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 Front Cover
 Title Page
 Disclaimer
 Preface
 Abstract
 Table of Contents
 List of maps
 List of Tables
 Abbreviations used in text
 Purpose of report and study...
 Methods
 Arrangement and contents of species...
 Oil pollution and marine birds...
 Recommendations for future...
 Acknowledgement
 Species accounts
 Literature cited
 Back Matter
 Back Cover















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.....t .. !lcl.. I- .IIClDTMARINEBIltDS OFTHE SOUTIIEAS.I'ERNUNITEDSIi\n8 AND GULF OF MEXICO Partm CIIARADRIIroRMatJhmIlsManagementservtceFlsh and WIdIIeservtce

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PURPOSEOFREPORTThepurposeofthisreportistosummarizethestatusof'22 coastal andmarinecharadriiformspeciesinthesoutheasternUnitedStatesandtoexplorethepotentialeffectsonthesespeciesofthedevelopmentofpetroleumresourcesontheOuterContinentalShelf(OCS).Thisentailedareviewofavailableinformationinorderto:1)determinewhere and whencoastalbirdsoccurinmarineareasthatmaybedevelopedforoilandgasproduction;2)ascertainwhichspecieswould be mostatriskfromoilspillsandthedevelopmentofoilresources;3)evaluatetheimportanceofpopulationsinthesoutheasternUnitedStatesinrelationtotheglobaldistributionandabundanceofthespecies;and4)summarizeinformationonthe historyofthespeciesmostlikelytobeadverselyaffectedbythedevelopmentofoilresources.Thismaterialispresentedina formthatenablestheBureauofLand Man agement(BLM)toidentifyaspectsofOCSdevelopmentthatmightthreatenpopulationsofmarinebirds.Itprovidesinformationthatwillaidmanagersinmakingdecisionsthatminimizedamagetothesepopulationsduringthedevelopmentofenergyresources.Acorollaryobjectiveistorecommendtopicsforfutureresearchinareaswhereinformationisparticularlyscarce.STUDYAREAThestudyareaencompassesthecoastalandoffshorewatersofthesoutheasternUnitedStates,fromthenorthernborderofNorthCarolinatotheMexicanborderofTexas.Manycoastalhabitatsoccurwithinthisarea:sandybar-rierislands;fresh,salt,andbrackishmarshes;openbeaches;coastalbays;dredge-spoilislands;mudflats;and mangroveislands.Thedominanthabitatsarediscussedbelow.HABITATSNorthCarolinaisdominatedby aseriesoffringingbarrierbeachesbehindwhichlielargeestuarieswithextensiveareasofshallowwaterandsaltmarsh.Thesefringingislands(theOuterBanks)arefarther(30-50kmor20-30mi)fromthemainlandthanaresuchislandsalongotherareasoftheAtlanticcoast(Warinneretal.1976).ExtensivestandsofsaltmarshwithdeeptidalchannelsarefoundsouthofCapeLookout,North Carolina, throughSouthCarolinaandGeorgia.Almostthree-quartersofthesalt-marshacreagealongtheAtlanticseaboardisfoundinthesethreestates.Thelargestareasof'saltmarshontheAtlanticcoastareinGeorgia,whichhas193,000ha(477,000ac),North

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FWS/OBS-83/30September1983MARINEBIRDSOFTHESOUTHEASTERNUNITEDSTATESANDGULFOFMEXICOPARTIIIbyRogerB.Clapp,DeborahMorgan-Jacobs,andRichardC.BanksMuseumSectionUnitedStatesFishandWildlifeServiceNationalMuseumofNaturalHistoryWashington,D.C. 20560ContractNo.14-16-0009-78-917ProjectOfficersJeffreyA.SpendelowCherryKellerNationalCoastalEcosystemsTeamUnitedStatesFishandWildlifeServiceNASA-SlidellComputer Complex 1010 GauseBoulevardSlidell,Louisiana70458ThisprojectwassponsoredbytheBureauofLand Management(MineralsManagementService)PreparedforNationalCoastalEcosystemsTeamDivisionofBiologicalServicesFishandWildlifeServiceU.S.DepartmentoftheInteriorWashington,D.C. 20240

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DISCLAIMERThefindingsinthisreportarenottobeconstruedasanofficialU.S.FishandWildlifepositionunlesssodesignatedbyotherauthorizeddocuments,nordoesthementionofanycommercialitemsindicateanendorsementbytheU.S.Government.LibraryofCongressCardNumber82-601888Thisreportshouldbecitedas:Clapp,R.B.,D.Morgan-Jacobs,andR.C.Banks.1983.MarineBirdsoftheSoutheasternUnitedStatesandGulfofMexico.PartIII:Charadriiformes.U.S.FishandWildlifeService,DivisionofBiologicalServices,Washington,D.C. FWS/OBS-83/30.xviand853pp.

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PartHIof thevolumesMarineBirds oftheSoutheasternUnited States and Gulf of Mexico,published by t"iieNationalCOarulJ::cosystelllsasynthesiil"'i'ii'(l"analysis of inforution aboutthe marine birdsin thisarea. Accountsfor22 species include information ondistribution,abundance,andsusceptibilitytooilpollution.Also included isinforlJlationonthebreedingbiologyof16 species abundantin the southeastasbreedingbirds, winter residents,ormigrants.Selectedbibliographiesfolloweachspeciesaccountandincludeadditionalsourcesofinformation.Anysuggestionsorquestions regarding thisreportshouldbedirectedInformationTransferSpecialistNationalCoastal l::cosystems TeamU.S. fish andWildlifeService NASA-SlidellComputer ColJlplex1010 Gausellou.levard Slidell.Louisiana 70458A badlyoiledgull(Larusap.).PhotographbyRogerB.Clapp.

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Informationonthe seasonal distributionandabundanceof22speciesof marine birdsoftheorder CharadriUorllles thatoccurinthecoastalsouth eastern UnitedStateshas been compiledandmappedfro.theliterature.In many instancesthisprovidesthefirst synthesis of knowledge aboutaspeciesforthisregion. We alsoprovide infon=ation onglobaldistribution,habitatandfoodforallspecies,andinclude information Onvariousaspectsoflifehistoryforthe16speciesthat we consider IllOst ill.portantincoastalareas. This information was gatheredinanattempttoassess the possibleeffectsofoffshoreoildevelopmentonpopulationsof llIadne birdsinthesoutheast.Thesusceptibilityofbirdstooildependsnotonlyontheirjuxtapositionintillleandspace,butalsooncurrents,clilllaticfactors,thestageof the lifeorannualcycle,sndthebehaviorofthespecies.Contaminationbyoil may resultinmattedfeathers with deathfollowingfromchilling,starvation,andingestionofoilduringpreening.Fewofthespeciescoveredinthisreportareatgreathazardfromthedirecteffectsofo11ing,butpopulacionsofmostof t;hese species are highlysusceptibletoenvironmentalchange.Largeconcentrationsofwintering,breeding,andmigrantgullsandternsoccurinthesoutheastandinsomeinstancesmakeupalargeproportionoftheglobalorNorth Am"erican population.Consequently,thisreportincludesmostofthemarinebirdsthat we believemostlikelytobedetrimentallyaffectedbythedevelopmentofoilresources.Oneoftheconclusionsreachedbythisreportisthat we stillknowverylittleaboutthestatusandpopulationsof SOllll'. ofthecharadriiformsthatoccurinthesoutheast.Additionalsurveysofcolonial _dne birdsinthe.southeastandnearbywatersare badly neededto ensure thatweknowenoughaboutthemtopreventtheir unti.ely lossfromourcoastalareas.ACommonTernnest.PhotographbyRogerB.Clapp.

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23BreedingRange Map forthe Brown NoddyandSootyTern24Breeding Kange Hap fortheBlack Sk.blller 65773225 \linter Distribution Hap fot" theBlack Sk1J:Der ..733Adult Laughing GulloffHatteras, North Carolina,inAugust 198!. PhotographbyRogerB.Clapp_

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CONTENTSPREFACEABSTRACTMAPSTABLESABBREVIATIONSUSEDINTEXTPURPOSEOF REPORT STUDYAREAHabitatsClimatesMETHODSARRANGEMENTANDCONTENTSOFSPECIESACCOUNTSpeciesIncludedScientificandVernacularNamesGeneralDistributionDistributionandAbundanceintheCoastalSoutheasternUnitedStatesSynopsisofPresentDistributionand AbundanceHabitatFoodandFeedingBehaviorImportantBiologicalParametersSusceptibilitytoOilPollutionSpeciesBibliographyOILPOLLUTIONANDMARINEBIRDSOFTHESOUTHEASTERN UNITED STATESVariabilityamongSpeciesinSusceptibilitytoOilPollutionRegionalandSeasonalDifferencesinOilingandMortalityofBeachedBirdsBirdKillsFollowingOilSpillsintheSoutheasternU.S.SourcesofVariationinMortalityfromOilPollutionEffectsofOilonContaminatedBirdsandTheirEggsPotentialHazardstoMarineBirdsfromOffshoreOilProductionRECOMMENDATIONSFORFUTURERESEARCHviiiivviiixxv11 134 78910 10111112 12131314 1518212225 2628

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StatusandBiologyofBreedingSpecies 28DistributionandBiologyofTransient,Wintering,andPelagicPopulations 33EffectsofOilonSoutheasternMarineBirds35ACKNOWLEDGEMENTS37SPECIESACCOUNTS39CHARADRIFORMESScolopacidae(Phalaropidinae)Red-neckedPhalarope(Phalaropuslobatus)account39bibliography52RedPhalarope(Phalaropusfulicaria)Laridae(Larinae)LaughingGull(Larusatricilla)Franklin'sGull(Laruspipixcan).Bonaparte'sGull(Larusphiladelphia)Ring-billedGull(Larusdelawarensis)HerringGull(Larusargentatus)GreatBlack-backedGull(Larusmarinus)Laridae-(Sterninae)Gull-billedTern(Sternanilotica)CaspianTern(Sternacaspia)viaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliography657689108 121133142151156176228 301313335 350 365 379

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RoyalTern(Sternamaxima) SandwichTern(Sternasandvicensis)RoseateTern(Sternadougallii)CommonTern(Sternahirundo)account.bibliographyaccountbibliographyaccountbibliographyaccountbibliography393 409 417 436 450 467 483 512ArcticTern(Sternaparadisaea)account551bibliography.559Forster'sTern(Sternaforsteri)account579bibliography593LeastTern(Sternaantillarum)account599bibliography617BridledTern(Sternaanaethetus)account637bibliography651SootyTern(Sternafuscata)BlackTern(Chlidoniasniger)Brown Noddy (Anousstolidus)Laridae(Rynchopinae)BlackSkimmer (Rynchopsniger)LITERATURECITEDviiaccountbibliographyaccountbibliographyaccountbibliographyaccountbibliography656 675 689 701717724731 747 757

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Number12 345678910111213 14 15161718 19202122MAPSBreedingRangeMapfortheLaughingGullWinterDistributionMapfortheLaughingGullWinterDistributionMapfortheBonaparte'sGullWinterDistributionMapfortheRing-billedGullBreedingRangeMapfortheHerringGullWinterDistributionMapfortheHerringGullBreedingRangeMapfortheGreatBlack-backedGullWinterDistributionMapfortheGreatBlack-backedGullBreedingRangeMapfortheGull-billedTernWinterDistributionMapfortheGull-billedTernBreeding Range MapfortheCaspianTernWinterDistributionMapfortheCaspianTernBreedingRangeMapfortheRoyalTernWinterDistributionMapfortheRoyalTernBreedingRangeMapfortheSandwichTernWinterDistributionMapfortheSandwich Tern BreedingRangeMapfortheRoseateTernBreedingRangeMapfortheCommonTernWinterDistributionMapfortheCommonTernBreedingRangeMapfortheForster'sTernWinterDistributionMapfortheForster'sTernBreeding Range MapfortheLeastTernviiiPage9192 143 157 198 199 302 303 336 337 368 369394 395 420 421 451 485 486 581 582601

