Analysis D G Homberger.xii + 236pp 1986.$30.00 Patterns andEvolutionary Significance of Geographic Variationin theSchistacea Group of theFoxSparrow (Passerella iliaca).R M Zink viii + 119pp 1986.$15.00 Hindlimb Myology and Evolutionof Old World SuboscinePasserineBirds (Acanthisittidae, Pittidae,Philepittidae, Eurylaimidae) R J Raikow viii + 81 pp 1987 $15.00 No 42 Speciation andGeographic Variationin Black-tailed Gnatcatchers J L Atwood vii + 74 pp 1988.$10.00 No 43 No 44 Nos 45 A Distributional Surveyof theBirdsof theMexicanStateofOaxaca L C Binford.viii + 418 pp 1989.$20.00 Recent Advances in theStudyofNeogene FossilBirds:I TheBirdsof theLateMioceneEarlyPliocene BigSandyFormation, MohaveCounty, Arizona(K J.Bichart);II Fossil BirdsoftheSanDiegoFormation, LatePliocene, Blancan, SanDiegoCounty,California (R M Chandler).vi + 161pp 1990.$20.00 & 46 Descriptions of Thirty-twoNew Species of Birdsfrom theHawaiianIslands:Part I Non-Passeriformes (S L Olsonand H F James),88 pp.; PartII Passeriformes (H F Jamesand S L Olson),88 pp 1991.Boundtogether(notavailableseparately) $25.00($22.50) No 47 No 48 No 49 No 50 No 51 No 52 No 53 Parent-Offspring ConflictandIts Resolution in theEuropean Starling.E Litovichand H W Power.71 pp 1992.$15.00($12.00) Studies in Neotropical Ornithology HonoringTedParker.J V RemsenJr.,Ed xiv + 918pp 1997.$49.95($39.95) AvianReproductive Tactics: FemaleandMale Perspectives P G Parker and N T Burley,Eds.v + 195pp 1998.$20.00($16.00) AvianCommunity, Climate,andSea-Level Changes in thePlio-Pleistocene oftheFlorida Peninsula S.D Emslie.iii + 113pp 1998.$20.00($16.00) A Descriptiveand Phylogenetic Analysisof Plumulaceous FeatherCharacters in Charadriiformes C J.Dove.iii + 163pp 2000.$19.95($15.96) Ornithology ofSabah: History,Gazetteer, Annotated Checklist, andBibliography F H Sheldon,R G Moyle,and J.Kennard.vi + 285pp 2001.$25.00($22.50) Evolution of Flightlessness in Rails (Gruiformes:Rallidae): Phylogenetic, Ecomorphological, and Ontogenetic Perspectives B.C Livezey x + 654 pp 2003 $10.00($9.00) Order from: Buteo Books, 3130 Laurel Road, Shipman, VA 22971, 1-800-722-2460;E-mail allen@buteobooks.com; or www.buteobooks.com Pricesin parentheses arefor AOU members For a completelist of Ornithological Monographsincludingbothin-printandout-of-printbooks,please visittheAmericanOrnithological Unionwebsiteat www.aou.org ORNITHOLOGICAL No No MONOGRAPHS TheBirdsofKentucky R M Mengel.1965.$25.00 Adaptations for Locomotion and Feedingin the Anhingaand the Double-crested Cormorant.O T O•vre 1967 $10.00 No No 10 No 11 No 12 A Distributional SurveyoftheBirdsofHonduras B L Monroe,Jr.1968.$25.00 TheBehavior ofSpotted Antbirds.E O Willis 1972.$10.00 Behavior, Mimetic Songsand SongDialects,and Relationships of the Parasitic Indigobirds (Vidua)ofAfrica.R B Payne.1973.$10.00 Intra-island Variationin theMascarene White-eye Zosterops borbonica F B Gill 1973 $10.00 No 13 Evolutionary Trendsin theNeotropical Ovenbirds andWoodhewers A Feduccia.1973 $10.00 No 14 No 15 A Symposium on theHouseSparrow(Passer domesticus) andEuropean TreeSparrow (P montanus) in NorthAmerica.S.C Kendeigh,Ed 1973.$10.00 Functional AnatomyandAdaptiveEvolution of theFeeding Apparatus in theHawaiian Honeycreeper GenusLoxops(Drepanididae) L P Richards.and W J Bock.1973 $10.00 No 16 TheRed-tailedTropicbird onKureAtoll R R Fleet.1974.$6.00 No 17 ComparativeBehaviorof the American Avocet and the Black-necked Stilt (Recurvirostridae).R B Hamilton 1975 $10.00 No 18 Breeding Biology andBehavior of theOldsquaw (Clangulahyemalis L.) R M Alison No 19 Bird Populations of AspenForests in WesternNorthAmerica.J A.D Flack.1976 1975 $6.00 $1o.oo No 21 SocialOrganizationand Behaviorof the Acorn Woodpecker in Central Coastal California M H MacRobertsand B R MacRoberts.1976.$10.00 No 22 Maintenance Behavior and Communication in the Brown Pelican R W Schreiber 1977 $6.00 No 23 No 24 No 25 No 26 Species Relationships in theAvianGenusAimophila L L Wolf 1977.$12.00 LandBird Communities of GrandBahama Island:TheStructureandDynamics of an Avifauna.J.T Emlen.1977.$10.00 Systematics ofSmaller AsianNightBirdsBased onVoice J.T Marshall.1978.$10.00 Ecology andBehavior of thePrairieWarbler,Dendroica discolor V Nolan, Jr 1978 $45.00 No 27 Ecology andEvolution ofLekMatingBehavior in theLong-tailed HermitHummingbird F G Stiles and L L Wolf 1979 $10.00 No 28 No 29 The ForagingBehaviorof MountainBluebirdswith Emphasis on SexualForaging Differences H W Power.1980.$10.00 The Molt of ScrubJaysand Blue Jaysin Florida.G T Bancroft and G E Woolfenden 1982 $10.00 No 30 AvianIncubation: EggTemperature, NestHumidity,andBehavioral Thermoregulation in a Hot Environment G S Grant 1982 $10.00 No 31 No 32 No 33 No 34 TheNativeForestBirdsofGuam.J.M Jenkins.1983.$15.00 TheMarineEcology ofBirdsin theRossSea,Antarctica D G Ainley,E F O'Connor andR F Boekelheide x + 97 pp 1984.$15.00 SexualSelection, LekandArenaBehavior, andSexualSizeDimorphism in Birds.R B Payne.viii + 52 pp 1984.$15.00 Pattern,Mechanism, andAdaptiveSignificance of Territoriality in HerringGulls(Larus argentatus) J.Burger.xii + 92 pp 1984.$12.50 (Continued on inside back cover) ... Editor With Ornithological Monographs #54, theAmericanOrnithologists' Union implementsa new philosophyin the productionof its monographseries.Sincethe seriesbeganin 1964 ,Ornithological Monographs. .. BradyTuckerFoundation,Inc Copiesof Ornithological Monographs may be orderedfrom ButeoBooks,3130 Laurel Road,Shipman,VA 22971.Priceof OrnithologicalMonographs54:$10.00($9.00for AOU members) Add... considerpublishingin Ornithological Monographs Ornithological Monographs is opento all aspectsof ornithology.All thatwe askis that theresearch involvegoodscience,havereasonablybroadornithologicalinterest,and