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TABLESNumber 1 Numberandpercentageofbeachedbirdsexaminedthatwereoiled16 2SusceptibilLtyofvariouslaridstooilingasindicatedbybandingrecoveries173ComparisonofregionalandseasonalvariationofbeachedbirdmortalityandincidenceofoilingintheeasternUnitedStates19 4Comparisonoffive-yearaveragesofbeachedbirdmortalityfordifferentareasintheeasternUnitedStates20 5OilspillsinthesoutheasternUnitedStates1978-1980comparedwithotherstates 23 6EstimatedpopulationsofcharadriiformmarinebirdsbreedinginthesoutheasternUnitedStatescomparedwiththeirstatusinotherareasadministeredby the U.S.297ApproximatenumberofRed-neckedPhalaropesrecordedbymonthforthecoastalsoutheasternUnitedStates 48 8DatesofoccurrenceforRed-neckedPhalaropesinthecoastalsouth-easternUnitedStates. 49 9ApproximatenumberofRedPhalaropesrecordedbymonthforthecoastalsoutheasternUnitedStates7210DatesofoccurrenceforRedPhalaropesinthecoastalsoutheasternUnitedStates 7211RecentestimatesofLaughingGullpopulationsnestinginthesouth-easternUnitedStates9312PeakconcentrationsofmigrantandwinteringLaughingGullsinthecoastalsoutheasternUnitedStates9613MeanclutchsizesreportedfortheLaughingGull10314RatesofhatchingsuccessreportedfortheLaughingGull1041516Weights(ingrams)ofLaughingGullsApproximatenumberofFranklin'sGullsrecordedby monthfortheareafromNorthCarolinatoMississippix106126

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Number17DatesofoccurrenceforFranklin'sGullsinthecoastalsoutheasternUnitedStates 12718PeakconcentrationsofmigrantFranklin'sGullsincoastalTexas12719Weights(ingrams)ofFranklin'sGulls13220PeakconcentrationsofwinteringBonaparte'sGullsinthecoastalsoutheasternUnitedStates14521MeanclutchsizesreportedfortheBonaparte'sGull14922 MeanclutchsizesreportedfortheRing-billedGull16823RatesofhatchingsuccessreportedfortheRing-billedGull17124ProductivityinRing-billedGullcolonies17325Weights(ingrams)ofRing-billedGulls17526HerringGullsnestinginNorthCarolina20127LayingperiodsreportedatsomeNorthAmericanHerringGullcolonies21428 MeanclutchsizesreportedfortheHerringGullinNorthAmerica21529ComparisonofmeanclutchsizesforearlyandlateclutchesinNorthAmericanHerringGulls21830 Rates ofhatchingsuccessreportedfortheHerringGullinNorthAmerica 21931ComparisonofhatchingsuccessforearlyandlatenestsofNorthAmericanHerringGulls22132Comparisonofhatchingsuccessforthree-andtwo-eggclutchesinsomestudiesofNorthAmericanHerringGulls22233ProductivityinNorthAmericanHerringGullcolonies22334Weight(ingrams)ofNorthAmericanHerringGulls227 35 NumberofdeadbirdsandnumberandpercentageofdeadHerringGullsfoundaftermajoroilingincidents22936DatesofoccurrenceforGreatBlack-backedGullsinthecoastalsoutheasternUnitedStates307xi

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Number37NumberofoiledbirdsandnumberandpercentageofdeadGreatBlack-backedGullsfoundaftermajoroilingincidents31438RecentestimatesofGull-billedTernpopulationsnestinginthesoutheasternUnitedStates342 3940MeanclutchsizesreportedfortheGull-billedTernWeight(ingrams)ofGull-billedTerns349351414243 4445RecentestimatesofCaspianTernpopulationsnestinginthesoutheasternUnitedStatesRecentestimatesofRoyalTernpopulationsnestinginthesoutheasternUnitedStatesPeakconcentrationsofwinteringandmigrantRoyalTernsinthecoastalsoutheasternUnitedStatesMeanclutchsizesreportedfortheRoyalTernWeights(ingrams)ofRoyalTerns371399 401 405 40846RecentestimatesofSandwichTernpopulationsnestinginthesouth-easternUnitedStates42447PeakconcentrationsofwinteringandmigrantSandwichTernsinthecoastalsoutheasternUnitedStates42648MeanclutchsizesreportedfortheSandwichTern43149RatesofhatchingsuccessreportedfortheSandwichTern432 50Weights(ingrams)ofSandwichTerns43551EstimatedformerandpresentpopulationsofRoseateTernsinnorth-easternNorthAmerica45952ApproximatenumberofRoseateTernsrecordedbymonthforthecoastalsoutheasternUnitedStates46253 54 55PeakconcentrationsofmigrantCommonTernsinthecoastalsoutheasternUnitedStatesMeanclutchsizesreportedfortheCommonTernIncubationperiods(indays)reportedfortheCommonTern.;:ii492 500 502

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Number56RatesofhatchingsuccessreportedfortheCommonTern..5035758FledgingsuccessinCommonTerncoloniesProductivityinCommonTerncolonies50550659Weights(ingrams)ofCommonTerns50960DatesofoccurrenceforArcticTernsinthecoastalsoutheasternUnitedStates.........55561ApproximatenumberofArcticTernsrecordedby monthforthecoastalsoutheasternUnitedStates55562PeakconcentrationsofwinteringandmigrantForster'sTernsinthecoastalsoutheasternUnitedStates58063RecentestimatesofForster'sTernpopulationsnestinginthesouth-easternUnitedStates58664MeanclutchsizesreportedfortheForster'sTern.59165 6667686970Weights(ingrams)forForster'sTernsRecentestimatesofLeastTernpopulationsnestinginthesoutheasternUnitedStatesMeanclutchsizesreportedfortheLeastTernRatesofhatchingsuccessreportedfortheLeastTernProductivityinLeastTern'coloniesWeights(ingrams)ofLeastTerns593608613 61461561871ApproximatenumberofBridledTernsrecordedby monthforthecoastalsoutheasternUnitedStates64872737475DatesofoccurrenceforBridledTernsinthecoastalsoutheasternUnitedStatesNumbersofSootyTernsfoundbreedingintheChandeleurIslandsRecent hreedine ofSootyTernsalongtheTexascoastPeakconcentrationsofSootyTernsinthecoastalsoutheasternUnitedStatesxiii649 663 664 667

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76Incubationperiods(indays)reportedfortheSootyTern66977RatesofhatchingsuccessreportedfortheSootyTern.67078Weights(ingrams)ofSootyTerns...67479PeakconcentrationsofmigrantBlackTernsinthecoastalsouth-easternUnitedStates..69280Meaoclutch sizes reportedfortheBlackTern69981Weights (in grams)ofBlackTerns...82 kecent estimatesofBlackSkimmerpopulationsnestinginthesoutheasternUnitedStates...73783PeakconcentrationsofmigrantandwinteringBlackSkimmersinthecoastalsoutheasternUnitedStates...73884Weights(ingrams)ofBlackSkimmers..746AnimmatureLaughingGull.PhotographbyRogerB.Clapp. dv

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ABBREVIATIONSUSEDINTEXTMostoftheabbreviationsusedinthetextareinstandarduseandwillbe knowntothereader;a fewmaybelessfamiliar.Theseare belowwithabriefindicationoftheirinterpretation.N,S,E,W,(capitalizedwithoutperiod)N.,S.,E.,W.(captitalizedwith period) compassdirectionsgeographicsitedesignation(e.g.,S.PadreIslandacad.AOUBOUBLMca.CBCcf.coll.compoCo.CSLPEHBAed./eds.et.seq.haIBBAimm.inUtt.inprep.msnNatLParkNatl.Seashore NERC nonad.NWROCSop.cit.Par.pers.comm.pers.observephotogr.prep.SDspec.sp./spp.St.Parksubad.acreadultAmericanOrnithologists'UnionBritishOrnithologists'UnionBureauofLand Management(circa)aboutChristmasBirdCount(confer)compare/seecollectedcompilerCountyCenterforShort-LivedPhenomenaEasternBirdBandingAssociationeditor/editorsandthefollowinghectareInlandBirdBandingAssociation inunature intheletters(of)inpreparationmanuscriptsamplesizeNationalParkNationalSeashoreNationalEnvironmentalResearchCouncilnonadultNationalWildlifeRefugeOuterContinentalShelf(operecitato)intheworkcitedParishpersonalcommunicationpersonalobservationphotographedpreparerStandardDeviationspecimenspecies(singular/plural)StateParksubadultxv

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subseq.subsequentunpubl.unpublished USF\!S UnitedStatesFishandWildlife Service USNFWLUnitedStatesNationalFishandWildlifeLaboratoryWAGBIWaterfowlers Association ofGreatBritainandIreland WMA WildlifeManagementAreaUnitSofmeasurementinthetextarepresentedas they wereinthesourcefromwhichtheywerederived,andarefollowedinparentheses by conversiontothemetricorEnglishsystems,asappropriate.AnadultHerringGull.PhotographbyJ.A.Spendelow.

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PURPOSEOFREPORTThepurposeofthisreportisto thestatusof22coastalandmarinecharadriiformspeciesinthesoutheasternUnitedStatesandtoexplorethepotentialeffectsonthesespeciesofthedevelopmentofpetroleumresourcesonthe.OuterContinentalShelf(OCS).Thisentailedareviewofavailable information inorderto:1)determinewhere andwhencoastalbirdsoccurinmarineareasthatmaybedevelopedforoilandgasproduction;2)ascertainwhichspecieswould be mostatriskfromoilspillsandthedevelopmentofoilresources;3)evaluatetheimportanceofpopulationsinthesoutheasternUnitedStatesinrelationtotheglobaldistributionand abundanceofthespecies;and4)summarizeinformationonthe historyofthespeciesmostlikelytobeadverselyaffectedbythedevelopmentofoilresources.Thismaterialispresentedina formthatenablestheBureauofLand Man agement(BLM)toidentifyaspectsofOCSdevelopmentthatmightthreatenpopulationsofmarinebirds.Itprovidesinformationthatwillaidmanagersinmakingdecisionsthatminimize damagetothesepopulationsduringthedevelopmentofenergyresources.Acorollaryobjectiveistorecommendtopicsforfutureresearchinareaswhereinformationisparticularlyscarce.STUDYAREAThe study areaencompassesthecoastalandoffshorewatersofthesoutheasternUnitedStates,fromthenorthernborderofNorthCarolinatothe Mexica9 borderofTexas.Manycoastalhabitatsoccurwithinthis area: sandybar/ rierislands;fresh,salt,andbrackish marshes; openbeaches;coastalbays;dredge-spoilislands;mudflats;and mangroveislands.The dominanthabitatsarediscussedbelow.HABITATSNorthCarolinaisdominated by aseriesoffringingbarrierbeachesbehindwhichlielargeestuarieswithextensiveareasofshallowwaterandsaltmarsh.Thesefringingislands(theOuterBanks)arefarther(30-50kmor20-30milfromthemainlandthanaresuchislandsalongotherareaSoftheAtlanticcoast(Warinrteret'al.1976).ExtensivestandsofsaltmarshwithdeeptidalchannelsarefoundsouthofCape Lookout, North Carolina, throughSouthCarolinaandGeorgia.Almostthree-quartersofthesalt-marshacreagealong the Atlanticseaboardisfoundinthesethreestates.Thelargestareasofsaltmarsh ontheAtlanticcoastareinGeorgia,which has193,000ha(477,000ac),North

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Carolina(64,000haor158,000ac),andSouthCarolina(176,000ha or 435,000ac)(West1977).Barrierislandsarealsoveryimportantcoastalhabitatin these threestates.Thelandareasofthebarrierislandsforeachstateare120,000ac(49,000ha)inNorthCarolina,124,000ac(50,000ha)inSouthCarolina,and153,000ac(62,000ha)inGeorgia(Warner1976),foratotalofabout397,000ac(161,000ha).Theareaofwaterbehindtheseislandsbecomessmallertothesouth(Warinneretale1976).NorthCarolina,SouthCarolinaandGeorgiahaveabout266 mi (428km),199mi(320km),and98mi(158km)ofopenbeach,respectively,alongtheirbarrierislands.Inotherpartsofthestudyarea(e.g.,partsoftheFloridaGulfcoast),beachesarefewornonexistent(WoolfendenandSchreiber1973).TheeastcoastofFloridaisdominatedby achainofbarrierislandsbrokenoccasionallybytidalpasses.Typically,theseislandsaresandyalongtheirouter perimeters. Largeareasofmarshandestuarineswamplielandwardoftheseislands(Warinneretale1976)andsaltmarshesgraduallygivewaytomangroveswampasone moves furtherlandward(Reimold1977).MuchoftheGulfcoastofFloridaisdominatedbysaltmarshesand mangroveSWamps(Warineretal.1976).Extensive.stretchesofopenbeacharefoundftomNaples ontheFloridapeninsulanorthalongthepanhandletoAlabama (Woolfenden andSchreiber1973).InAlabama,tidalsaltmarshes,sandybeaches,andoffshoreislandsarecommoncoastallandforms.Mississippi'sGulfcoastconsistsalmostentirelyofbarrierislandsthathavesaltmarshesintheircenters.TheMississippishorelineisextensivelydevelopedbutstillcontainsfresh,salt,andbrackishmarshes(Warinner et al.1976).Only alimitedamountofsaltmarshisfoundbetweennorthernFloridaandMississippi.Mostmarshesaresmall,disjunct,andinalluvialpocketsprotectedbybayshores(West1977).LouisianahasmoremarshandestuarineareathananyoftheotherUnitedStatesexceptAlaska(Warinneretal.1976),andcontainsnearlyhalfthetotalacreageofsaltmarshinthecontiguousUnitedStates.Insomeplacesthemarshesextendinlandasmuchas40-50km(25-30mi)(West1977).Thecoastlinealongthewesternthirdofthestateissandy,buttherestoftheareaisdominatedbybarrierislandsandmarshthatarestronglyinfluencedbytheenormous amountsofmudandsiltdepositedbytheMississippi River (Warinneretale1976).TheLouisianacoastisoneofthemostproductiveareasformarinebirdsinthecontinentalUnitedStatesandsupportsenormouswinteringpopulationsofwaterfowl.ThecoastofTexas makes up alargeportionofthewesternshoreoftheGulfofMexico.Sandybeachesandoffshorebarrierislandsareabundant.Twosemi-landlockedlagoons,theUpper and Lower Laguna Madre, and alargelow-salinityestuary,SabineLake,areareasofgreatimportancetowinteringwaterfowl.Anestimated 78% oftheworld'spopulationofRedhead(Aythya amer-. icana)duckswintersintheLagunaMadre,and 13% oftheworld'sshrimp harvest comesfromTexaswaters(Warinneretale1976).AlimitedamountofsaltmarshispresentinTexasalongbayshoresenclosedbyoffshorebars(West1977).2

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CLIMATESTheclimaticregime,likethelandform,differswidelyfrom onepartofthestudyareatoanother.Thenortheasternportionisthe coldest. ThelowestmidwintertemperaturesalongthecoastofNorthCarolinaareontheorderof 2ifF (-70C)andtheaveragedailymaximumduringmidsummeralongtheextremesoutherncoastisonly86F(300C),some 60Flessthanisusuallyrecordedintheinterior.Julyisthewettestmonth andOctoberthedriest.Alongthecoast,snow andsleetusuallyfallonlyonceortwiceayearandareusuallyassociatedwithnortheasterlywinds.PrevailingwindsinNorthCarolinablow fromthesouthwestmostoftheyearand fromthenortheastinSeptember andOctober(Hardy1974).TheweatheralongthecoastofSouthCarolinaissimilartothatinNorthCarolinawithsomevariation.AverageannualtemperaturesalongtheSouthCarolinacoastareabout 68PF(200C), withanaveragedailymaximuminJulyof880F(3l0C)andaveragedailyminimumsinJanuaryfrom 3SO F (l.JOC) inthenortheastto42F(6C)inthesoutheast.Marchisparticularlyrainyalongthecoast,andOctoberand Novemberarethedriestmonths.PrevailingwindsinSouthCarolinaarefromthesouthwestandsouthinspringand summer,predominantlyfromthenortheastinautumn,and fromthenortheastandsouthwestinwinter(Landers1974).TheclimateinGeorgiaischaracterizedbyshortmildwintersandwarmhumid summers. Thecoastalareabecomesprogressivelydrierand warmer fromnorthtosouth.Peakperiodsofprecipitationoccurinwinterandearlyspring;theaverageannualrainfallrangesfrom75in(190cm)intheextremenortheasternpartofthestateto53in(135cm)alongthelowereastcoast.Average summertemperaturesrangefrom 73PF(Z3O C)intheextremenorthto 8ZOF (Z80C)inpartsofsouthGeorgia;averagetemperatureforthethreewintermonthsrangesfrom 4loF(SOC) inthenorthto560F (lJOC) onthelowereastcoast.AreasinnorthernGeorgiahavefreezingtemperaturesduringthedayforalmostathirdoftheyear'butthelowercoastonlyhasabouttendaysoffreezingtemperaturesannually(Carter1974).Floridahasawiderrangeofclimatethananyotherstateinthesoutheast.Theclimaterangesfromtemperatetosubtropicalinthenorth,totropicalintheFloridaKeys. Summersarewarm,humid,andlong,andwintersaremildandbrief.Rainfallisabundant,especiallyfromJunetoSeptember.Meanannualtemperaturesrangefromtheupper60's(F)innorthernFloridatothemid-70'sinthesouthandreachnearly"780F(Z60C)atKeyWest.Rainfallvarieswidelyfromareato area andfromyeartoyear,withmostareasusuallyreceivingbetween 50 and65in"(130-170cm). ThedrierKeyshaveanaverageannualrainfallofonlyabout40in(100cm).Onthesouthernpartofthepeninsulaprevailingwindsarefromthesoutheastandeast;elsewheretheyaremoreerraticand "tendtobe fromthenorthinwinterand fromthesouthin summer. Tropicalstormsfrequentlycausegreatdamage; fewyearspasswithoutahurricaneaffectingpartofthestate(Bradley1974).The GulfofMexicohasamaritimetropicalclimatewithmeanwinteraturesofabout 700F (ZlOC) and mean summertemperaturesof84F(ZgoC).tivetoseasonsinotherpartsofthestudyarea,both summer andwinterhotand humid;humidityisgreatestduringspringand summer, andlowest3temperRelaareduring

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latefallandwinter(BLM1978a).RainoccursfairlyevenlythroughouttheyearalongthewesternandnorthernGulf,withapeakfromJunethroughAugust(BLM1978a).FurthereastintheGulfthepeaktendstobelaterandfallsinAugustandSeptember(BLM1978b).Theareabecomesprogressively fromthesouthwesttothenorthandcentralportionsofthenorthernGulf.ThedriestareaoftheTexascoastextendsfromBrownsvillenorthtoaboutCorpusChristi;themost humidareafromGalvestontotheSabineRiver(Chaneyetal.1978).Averageannualprecipitationrangesfromabout69cm(27in)atBrownsvilleto137cm(54in)at New Orleans(BLM1978a)and170cm(67in)inMobile(BLM1978b).Tropicalstormsandhurricanesregularlyoccurduringlatesummer andfallandoftenravagecoastalhabitats.MostofthesestormsentertheGulfthroughtheYucatanChannelandStraitsofFlorida(BLM1978a).SoutheasterlywindspredominateoverthenorthernGulfduringthesummer.Easterliesaremore commonduringthewinter,andprevailingwindsfromthewestandsouthwestarerareatanytimeofyear(BLM1978a).METHODSMost.oftheinformationwasobtainedby aliteraturesearch.Additionalinformationonoilingofindividualspeciesofbirdsandtheirdistributionwasobtainedthroughexaminationofmuseumspecimensandinterviews,butthesewerenotmajorsources.Severalcomputerizedinformationretrievalsystemswereinvestigated,butdidnotmeetourneeds.Thesesourceswereparticularlyweak onthelocaldistributionofbirds,muchofwhichistobefoundinregionaljournalsnotcoveredbycomputerservices;thetemporalcoveragewasalsoinadequateforthisstudy.AsBartonekandLensink(1978)pointedout,visualsearchesofperiodicals"provedfarmoreproductivefromthestandpointofbothnumbersofcitationsandthoroughnessofthesearch."Weobtainedliteraturecitations-primarilybyscanningtheliteratureand byconsultingbibliographiesinrelevantpapers.Theprimarysourcesforthejournals,books,andpaperswerethelibrariesandreprintfilesoftheBirdDivisionsoftheSmithsonianInstitution,Washington,D.C.andtheAmericanMuseumofNaturalHistory,NewYork.OthermajorsourcesofinformationwerethelibraryoftheDepartmentoftheInterior,theLibraryofCongress,andtheBirdLibraryandreprintfilesofthePatuxentWildlifeResearchCenter,Laurel,Maryland.TheWelderWildlifeFoundation,Sinton,Texas,andthelibraryofgovernmentpublicationsandreportsmaintainedbytheNationalCoastalEcosystems Team,Slidell,Louisiana,wereparticularlyrichsourcesofinformationotherwisedifficulttoobtain.Unpublishedreportsandpaperswereobtainedfrom:TheFloridaAudubonSociety,VeroBeach;theFloridaFishandGameCommission,Gainesville;EvergladesNationalPark,Homestead; andindividualslistedintheacknowledgments.SeveraldozenvaluablebutunpublishedthesesWereobtainedfromeducationalinstitutions.

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Searchesweremadeofseveralsecondarysourcesofliteraturecitations.Literaturereviewsectionsofmajorornithologicaljournals,particularlyTheAuk,TheIbis,andtheJournalofFieldOrnithology(Bird-Banding)wereespeciallyuseful,aswasWildlifeReview.Wealsomadeextensive ofCurrentContents,OilPollutionAbstracts,andDissertationAbstracts.TheZoologicalRecord,BiologicalAbstractsandEcologicalAbstractswerealsoconsultedbutwerelessefficientsourcesofinformation.AllstatebirdjournalsdealingwiththesoutheasternUnitedStates(seelistbelow)werescanned;thesejournals,alongwithAmericanBirds(AudubonFieldNotesinearliervolumes),providedmuchoftheinformationonlocaldistributionineachstate.Weplacedconsiderableemphasisonrecentnessofinformationintheliteraturesearch.A fewjournals(e.g.,WilsonBulletin,Bird-Banding)wereexaminedforatleast30yearsintothepast,TheAukfrom 1930tothepresent.Manyothers,dependingonthedegreetowhichtheyyieldedusefulinformation,werescannedforonlya fewrecentyears.Wecoveredtheforeignliteratureasthoroughlyaspossible.Mostofthespeciestreatedinthisreporthavea widegeographicdistribution,andmuchofwhatisknownoftheirbreedingbiologyistobefoundonlyinforeignperiodicals.Thelinguisticlimitationsoftheauthors,aswellasthetemporalandfiscallimitationsinvolvedintheproductionofthisreport,precludedfulluseofthismaterial.Listedbelowaretheserialpublicationscoveredextensively.Whereappropriate,thoseareasoftheworldthatthesejournalscovermostthoroughlyarelistedinparentheses.ActaOrnithologica(Poland,U.S.S.R.)AlabamaBirdsAlauda(France,FrenchAfrica)AmericanBirds[AudubonFieldNotes](UnitedStates,Canada) AnimalBehaviorArdea(westernEurope)AtlanticNaturalist(DelawaretoVirginia)AtollResearchBulletinAuk(NorthAmerica)AustralianBirdWatcherBehaviourBiologia(Bratislava)(SeriaB)(Czechoslovakia)BiotropicaBirdStudy(GreatBritain)BlueJay(centralCanada)BritishBirdsBulletinoftheBritishOrnithologists'Club(world)BulletinoftheKansasOrnithologicalSociety5BulletinoftheOklahomaOrnithologicalSocietyBulletinoftheTexasOrnithologicalSocietyCaliforniaFishandGameCanadianField-NaturalistCanadianJournalofZoologyChat(NorthandSouthCarolina)ColonialWaterbirds(easternNorthAmerica)Condor(NorthAmerica,neotropics)Corella[AustralianBird-Bander]DanskFugle(Denmark) DanskOrnithologiskForeningsTidsskrift(Denmark)EcologyEkologiaPolska(Poland)Elepaio(Hawaii)Emu(Australia,NewGuinea) Estuarieg [ChesapeakeScience](U.S.AtlanticCoast)

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FieldNaturalistFloridaNaturalistFloridaScientistGerfaut(westernEurope,Africa)Hornero(Argentina)Ibis(OldWorld,Africa)IrishBirdsJack-PineWarbler(Michigan)JournalfurOrnithologie(Germany)JournalofAnimalEcologyJournalofAppliedEcologyJournalofEcologyJournalofFieldOrnithology[BirdBanding](UnitedStates)JournalofWildlifeManagement(NorthAmerica)Kingbird(NewYork)Larus(Yugoslavia,easternEurope)Limosa(Netherlands)L'OiseauetlaRevueFrancaised'Ornithologie(France)Loon[Flicker](Minnesota)LouisianaOrnithologicalSocietyNewsMarinePollutionBulletinMarylandBirdlifeMississippiKiteMississippiOrnithologicalSocietyNewsletterMurrelet(Pacificnorthwest,Alaska,westernCanada)NosOiseaux(France,westernEurope)Notornis(NewZealand,Pacificislands)Oikos (Denmark,Scandinavia)Oriole(Georgia)OrnisFennica(Finland,Balticarea)OrnisScandinavica(Scandinavia)OrnithologischeBeobachter(Switzerland,middleEurope)OrnithologischeMitteilungen(world)Ostrich(SouthAfrica)ProceedingsoftheAnnualConference,SoutheasternAssociationofGameandFishCommissioners(southeasternU.S.)ProceedingsoftheLouisianaAcademyofScienceRevueSuissedeZoologie(Switzerland,centralEurope)Ring(Europe).Ringing&Migration(GreatBritain)RivistaItalianadeOrnitologia(Italy)ScottishBirdsSouthAustralianOrnithologistSouthwesternNaturalist(southwesternU.S.)SovietJournalofEcologySterna(Norway)SuomenRiista(Finland,Balticarea)TexasJournalofScienceTori(Japan)TransactionsoftheNorthAmericanWildlifeandNaturalResourcesConferenceVarFagelvarld(Sweden)VestnikZoologii(U.S.S.R.)Vogelwarte(westernandcentralEurope)WesternBirds(westernU.S.)WildfowlWilsonBulletin(NorthAmerica)ZeitschriftfurTierpsychologieZoologicheskiiZhurnal(U.S.S.R.)Thereprintfilesofseveralinstitutionssuppliedsomelesseasilyobtainablematerial.The mostusefulofthesewerefilesoftheMuseumSectionoftheDenverWildlifeResearchCenter(formerlytheNationalFishandWildlifeLaboratory),theBirdDivisionofthe MuseumofNaturalHistory,theAmericanMuseumofNaturalHistory,andtheBirdLibraryoftheGabrielson6

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LaboratoryofthePatuxentWildlifeResearchCenter.Inall,about10,000citationsdealingdirectlywiththespeciestreatedareincludedinthethreepartsofthisreport.The more generdl articlesfoundintheLiteratureCitedsectionsattheendofeachvolumewillprobablycontainatleastanadditional1,000citatons.Certaindata(e.g.,weights)reportedinthisstudywereobtaineddirectlyfromdatainthefilesandcollectionsofmuseums,particularlythoseoftheU.S.NationalMuseumandLouisianaStateUniversity.ARRANGEMENTANDCONTENTSOFSPECIESACCOUNTSEach volumeofthisseriescoversgroupsofbirdsforwhichtheproblemsofpreparingthisreporthavebeendifferent,eventhoughallspeciesincludedsharethecharacteristicofbeingatleastoccasionallyfoundalongthewatersofthesoutheasternUnitedStates.Volume 1(Clappetal.1982b)dealswithpelagicseabirdswhosestatusandoccurrenceinthisareaareonlynowbeingdocumented.Thesespecies,forthemostpart,donotbreedcommonlyinsoutheasternwaters,andhavereceivedlittlestudy.Volume 2(Clappetal.1982c)dealswithducks,geese,andswans,mostofwhichareimportantinsoutheasternwatersbecauselargenumberswinterormigratethroughthisarea.MostofthesespeciesarecoveredextensivelyintheNorthAmerican andforeignliterature,andmuchisknownabouttheirbreedingbiologyanddistribution.Someofthem,primarilyspeciesthatareseldomhunted(e.g.,mergansers,scoters),havebeenlittlestudiedandareamongthosespeciesknowntobemostadverselyaffectedbyoilpollutioninotherareas.Informationondistribution,status,andbreedingbiologyofmanyspeciesofwaterfowlhasbeensummarizedina numberofrecenthandbooks(e.g.,Bellrose1976;Palmer1976a,1976b;Crampetal.1977).Thespeciesaccountspresentedinvolume 3varyinlengthanddetailwiththequalityandquantityofmaterialexaminedand with thespeciesabundanceandvulnerabilitytooilinginsoutheasternwaters.Theaccountstendtobelongerthanthoseintheearliervolumesbecausemostofthespeciesincludedherehavebeenstudiedmorethoroughlyandhavea moreextensiveliteraturethanthespeciesincludedinVolumeI,becausefewhavereceivedmonographictreatmentashavethoseinVolumeII,andbecausemostareregularlyorseasonallyabundantinthesoutheasternUnitedStates.Muchofthedataonthestatusandbiologyinthesoutheastofthespeciesincludedhereisofrecentorigin.Muchusefulinformationwaspublishedwhilethisreportwasbeingcompiled(e.g.,Duncan and Havard 1980,Rowlett1980,Portnoyetal.1981);stilllittleisknownaboutoffshoredistributionformanyspecies.Althoughwehavetriedtomakethisreportastimelyandthoroughaspossible,furtherresearchprobablywillrevealtheinadequacyofourpresentknowledgeofthestatusofmanyspeciesinsoutheasternwaters.7

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SPECIESINCLUDEDThisreportcovers22membersoftheorderCharadriiformes:2phalaropes,6gulls,13terns,andtheBlackSkimmer. Thebirdsdealtwithherearespeciesforwhich most oftheNorthAmerican(andsometimesworld)populationbreeds,winters,ormigratesthroughthesoutheasternUnitedStates.Severalspecies(e.g.,thephalaropes,ArcticandBridledTerns)occurlargelyasoffshoreorpelagicmigrants,others(e.g.,Bonaparte'sGull,CommonandBlackTerns)occurlargelyascoastalmigrants.Somespecies(e.g.,Herringand Hing billedGulls)breedmostlyorentirelynorthofthestudyarea,butalargeportionoftheirNorthAmericanWinteringpopulationoccursinthesoutheasternUnitedStates.Largeportionsofthepopulationsofseveralotherspecies(e.g.,LaughingGull;Royal,Forster'sandLeastTerns)breedinthesoutheasternUnitedStates.Otherspecies(Gull-billed,Caspian.Sandwich.andSootyTerns;Brown Noddy;BlackSkimmer)breedinthesoutheasternUnitedStatesinsuchlargenumbersthattheirpopulationstheremake up asignificantportionoftheNorthAmericanpopulation.Someofthespeciesincluded(e.g.,LaughingGull;Caspianand RoyalTerns;BlackSkimmer)withimportantbreedingpopulationsinthesoutheastalsousethe areaps amajorWinteringground.TheRoseateTern,uncommontorareinthesoutheast,wasincludedbecauseitspopulationsaredeclininginmanyportionsofitsrangeandbecauseithasbeensuggestedasacandidateforlistingasanendangeredorthreatenedspecies. We originallyintendedtoincludeaccountsforallspeciesofseveralsubfamilies (Stercorariinae, Larininae,Sterninae,Rynchopinae)ofthefamilyLaridaeandforall ofmurres.guillemots,andotheralcids(AIcidae)thathadbeenrecordedatleastonceincoastalportionsofthesoutheasternUnitedStates.WealsointendedtoincludeanaccountfortheAmerican Cpot(Fulicaamericana)[Rallidae),aspeciesthatisabundantinmarshycoastalhabitatsinmanyofthesoutheasternUnitedStates.Thelossofpersonnel,andthelossofuse,increasedcostof_repair,anddeteriorationofwordprocessingequipmentandsuppliesmadeitnecessaryforustodeletemanyspeciesfromthisvolume andtocompletethisreportatascalelessthanoriginallyenvisoned.Thedeletedspeciesincludedthefollowing:AmericanCoot,PomarineJaeger(Stercorariuspomarinus),ParasiticJaeger(Stercorariusparasiticus), 1ong tailedJaeger(S.longlcaudus),Skuaspp.(Catharactaspp.),LittleGull(Larusminutus), Commo; Black-headedGull(L.ridlbundus).Band-tailed Gull (L.belcheri),MewGull(L.canus),CaliforniaGUll(L.californicus).ThayerTsGull thayeri),IcelandGull glaucoldes),LesserBlack-backedGull fuscus),GlaucousGull(L.hyperboreus),Black-leggedKittiwake(Rissatridactyla). sabine's Gull(Xemasabini),ElegantTern(Sternaelegans),White-wingedTern(Chlidonias leueopterus), BlackNoddy (Anousminutus),Dovekie(AIleaIle).CommonMurre(Uriaaalge),Thick-billedMurre (U.lomvia), RazorbIII (AIcatorda),and (Cepphusgrylle). The preliminarybibliographiesforthesespeciestotalabout170pagesandhavebeenretainedinthefilesoftheMuseumSection,DenverWildlifeResearchCenter.8

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SCIENTIFICANDVERNACULAR NAMES. EachspeciesaccountisheadedbythecommonEnglishand nameofthespecies,followedbyvernacularnamesinotherlanguagesandalternativecommonEnglishnamesusedintheUnitedStatesandinotherEnglishspeakingcountries.Scientificandvernacularnamesfollowthe34thsupplement(AOU1982)totheAOUChecklist(AOU1957),asdoesthesequenceinwhichwelistorders,families,andsubfamilies.Widelyusedalternativescientificnamesarealsonoted.Explanationismadeinfootnoteswherechangesinscientificnames have beenadoptedrecently.Scientificnamesof other organisms(e.g.,plants,fish,crabs,molluscs)giveninthe text areeitherthoseusedintheworkscitedorare from standsrdrecentreferencesorregionalguides.Missingscientificnameshavebeensuppliedonlywhenwewerecertainwhatspecieswas meant bythevernacularnameusedintheoriginaltext.Theprimarysourceformostofthenon-EnglishvernacularnamesistheNomina AviumEuropaearum(Jorgensen1958);othersourcesconsultedincludeDement'evandGladkov(1951),Austinand Kuroda(1953),Edwards(1972),and Crampetal.(1977).Theabbreviationsforthelanguagesandothergeographicalusesareasfollows:DA:Danish IC:IcelandicPR:PortugueseDU:DutchIT:ItalianRU:RussianEN:English(OldWorld)JA:JapaneseSP:SpanishFI:Fi.nnish NW: NorwegianSW:SwedishFR:FrenchNZ:NewZealandUS:UnitedStatesGE:GermanPO:PolishWith fewexceptions, the commonnamesgivenarethoseinwidestuseintheornithologicalliteratureofthecountriesindicated.InseveralinstancesweincludetransliteratednamesfromlanguagesinwhichRomancharactersarenotused(Japanese,Russian).ForJapanesenameswerelieduponAustinand Kuroda(1953)and ror RussiannamesweusedthenamesusedinthetranslationofDement'evandGladkov(1951).Insomeinstancesmorethanoneforeignvernacularnameforaspecieshasbeen ThisisparticularlytrueforSpanish,inwhichvernacularnamesmayvaryconsiderablyfromareatoarea.Theprimaryreasonforsupplyingthesealternativenamesistoassistfuturecomputer-basedliteraturesearches.Someofthe translationsofforeignnames(whicharethoseenteredoncomputers)implyadifferentspeciesthanthenamewouldnormallysuggestto a readerofEnglishorcannotbereadilyassociatedwith an Englishname(e.g.,thetranslationoftheRussiancommonnameforLarusridibundusisLaughingGull,a namethatinEnglishindicatestheNorth AmericanLarus atricilla).Inaddition,manyofthemoreregionally-orientedforeignlanguagejournals,likethoseintheUnitedStates,9

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failtolistthescientificnames.Asaresult,searchesofcomputerliteraturesystemsbyscientificnamealonemayfailtoindicateimportantnotesorpapersdocumentingrecentchangesindistribution.Alternativescientificnameswidelyorrecentlyinusearesuppliedasanotheraidtoliteraturesearches.TheCaspianTernappearsinrecentliteratureasSternacaspia,Sternacaspius,Sternatschegrava,Hydroprognetschegrava, caspia.Onecomputersearchwemaderevealednolessthanfourdifferent wheneachscientificname wasusedasakeyword.GENERALDISTRIBUTIONThissectionisdividedintotwoparts,oneg1v1ngoccurrenceinNorthAmerica,theothergivingoccurrenceelsewhereintheworld.Mostofthisinformationhasbeentakenfromstandarddistributionalworks,butithasbeensupplemented,wherepossible,withmorerecentliterature.Breedingandwinteringrangesareemphasizedinthissection,withlessinformationgivenaboutareasofoccurrenceduringmigration.MaterialrelatingtoNorthAmericaismoredetailedandmorecompletethanforotherareasoftheworld.DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESInthissectionwepresentmoredetailedremarksondistributionandabundanceinthesoutheast.Asmuchrecentinformationthrough1979aspossiblehasbeenincorporated.Someaccountsprovideevenmorerecentinformationbuttheaccountswerecompletedoveranumberofyearsand,asaresult,thedepthofcoverageisnotconsistent.Thissectionisbaseduponthemostrecentstateornithologicalhandbooksandchecklists;italsoincludesmuchinformationonseasonalobservationspublishedinAmericanBirdsandstatejournals,breedingdatafromtheColonialBirdRegisteratCornellUniverSity,anddatafromunpublishedmanuscripts.Thissectionalsoincorporatesinformationonseasonaloccurrence,breedingstatusandnumbers,andhabitats.Coastalareasareemphasized,butinsomecasesstatuselsewhereisalsomentioned.Availabledataformanyspeciesareoftenunsatisfactory,incomplete,orextremelyscanty.Thisisparticularlytruefortransientsandwinteringpopulations.InformationisgivensequentiallybystatefromNorthCarolinatoTexas.Wehavenotlistedstatesinwhichaspecieshasnotbeenrecorded.InformationfromFloridaisusuallypresentedintwosubsections,onedealingwiththeAtlanticcoast,theotherwiththeGulf.Thiswasdonebecausethestatusofaspeciesmayvaryconsiderablyfromcoasttocoast.InsomeinstancesasectiondealingwiththeKeysisalsoincludedwhen aspecies'statusthereisdifferentfromthatoneithercoast.10

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SYNOPSISOF PRESENTDISTKIBUTION AND ABUNDANCE Thissectioninthespeciesaccountssummarizesinformationgivenintheprevioussections,oftenwithadditionaldataonworld-widepopulationlevelsdependinguponourpresentknowledgeofthespecies.Forsomespeciestabularinformationispresentedaboutseasonaloccurrence,abundance,andareasofconcentrationinthesoutheast.Informationonthesetopicsisgenerallylimitedandshowswherefurtherdatashouldbeobtained.Twotypesofmapsindicatewhereconcentrationsofmarinebirdshavebeenreported;onedealswithbreedingbirds,theotherwithwinteringpopulations.Breedingcolonymapsarebasedonhighlydiversesourcesofinformation,includingtheColonialBird Kegister andaconsiderablenumberofpublishedandunpublishedcensuses(e.g.,Portnoy1977,ParnellandSoots1979ms).Otherdatawerefoundinrecentlypublishedpapersorwereobtainedfromlocalornithologists.Thesemapsgiveanestimateofthenumberofbreedingbirdsandindicatetheyearorperiodforwhichthisestimateapplies.Thelargestestimateavailableinthelastfewyearsisusedinsteadofarangeormeanbecauseestimatesarefewandbecausewewishtoemphasizeareasknowntocontainlargecoloniesorconcentrationsofcolonies.Notalldatawereplottedbecausesomespeciesoccurinsomanycoloniesthatitwasnotfeasibletoplotthemonour maps. The maps maycontainsomeinaccuracies.Theyarenotintendedasanatlas;theirprimarypurposeistoprovideanoverviewofwhereconcentrationsofbreedingmarinebirdsoccurinthesoutheasternUnitedStates.MostofthewinterdistributionmapsarederivedfromBystrak(1974),whobasedhismaps onananalysisofNationalAudubonSocietyChristmas Bird Countsforoneormoreoftheyearsfrom1970-1972.Wechose45of58coastalChristmasBirdCountsinthestudyareaandcompiled5-yearmeansfor1973-1977.Insomeinstancesfewerthan5yearsofcountswereavailableandthemeanisforashorterperiod.Localitieswerechosentoshowgeographicvariationinnumbersandtoemphasizewherethelargestconcentrationswerefound.Thefiguresshouldnotbeconstruedtoindicatethetruesizeoflocalpopulations.TheChristmasBirdCountsvaryconsiderablyintheamountofestuarine,coastal,andmarinehabitatcovered,butwetriedtoallowforthisbychoosingcountsthatcontainedthemostmarinehabitat.ThenumbersreportedinanygivenyearmaynotbeprecisebecauseofthelimitationsofChristmasBirdCounts.Thesemapsareintendedtoserveprimarilyasanindexofwherewinterconcentrationsaremostlikelytobefoundandhowthisdistributionvariesthroughoutthesoutheast.HABITATThissectiondealswiththenesting,feedingandnonbreedinghabitatsandtheoffshorehabitatsusedbythespecies.Theextentanddetailofinformationreportedisusuallygreatestfornestinghabitat,lessforfeedinghabitat,andleastfornonbreedinghabitatssuchasthoseusedbymigratingorwinteringbirds.WeoftencouldgiveonlysparseinformationforpoorlystudiedspeciessuchastheBonaparte'sGull,andfoundlittleadequateinformationabout11

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offshoreoccurrencesformanycommonspecies.HabitatsusedinthesoutheasternUnitedStateshavebeenemphasizedwhendatawereavailable;otherwiserelevantinformationfromotherareasisused.FOODAND FEEDING BEHAVIORWegiveatleastabriefgeneralstatementaboutfoodseatenandthefeedingmethods.Insomeinstancesweincludemoredetailedinformationonfoodhabits,brieflyabstractingrecentstudiesandindicatingproportionsofdifferentfoodseaten.FormostspeciesfewdataonfoodhabitsinthesoutheasternUnitedStatesaregiven,primarilybecauselittleisknownaboutthebirds'dietinthisarea.Consequently,datafromotherareashavebeengivenontheassumptionthatsimilarfoodsareeateninthesoutheast. Food habitshavebeensummarizedbygeographicareaforafewspeciesforwhichmuchrecentinformationisavailable.Forspecieswhosefoodhabitsarewelldocumented,wepointedoutdifferencesinfoodhabitsofadultsandyoung,andcommentedonseasonalvariationanddifferencesinfoodseatenindifferenthabitats.IMPORTANTBIOLOGICAL PARAMETERSIbis sectionpresentsbasicinformationtoallowbiologiststopredicttheeffectsofdevelopmentofoilresourcesonpopulations.Thesixspeciesforwhichwedonotgiveimportantbiologicalparameterseitheroccurlargelyasoffshoremigrants(RedandRed-neckedPhalaropes,ArcticandRoseateTerns)oraremuch moreabundantoutsidethesoutheasternUnitedStates(GreatBlackbackedGullandBrownNoddy).Muchoftheinformationcompiledinthissectionisderivedfromresearchconductedelsewherebecausefewadequatestudiesofthebreedingbiologyofcoastalseabirdshavebeenmadeinthe southeast. Foronespecies,theHerringGull,forwhichtherearemanydatafromboththeOldandNewWorlds,wehaveincludedonlyinformationfromNorthAmericanstudies.Thedatainthissectionconsistofbriefsummariesoftheegg-layingperiod,meanclutchsize,incubationperiod,hatchingsuccess,ageatfledging,fledgingsuccess,mortalityofeggsandyoung,renesting,ageatfirstbreeding,maximumnaturallongevity,andweight.Dataonegglaying,incubationperiod,andageatfledgingallowonetoestimateperiodswhenbirdsbreedingwithinthestudyareaaremostvulnerabletodisturbance.Clutchsizeandhatchingandfledgingsuccessaregoodindicatorsofproductivity.Informationonmortalityandrenestingindicatethepotentialforrecoveryfollowingalargenestingfailure.Figuresforknown maximumnaturallongevityallowacrudecomparisonbetweenspeciesoftheirtotalreproductivepotential.The maximumnaturallongevityisgivenintermsof"estimatedminimumage"inyearsandmonthsfollowingKennard(1975)andClappetal.(1982a),andmaylistinformationbasedonbandingrecordsfromtheUnitedStates,CanadaandtheOldWorld.Finally,informationonweightsisincludedbecausethis,andpopulationdatagivenelsewhereinthereport,willallowplannerstocomparespeciesintermsofbiomassaffectedbyanygivenoil-relatedactivity.12

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SUSCEPTIBILITYTOOILPOLLUTIONWeemphasizeoilingrecordsfromsoutheasternwatersbutmayhavemissedreportsofoilingforsomespecies.MuchoftheOldWorldliteraturereportsoiledbirdsonlybyspeciesgroups(e.g.,gulls,ducks)andcombinesinformationonindividualspeciesinthesetotals.SomeinformationmaybefoundinOld WorldregionalperiodicalsunavailableintheUnitedStatesandnotcoveredbycomputer-basedliteratureretrievalsystems.Foreachspecieswealsoprovideanestimateofthepotentialeffectofoilpollutionandthedevelopmentofoilresourcesinthesoutheast,takingintoaccounttheknownorsuspectedvulnerabilityofthespecies,itsabundanceinthesoutheast,anditsabundanceelsewhere. SPECIES BIBLIOGRAPHYAttheendofeachspeciesaccountisaspeciesbibliographywhichincludescitationsthatprovideadditionaldataonthetopicsbrieflycoveredinthetext,aswellasonvarious'otheraspectsofthebiologyofthespecies.AllcitationsinthetextappearintheLiteratureCitedattheendofthisreport.Thebibliographiesinthisvolumearenotexhaustive,butcovertheworldliteraturebecauselittleisknownofthebiologyofmarinebirdsinthecoastalsoutheasternUnitedStatesandbecausemoreextensivebibliographiesshouldbehelpfulinplanningfutureresearchonthesebirds.Asintheprecedingvolumesinthisseries(Clappetal.1982b,1982c),thebibliographiesstressdistribution,ecology,andbehaviorofthespeciesbutprovidemoreinformationontopicssuchasidentification,hybrids,diseases,andtheeffectsofpesticides.Oursearchoftheliteraturestressedrecentnessofinformation.Eachspeciesbibliographyshouldberelativelycompletethrough1981andmid1982.Somereferencespublishedmorerecentlymaybeincluded,buttheseandotherreferencesfrom1981and1982maynothavebeenusedinwritingtheaccount.Wealsolistedotherimportantpapersdealingwiththebiologyofthespeciesthroughtheearlypartofthecentury,andincludedolderreferencesthatarestillthemajorsourcesofinformationonthespecies.WeweremorecompletewithpaperswritteninEnglish.Thespeciesbibliographiesarearrangedfrompresenttopastwithauthorslistedalphabeticallyundereachyear,ratherthaninthemoreconventionalalphabeticalandchronologicallistingusedintheLiteratureCited.Wearrangedthemthiswaytomakeiteasierforthereadertofindthemostrecentinformation.Wecheckedallreferencesusedinthetextoftheaccountsaswellasmostoftheremainingreferences.Somecitationsfromsecondarysourcesremainunchecked.Weestimatethatthethreevolumesinthisseriescontainontheorderof10,000references,andourtemporalandfiscallimitationsweretoogreattoallowcompleteverificationofallreferences.13

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OILPOLLUTIONANDMARINEBIRDSOFTHESOUTHEASTERNUNITEDSTATESWiththepossibleexceptionofmarineturtles,marinebirdsarethevertebratesmostseverelythreatenedbyoilpollutionandthedevelopmentofoilresources.TheworkofOldWorldbiologistspresentsclearevidenceofsubstantialdamagetoseveralpopulationsofmarinebirds.SpecificinformationontheeffectsofoilingandoilspillsonavianpopulationsintheNewWorldisverylimited,especiallysoforpopulationsinthesoutheasternUnitedStates.SystematicsurveysofbeachedbirdsandseabirdkillshavebegunonlyrecentlyintheUnitedStates.Further,dataonoilingofmarinebirdsarescatteredthroughadiversebodyofliterature.Manydistributionalnotesreportingthefirstspecimenofaspeciesfromageographiclocalityparentheticallynotethatthespecimenwasoiled.OtherinformationisscatteredthroughregionaldistributionalworksandyetmoredatalieinthebandingandrecoveryfilesoftheBirdBandingLaboratoryoftheU.S.FishandWildlifeService.OilpollutionkillsmanythousandsofmarinebirdseachyearinDenmark(Riisgard1979)andelsewhere.Duringthewinterof1980-1981,fiveoilingincidentsinnorthwestEurope(Mead1981)ledtoanestimated60,000oiledbirdsonthebeaches(MeadandBaillie1981).Mostofthesebirdswerevariousspeciesofauks,butmanyBlackSeaters(Melanittanigra)werekilledinWesternDenmarkandmanylive,oiledBlack-leggedKittiwakes(Rissatridactyla)wereseeninHollandandBelgium(MeadandBaillie1981).ThesespillsprobablykilledmoremarinebirdsinwesternEuropethanhadanyincidentsduringtheprevious12years(Anon.1982a).AnotherrecentspillthatwashedashoreontheMagdalenIslandsintheGulfofSt.LawrenceoffeasternCanadakilledmorethan1,500seabirds,mostofthemevidentlyDovekies(AIleaIle)andmurres(Uriaspp.)(Anon.1982b).----OilinghasbeenamajorfactorinreducingpopulationsofCommonEiders(Somateriamollissima)intheDanishWaddensea(Joensen1973),CommonEidersandBlackScotersinHolland(SwennenandSpaans1970),andAtlanticPuffins(Fraterculaarctica)inFrance(Bourne1976).OtherexamplesofsignificantreductionsinavianpopulationsduetooilpollutionaregiveninreviewsbyBourne(1968a,1976),Croxall(1975),VermeerandVermeer(1975),andtheFoodandAgriculturalOrganizationoftheUnitedNations(1977).Smallerbutsignificantlosseshavealsobeenrecordedinlocalizedpopulations.Anestimated25-50%oftheCommonLoons(Gaviaimmer)winteringinShetland,offScotland,diedfollowingtheESSOBERNICLAoilspill(StoweandMorgan1979).AlllocalMallards(Anasplatyrhynchos),European Coots (Fulicaatra),andMoorhens(=CommonGallinule,Gallinulachloropus)diedfollowinganoilingoftheAmerRiverintheNetherlands;itwasestimatedthatapproximately88%oftheGreylagGeese and71%oftheBewick'sSwans(Cygnuscolumbianusbewickii)alsowerelost(Belterman1972).Duringthewinterof1981-82an air-sPill inBelgiumkilledabout300birdsandaffectedasmanyas3,000,mostlygullsandtheentirelocalpopulationofLittleGrebes(Tachy-baptusruficollis)(Anon.1982c).---14

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Despitecontinuingreportsofhighseabirdmortalityfromoilpollution,particularlyinthecolderwatersoftheNorthAtlantic,currentopinionisthatthelong-termeffectsofoilspillsontheenvironmentmaybescant.TheRoyalCommissiononEnvironmentalPollution(1981inBourne1982)statedthat"Wehaveconcludedthatoilspillsareunlikelytocauselong-lastingdamage;theenvironmenthasaremarkablecapacityforrecoveringfromeventhelargestoilspillages. We havefoundnoclearevidencethatoilspillshavesignificantlyaffectedpopulationsofseabirdsorothermarinespecies Theshorttermconsequencesofoilpollutionareseriousandaretypifiedbythepollutionofbeachesontheonehand,anddamagetoseabirdsontheother"Clark(1982)pointedoutthatalthoughtensofthousandsofmarinebirdsdieannuallyfromoilpollutioninwesternEurope,naturalmortalityaccountsforhundredsofthousandsperyear.Nonetheless,Nettleship(1977)believedthatoilpollutionoftheseastillconstitutesthesinglelargestthreattoseabirds.VARIABILITY AJl0NG SPECIESINSUSCEPTIBILITYTOOILPOLLUTIONSurveysofbeachedbirdsarebiasedindicatorsoftheproportionofapopulationaffectedbyoiling(Bourne1976,PowersandRummage1978).RecentstudiesbyManometBirdObservatory(PowersandRumage1978,Powersetal.1980)documenteddifferencesbetweenthespeciescompositionofbirdsfoundoiledonbeachesandbirdsfoundoiledatsea.InDecember1976andinJanuary1977,181oiledbirdswerefoundonthebeachesofNantucketIslandandMartha'sVineyard,Massachusetts,followingthespilloftheARGOMERCHANT.Mostofthesebirdswerealcids(48.1%),gulls(27.1%)andloons(18.2%),but7seaducks,2NorthernGannets(Sulabassanus),2cormorants,and1grebewerealsofound.Most (92%of1120)ofthebirdsseenoffshoreweregulls;ofthese,59%,41%and9%,respectively,oftheHerringGulls,GreatBlackbackedGulls,andBlack-leggedKittiwakes(Rissatridactyla)wereoiled.BothNorthernGannetsandNorthernFulmars wereseenoffshoremorefrequentlythanobservedonthebeaches,makingup6%and1%,respectively,ofthebirdsseenoffshore.Twelvepercentofthegannetsseenandatleastsomeofthefulmarswereoiled.Alcidsmadeuponly12%ofthebirdsseen,butonly12oiledDovekies(AIleaIle)wereobserved.PowersandRumage(1978)pointedout,however,thatoilonthelargeralcidscouldhavebeenoverlookedeasily.Althoughbeachedbirdsurveys(Table1)givesomeideaofdifferencesinsusceptibilitytooilingbetweengroupsofbirds,therecentworkbyPowersetal.(1980)suggestshowdifferentthesituationmaybeoffshore.Offshoresurveysmaybemorereliableforindicatingthemagnitudeoftheoilpollutionprobleminagivenareaandforprovidingusefuldataontheincidenceandextentofoilpollution.Speciessuchasloons,grebes,auks,andseaducksarefrequentlykilledbyoil.Others,suchasgulls,arefrequentlyoiledbutdiemuchlessoftenfromthedirecteffectsofoiling(Table2).Terns(Table1,Bourneinlitt.)areseldomfoundonbeachedbirdsurveys,andiftheydodieareperhapslesslikelytobefound.Theyevidentlyseldomdiefromthedirecteffectsofoiling(Table2).BlackSkimmersareprobablyintermediatebetween15

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Table1.Numberandpercentageofbeachedbirdsexaminedthatwereoiled(a).GreatBritainTotal%foundoiledKindsofbirdsLoons(Divers)GrebesAlbatrossPetrels(c)NorthernFulmarShearwatersStorm-petrelsGannetsCormorantsBrownPelicanWildfowlPhalaropesJaegersKittiwakesGullsTernsSkimmer Auks 152 54 337 182 21811372448 6171 94 591750 45 76 30 80SouthOregon-AtlanticCoastWashingtonCaliforniaUnitedStatesCoast(b)CoastTotal%Total%Total%foundoiledfoundoiledfoundoiled114 4 3 33 175LO14 64 14 36 798 5 0 0 80 0 2 50O?0 570 28 301 4 14 0 0 623 22 0 4 25 40 0 61760 0130.517 0 38 0514 26 92 296 7 0 119 3 1 0 0 80 0 L052133 241310163111972370 0 1 0 0 104 94 2848 19(a)DataforGreatBritain,thesouthAtlanticcoastoftheUnitedStates,theOregon-Washingtoncoast,andtheCaliforniacoastarefromBourne(1976),MalcolmSimons(inlitt.),Harrington-Tweit(1979),andAinley(1976),respectively;theperiodscoveredare1968-1970,December1977August1978,mid-winter1976,and1971-1975,respectively.Dataforthesoutheasterncoastthrough1 December 1977arebasedonsurveysfromCapeHatteras,NorthCarolina,toCapeCanaveral,Florida,thereaftersouthtoJensenBeach,Florida.(b)MostmortalityoffulmarsandBlack-leggedKittiwakesfollowingthisincidentwasnotbelieved.tobe duetooil;mostwildfowlandalcidmortalitywasattributedtooiling(Harrington-Tweit1979).(c)AlthoughBourne(1976)didnotspecify,histerm'petrels'probablyincludedallProcellariidae(e.g.,petrels,shearwaters,fulmars)andmayhaveincludedHydrobatidae(storm-petrels).Histerm'gulls'probablyincludedallLaridae(gullsandterns).Forothermaterialsummarizedhere,'petrels'referstoPterodroma,'shearwaters'toPuffinus,'gulls'toLarus,and'terns'toSterninae.16

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Table2.Susceptibilityofvariouslaridstooilingasindicatedbybandingrecoveries(a).Numberrecov-SpecieseredLaughingGull2,269(Larusatricilla)Ring-billedGull23,531(Larusdelawarensis)CaliforniaGull2,658(Laruscalifornicus)HerringGull34,713(Larusargentatus)WesternGull2,916(Larusoccidentalis)Glaucous-wingedGull18,979(Larusglaucescens)GreatBlack-backedGull1,426(Larusmarinus)CaspianTern2,390(Sternacaspia)RoyalTer-n---4,230(Sternamaxima)Common Ter;:;-56,067(Sternahirundo)SootyTern30,461(Sternafuscata)Brown Noddy 942 (Anousstolidus)BlackSkimmer 927(Rynchopsniger)Numberdeadfromoiling4196588233 3 514112Percentdeadfromoiling0.180.080.230.170.270.120.210.130.120.030.0030.110.22Areasfromwhichdeadoiledlaridswerereported(b)NewJersey,Louisiana,Texas,PanamaOntario(4),NewYork(3),Florida,NorthCarolina,Michigan(2)California(4),Washington(2)NewYork(10),Massachusetts(8),Ontario(5),Newfoundland,Texas(4)California(6),Washington,OregonBritishColumbia(9),Washington(7),Oregon(4)NewYork(2),QuebecCalifornia(2),MichiganVirginia,NorthCarolina,SouthCarolina,FloridaCaribbean(5),Brazil(2),Venezuela(2),NewJersey(2)FloridaCaribbeanFloridaa)ThistableisderivedfromaprintoutproducedbytheBirdBandingLaboratorythatlistedallbirdswhoserecoverythrough11April1978wasbelievedduetooiling,andfromClappetal.(1982a)whichgivesthenumberofrecoveriesforeachspeciesprocessedbytheBBLthroughDecember1981.Thepercentagesmayunderestimatethenumbersrecoveredduetooiling,butsuggestthedegreetowhichoilingcausesdirectmortality.NoFranklin'sGulls(250recoveries), Roseate Terns(1,286),ArcticTerns(291),orWhiteTerns(Gygisalba)(376)werereporteddeadfromoiling.b)Forspeciesrecoveredinlargenumbersweonlylistareasfromwhichthemostoiledbirdswerereported.17

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thesetwogroupsintheirsusceptibilitytooiling.REGIONALANDSEASONALDIFFERENCESIN OILINGANDMORTALITYOFBEACHEDBIRDSAlthoughbeachedbirdsurveysintheeasternUnitedStateshavebeenconductedforonlyashorttime,theextentofoilingamongbirdsfounddeadalongthesouthernAtlanticcoastappearslowcomparedwithotherareasintheUnitedStatesandEurope.Only4%of400birdsfounddeadalongthesoutheasternAtlanticcoastfromJanuary1976throughAugust1978wereoiled.Incontrast,oilingoccurredon82%of667birdsfoundalongthePolishBalticcoastfromNovember 1974toAugust1975(Gorskietal.1977),on26%of162foundalongIrishcoastsfromDecember 1977toMarch 1978(O'Keeffe1978),on79%of3,431foundontheinternationalbeachedbirdsurveysinNorthwestEuropeinJanuaryMarch 1975(Lloyd1976),andon18%of2,420foundalongtheCaliforniacoastin1975(Ainley1976).BirdmortalitypermileofbeachalsotendstobelessinthesoutheasternUnitedStatesthaninotherareas(Table3).Mortalityfiguresforaheavilypollutedarea,thePolishBalticcoast(3.2birds/kmor5.1birds/mi;Gorskietal.1977),areconsiderablyhigherthanforanywhereinthesoutheast.OtherareasinnorthwesternEuropevaryconsiderablyinmortalityrecordedduringbeachedbirdsurveys;thesemortalitiesareusuallygreaterthanthosefoundinthesoutheasternUnitedStates,butresemblethosefoundalongtheNorthAtlanticcoastoftheUnitedStates.Lloyd(1976)reportedarangeinthenumberofdeadbirdsfoundof0.17birds/km(0.3birds/mi)inpartofFranceto4.06birds/km(6.5birds/mi)inWest Germanyduringthewinterof1975.ForGreatBritain,1968-70,theaveragewas1.3birds/km(2.1birds/mi)(Bourne1976),similartomortalityinthesoutheasternUnitedStates.Mortalityduring1976-1980alongthesoutheasternAtlanticcoastandFloridaGulfaveraged1.3birds/mi(0.79birds/km)and1.2birds/mi(0.77birds/km),respectively(Simons1981ms).MortalityalongtheCaliforniacoastisalsogreater than inthesoutheast;surveystherereportedanaverageof3.5birds/mi(2.2birds/km)from1971to1975(Ainley1976).ThedisparitybetweenbeachedbirdmortalityratesinCaliforniaandEuropeandthesoutheastmayarisepartlyfromdifferencesinprevailingwindsandcurrents.InpartsofNorthAmericawhereprevailingwindsblowoffshore,mostmortalityisfoundaroundenclosedinlets.OnislandsoffshoreinwesternNorthAmericaandinnorthwestEurope,whereprevailingwindscarrydyingbirds(andoil)toshore,bothchronicoilpollutionandtherecordedmortalityofmarinebirdsisgreater(Bourne1976).Thecolderwinterclimatesintheseareasarealsoalmostcertainlyafactorintheincreasedmortality.BothoilpollutionandbirdmortalityaregreatestinnorthwestEuropeduringthecoldestpartoftheyear(Bourne1968a).Oiling(andpresumablymortality)ofmarinebirdsoffthenorthAtlanticcoastoftheUnitedStatesisgreatestduringwinterandspring(Powersetal.1980);mortalityisalsogreaterduringwinterandspringalongthesouthAtlanticandGulfcoasts(Tables3,4).18

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Table3.Comparisonofregionalandseasonalvariationofbeachedbirdmortalityand incideT.lce ofoilingintheeasternUnitedStates(a).AtlanticCoastAtlanticCoastNofCape SofCapeFloridaGulfTexasGulfHatterasHatterasCoastCoastDead Dead Dead Deadbirds/%birds/%birds/%birds/%PeriodmileoiledmileoiledmileoiledmileoiledSPRINGMar.-May19822.196 1.32 0.02.143.330.0Mar.-May1981 1.48 34.3.577.7 1.00 0.02.95 11.1 Mar.-May1980 1. 520.00.552.75Mar.-May197951.40.9220.0.820.0Mar.-May 197866.8(b)1.580.0 1.77 2.000.0Mar.-May19772.505.50.950.0.750.0SUMMERJune-Aug.19812.100.0.150.00.270.0June-Aug.19804.270.0.390.0.390.02.800.0June-Aug.19794.40 1.2 0.385.60.530.03.00100.0(c)June-Aug.19786.370.0 1.43 0.01.300.0June-Aug.19776.810.90.140.00.50FALLSep.-Nov.19812.174 1.07 2 1.47 0.00.400.0Sep.-Nov.1980 1.481.041.00 Sep.-Nov.19790.9813.41.430.00.590.00.360.0Sep.-Nov.19781.050.01.490.01.005.64.000.0Sep.-Nov.19770.240.00.600.01.250.00.750.0WINTERDec.-Feb.81-822.513.9 1.54 0.90.0Dec.-Feb.80-81 1. 7612.5.290.01.390.0Dec.-Feb.79-80.57 1. 56Dec.-Feb.78-792.192.31.84 1.1 1.740.01.000.0Dec.-Feb.77-782.706.52.87 1.4 3.401.000.0Dec.-Feb.76-779.335.51.750.02.880.019

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Table3.Concluded.(a)ThiscomparisonisbasedonBulletinsoftheAtlanticandGulfCoastBeachedBirdSurveyProjectandSimons(1981ms,inlitt.).Thesedata,whileuseful,areincompleteandhavesometimes(particularlyinTexas)beenbasedonsurveysofsofewmilesofbeachthattheresultsobtainedareprobablynotcomparablefromregiontoregion.Dashesindicatelackingdata.Severalfiguresfromthefirsttwovolumesofthisserieshavebeencorrectedandsomeadditionalinformationisgiven.(b)ThishighfigureistheresultofanoilspillintheChesapeakeBayinFebruary1978.(c)ThishighfigureispresumablytheresultoftheIXTOCoilspillinthesouthernGulfofMexico.Table4.Comparisonoffive-yearaveragesofbeachedbirdmortalityfordifferentareasintheeasternUnitedStates(a).Deadbirdsfoundpermileonthe:Five-AtlanticAtlanticAverageyearCoastNCoastSFloridaforperiodofCapeofCapeGulfentireSeasonendingHatteras HatterasCoastareaSpring19822.791.381.12(Mar.-May)Summer19806.310.610.83(June-Aug.)Fall19800.801.220.93(Sep.-Nov.)Winter19813.702.572.25(Dec.-Feb.)2.081.003.04(a)ThesefiguresarecompiledfromBulletinsproducedbytheAtlanticandGulfCoastBeachedBirdSurveyProject.ToofewdataareavailablefortheTexasGulfcoastforthesetobemeaningful.ThehighaveragemortalityinsummerinthenortheasternUnitedStatesispartlytheresultofdisturbanceofnestingcolonies.20

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Dataonoccurrenceandseasonalvariationofoiledbirdsatseaarelargelyunavailable,butapreliminaryreportbyPowersetal.(1980)ofat-seaobservationsfromFebruary1976toDecember1979intheGeorgesBankareaoffMassachusettsprovidessomeusefulinformation.Themajorityofoiledbirdsseenweregulls.Insomemonths(JanuaryandFebruary)gullsmadeupnearlyalloiledbirdsseen,butveryfewoiledgullswereseenduringthesummer(June-August).OiledNorthernFulmarswereregularlyseeninspringandsummer.OiledGreaterShearwaters(Puffinusgravis)wereseeninJuneandOctober.Oiledseaducks(CommonEider [SomaterIalnollissima], Red-breastedMergansers[Mergusserrator],White-wingedScoters[Melanittafusca]andOldsquaws [CIangUfa hyemalis])wereseeninDecemberand GannetsandPomarineJaegers(Stercorariuspomarinus)wererarelyoiled. BIRD KILLSFOLLOWINGOIL SPILLSIN THE SOUTHEASTERNUNITEDSTATESThefewreportsoflargebirdkillsfollowingoilspillsinsoutheasternwatersareusuallyinadequate.A,typicalexampleoccurredinlateDecember1968,when abargespilledcrudeoilalongthecoastofWakullaCounty,Florida.Thisresultedin"manyduckssnipeandotherbirdssocoveredwithoilthattheywereunabletofly.Smallerbirdswereunabletowalkintheheavyoil"(CenterforShort-LivedPhenomena1969).Reportsofmorerecentspillsarenobetter.Duringthespringof1981,39oiledCommonLoons(Gaviaimmer)werepickedup onthebeachesofPeaIslandNWR,NorthCarolina.Despit-e---treatment,only2survived(Simons1981).Howmany morediedatseaorwhatotherspecieswereaffectedisunknown. A"largetoll"ofwinteringCommonLoonsandRed-throatedLoons(Gaviastellata)wastakenbyanoilspillnearMyrtleBeach,SouthCarolina(Anon.1981).Inyetanotherincident,morethan175,000gallonsofheavyfueloilspilledintoamarshontheCapeFearRiverinSouthCarolina.ThisspillwasfollowedbytheindirectoilingofBrownPelican(Pelecanusoccidentalis)eggswhenoiledadultsreturnedtoincubatetheireggs.Whetherthisoilingaffectedthecolonyorotherwaterbirdsnestingnearbyisnotknown(Anon.1982d).WeknowofonlytwoinstancesofmajoroilspillsinornearthesoutheasternUnitedStatesforwhichthereisevenfairinformationaboutthenumberandspeciesofbirdskilled.ThefirstoftheseoccurredinearlyFebruary1976inthelowerChesapeakeBay.FollowingthesinkingofabargenearthemouthofthePotomacRiver,spilledfueloilresultedinthewidespreadcontaminationofmarshesandbeaches,andthedeathofanestimated20,000to50,000birds(Rolandetal.1977).Perryetal.(1979)madeestimatesforeachspeciesthatdiedduringthisspill,fivespillsthatoccurredintheDelawareRiver,andanotherlargespillinChesapeakeBay.Mostofthevictimsofthesevenspillswereducks(69.02%ofca.52,500birds),HornedGrebes(Podicepsauritus)(27.75%),andCommonLoons(1.30%).Fewcharadriiformsdied(315HerringGulls,30GreatBlack-backedGulls,10Ring-billedGulls,andoneAmericanOystercatcher(Haematopuspalliatus).21

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ThesecondmajormortalityfollowedthespillinTampa Bayofsome80-100tonsofoilwhentheGreektankerDELIANAPOLLONranagroundinmid-February1970andruptureditshull(Wallace1970,Clark1973).Oilspreadingoverthebayresultedinatleast4,500birdsbeingbroughttocleaningandrehabilitationstations(Sims1970).Asmanyas9,000birdsmayhavedied(Clark1973).Thebirdsbroughttocleansingstationswerelargelyducks,CommonLoons,andHornedGrebes(Sims1970).Severalofthesoutheasternstatesareamongthosewiththegreatestnumberofoilspillsandamountofoilenteringtheecosystemyearly(Table5).However,solittleinformationontheeffectsofoilingonseabirdsofthisareaisbeingrecordedandreportedthatitisimpossibletoassessadequatelytheoveralleffectofoilpollutioninthesoutheast.SOURCESOFVARIATIONINMORTALITYFROMOILPOLLUTIONAlargenumberoffactorsareinvolvedindeterminingthemagnitudeofdetrimentaleffectsofoilpollutiononmarinebirds.Birdsoiledincoldweatherorcoldwaterhavea muchhigherfatalityratethandothoseinwarmweatherandwarmwater.Evensmallamountsofoilmayleadquicklytodeathunderthestressofcold(Levy1980),butbirdsinwarmerareasmaysurvivethesamedegreeofoiling(R.Clapp,pers.observ.;C.Harrison,pers.comm.).ReportsfromEurope(BourneandBibby1975,Riisgard1979)indicatethatmortalityfromoilingisgreaterinwinterthaninsummer.Oilspilledincoldwaterremainsliquidlongerthaninwarmwaterandislikelytocausemore damageasaresult.Itfirstformsa"chocolatemousse"water-in-oilemulsionandthenitformstarballs.Althoughtheseformsofoilmaypresentsomehazardtobirds(BourneandBibby1975),thehazardisapparentlymuchlessthanwithfreshoil.Bourne(1976)summarizedsomeofthechangesindaily,annual,andlifecyclesofmarinebirdsthatmayincreasetheirvulnerabilitytooilpollution.Localcurrentsandwindsmaybringdriftingslicksintoraftsofbirdsroostingonthewater.BourneandDevlin(1969)suggestedthatmostmortalityfromoilingoccurswhenroostingorfeedingbirdsaretrappedbydriftingslicks.Birdsthatconcentrateinflocksof1,000ormoretoscavengebehindfishingtrawlers(e.g.,gannetsandgulls[Powersetal.1980]),maybedrawnintotheareaofanoilspill,causingdisproportionatelylargenumberstobeoiled.Conversely,oiledbirdsmayfollowtrawlersawayfromthesiteofaspillandthusbeunderrepresentedamongthesickanddyingcomingashore(PowersandRumage1978).Bourne(1976)pointedoutthatmarinebirdsareparticularlysusceptibletodamagefromoilwhentheyaremolting.Whenbirdslacktheirusualinsulation,smallerthanusualamountsofoilmayleadtodeathfromchilling,shock,andstarvation.22

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Table5.OilspillsinthesoutheasternUnitedStates1978-1980comparedwithotherstates(a).PercentPercentPercentofoilPercentNumberofspillsofallVolumespilledofoilofoiloccurringU.S.spilledinthespilledspillsintheoil(gallons)south-inU.S.State1978-80southeastspills1978-80eastwatersSoutheasternstatesNorthCarolina2972.4SouthCarolina2822.3Georgia2592.1Florida2,01816.2Alabama 4343.5Mississippi1511.2Louisiana5,66445.5Texas3,35026.9Total-Southeast12,455100.1OtherheavilypollutedstatesNewJersey1,052Massachusetts792NewYork1,081California2,249Kansas120Kentucky508Missouri292 RhodeIsland253Alaska798Pennsylvania800Total-U.S.Waters30,7771.0139,1651.30.4 0.9105,7911.00.3 0.8474,2064.41.56.6507,6934.71.61.4518,8294.81.60.565,7500.60.218.45,325,74749.316.5 10.93,662,89633.911.340.510,800,077100.033.43.43,921,30812.22.61,992,6556.23.51,334,9704.27.31,083,1293.4 3.9901,0962.81.7861,3292.70.9809,9872.50.8700,1052.22.6639,8602.02.6614,1291.932,207,039(a)ThistableiscompiledfromU.S.CoastGuard(1980,1981)data.Otherheavilypollutedstateschosenforcomparisonwerethosethathadthegreatestamountofoilspilledfrom1978through1980.Thefiguresfor1980werepreliminaryfiguresonly.ThefigureforTexasprobablyissomewhatinflatedduetotheIXTOCblowout.23

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Breedingbirdsareparticularlysusceptibletooil.Thelossofonememberofapairmay meancompletelossoftheirreproductivepotentialforthatyear.Althoughthislossmayberecoupedinfuturegenerations,mostmarinebirdshavelowproductivityandpopulationstakeyearstorecoverfromasingleoilingincident.Oilinthevicinityofbreedingcoloniesalsomaydiminishreproductivesuccessbydecreasingthehatchingsuccessofcontaminatedeggs,bydisturbancetothecolonyfromattemptstocontrolpollution(Bourne1976),andbydebilitatingoiledbirdssomuchthattheymaynotattempttobreed(Stowe1982).Fordetal.(1982)recentlydevelopedamathematicalmodelthatdealtwiththeeffectsofoilspillsonbreedingpopulationsofseabirds,usingguillemotandkittiwakepopulationsintheBeringSeaasparadigms.Theirmodelsuggeststhatacatastropicmortalityofadultsrequiresalongerrecoverytimeforapopulationthandoesasimilarmortalityofyoung.Acompletebreedingfailurebyguillemotsinoneyearwouldhaveasmallereffectonrecoverytimethanwoulda5%mortalityofadults.Althoughtheseremarksarebasedonalcids,agrouphighlysusceptibletooilpollution,theauthorspointedoutthattheirresultsprobablyareapplicabletootherspecieswithhighadultsurvivalrates.Highadultsurvivalrateshavebeenshownforseveralspeciesthatoccurinthesoutheast,NorthernFulmar,ManxShearwater(Puffinuspuffinus),HerringGull(Dunnet1982),andCommonTern(DiCostanzo1980),andarelikelytooccurinspeciesofgullsandternsbreedinginthesoutheast.Fordetal.(1982)alsomodeledtheeffectsofchroniclow-levelpollutionfollowingaone-timeadultmortality(asmightbecausedbyanoilspill).Theirresultsindicatedthatsmalldecreasesinadultsurvivalorfecunditygreatlyincreasedtheamountoftimerequiredfortheaffectedpopulationtorecover.Theyalsopointedoutthat"Chroniclow-levelpollutionmayproducepermanentchangesinthesurvivorshipandfecundityschedulesofapopulation,whichcanaltertherecoverytimeandextinctionpointforone-timeperturbations."Fordetal.(1982)concludedbystatingthattheirmodelshouldberegardedasnomorethanapreliminaryestimatethatpredictsonlythemagnitudeofapopulation'sresponsetoanoilspill.Theyindicatedthatmoreprecisepredictionsarenotpossiblewithoutadequatedata.Fewobservationsofthebehaviorofbirdsencounteringoilhavebeenreported.Availableinformationindicatesthatdifferencesinbehaviorbetweenspeciesaffecttheirlevelofvulnerabilitytooil.AccordingtotheInternationCouncilforBirdProtection(1960),Long-tailedDucks(=Oldsquaw,Clangulahyemalis)choosetolandonoilslicks.Iftrue,thismayaccountforsomeoftheveryhighoil-relatedmortalitiesreportedforthisdivingduck.Guillemots(=CommonMurres,Uriaalgae)divetoescapefloatingoilbutsuffertheriskofsurfacingintoitandthusbecomingseverelycontaminated(Bourne1968b).Hainard(1959)reportedthatsomedivingducks(TuftedDuck[Aythyafuligula)andPochard[A.ferina])avoidedpatchesofoilfloatingdown ariver.Gulls(Bourne1968b) and ManxShearwaters(Puffinuspuffinus)(Casement1966)alsoactivelyavoidthicker,morenoticeableoilslickswhenflying.Someofthesebirdsalsoavoidoilwhenswimming;aHerringGullandaBlack-leggedKittiwake(Rissatridactyla)that apatchoffloating24

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oilimmediatelytookflight(Bourne1968b,BourneandDevlin1969).Sanderlings(Calidrisalba)andWillets(Catoptrophorussemipalmatus)oiledontheTexascoastspentlesstimefeedingandmoretimeroostingandmakingcomfortmovementsthandidunoiledbirds(Chapman1981).Thenumberofbirdsthatdiefollowinganoilspillisalsorelatedtothetypeofpetroleumspilledandhowlongitremainsintheenvironment.Crudeoilislesstoxicthanrefinedoils(dieseloil,No.2fueloil,Bunker"C")(Hay1979),andfreshoilcausesmoredamagethanolder,weatheredoils(BourneandBibby1975).Someoilsmaybeinnocuousenoughthatoiledbirdsarenotkilledandareevencapableofcleaningtheirplumage(Birkheadetal.1973,Phillips1974).Thenumberofdeathsfromoilingfollowingaspillisnotnecessarilyrelatedtotheamountofoilspilled;largespillsmayresultinfewdeaths,whilesmallerspillsmaycauselargelosses,particularlywhensubstantialnumbersofbirdsareconcentratedinsmallareas(Croxall1975,Salomonsen1979).Inaddition,largeoilsp'i11smaycausenogreaterlossofmarinebirdsthandoeschronicoilpollutionoftheenvironment(Nelson-Smith1973,Croxall1975,HolmesandCranshaw1977).EFFECTSOFOILONCONTAMINATEDBIRDSANDTHEIREGGSTheprimaryeffectofoilonbirdsisalossofbuoyancyandinsulationwhentheplumagebecomesmatted.Oiledbirdsthenmayquicklychillanddiefromexhaustionandexposure.Ingestionofoilbybirdsalsoleadstoavarietyofphysicalandphysiologicaldisordersthatmakedeathmorelikely(Clappetal.1982c,EastinandMurray1981).Experimentalstudiesalsohavedealtwiththesecondaryeffectsofoilingonreproductionofmarinebirds.Smallamountsofoilwillreducethehatchingsuccessofduck,heron,gull,tern,andaukleteggs(EastinandHoffman1978,StickelandDieter1979,Ainleyetal.1981,HoffmanandEastin1981).Toxicityofoilsisgreaterforneweggsthanforthosefurtheralonginincubation,andolderweatheredoilsarelesstoxicthanfreshones.Oilingofincubatinggullscausessignificanteggmortalitywhentheoiledfeatherscomeincontactwiththeeggs.Oilingofeggsalsomayresultindeformedchicks.Deformedbills,incompletelyossifiedwingorfootbones,abnormallysmallliverlobesandstuntingwerethemostcommonabnormalitiesfoundinexperimentalstudies(StickelandDieter1979).OtherstudieshaveshownthatthenumberofeggslaidbyMallards(Anasplatyrhynchos)decreaseswhentheyarefeddietscontaining2.5%SouthLouisianacrudeoil(EastinandHoffman1978,StickelandDieter1979).Ducklingsfeddietscontaining5%ofthiscrudeoilgrowmoreslowlythanusualandmaydevelopanumberofinternaldisorders(EastinandHoffman1978).Theseamountsofoilseemlarge,butotherworkhasshownthatsmallerdoses,similartowhatmightbeencounteredinnature,mayalsohaveadverseaffectsonhealthandreproductivebiology.25

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HerringGullchicksatLittleDuckIsland,Mainewerefed1 mldosesofacrudeoil/cornoilmixture.Chicksfed0.2and0.5mlofweatheredSouthLouisianacrudeoilgrewmoreslowlyfor7-9daysaftertreatmentthandidcontrolsfedonly1 mlofcornoil(Butleretal.1978).Chicksfed0.2mlofcrudeoilrecoveredwithintwoweeks,butthosefed0.5mlshowed areducedgaininweightforaboutthreeweeksaftertheinitialdosing.Bothgroupsfedcrudeoilalsoexhibiteddecreasedculmengrowth.Nodifferencesinbehaviorwerenoted.Peakalletal.(1978)triedtodeterminewhichcompoundsincrudeoilinhibitedgrowth.HerringGullchickswerefed1 mloftwoSouthLouisianacrudeoils,orthealiphaticoraromaticfractionscontainedwithin1 mlofeach.Onecrudeoilcausednosignificantreductioninweightgain,butboththecrudeoilandaromaticfractionsoftheothercausedsignificantdecreasesinthegrowthrate.Hallett(inPeakalletal.1978)foundthatpolynucleararomaticswith3ormoreringswerepresentintheoilthatcausedreducedgrowth,butnotintheother.Thisvariationintheadverseeffectscausedbytwopresumablysimilarcrudeoilssuggeststhattheeffectsonlocalbreedingpopulationsencounteringoilspillsmaybehighlyvariableandnotpredictablebythedegreetowhichthebirdsbecomecontaminated.AdditionalinformationonexperimentalworkshowingtheeffectsofoilonmarinebirdsofthesoutheastisgiveninthespeciesaccountsforGreatBlack-backedGull,LaughingGull,andSandwichTern.POTENTIALHAZARDSTOMARINEBIRDSFROMOFFSHOREOILPRODUCTIONAbouttwo-thirdsoftheoilincoastalwaterscomesfromrunoffandeffluentfromterrestrialsources.Tankeroperationsaccountforabout26timesmoreoilinmarinewatersoftheUnitedStatesasdooffshoreoperations(Ohlendorfetal.1978),butmaycauseadisproportionatelylargeshareofavianmortality.Ohlendorfetal.(1978)suggestedthat,forthemarineenvironment,itmaybesafertoproduceoiloffshorethantoimportit.Widespread,severeeffectsofchronicsmallspillsatproducingoffshoreoilfieldshavenotyetbeenfound(DicksandHartley1982)andthesefieldsmaycauserelativelyfewlong-termbiologicalproblems(Dicks1982,DicksandHartley1982).The samemaynotbetrueforchronicdischargesincoastalwaters,particularlyatspecificsiteswhereoilisproduced,handled,andrefined.Theretheeffectsmaybelocalandsubtleorsevereandlong-lasting(DicksandHartley1982).LongleyandJackson(1980)reviewedtheproblemscausedbythedevelopmentofpetroleumresourcesinbrackishmarshesandsuggestedalleviativemeasuresthatcouldbetaken.Effectsincludedirectlossofvegetationandanimals(e.g.,bydredgingconstructionofpipelinesandroads);additionofdissolved,particulateandtoxicmaterialstotheenvironment;andchangesinwaterflows.Changesinwaterflowwereconsideredthemostdamaginghazardbecausetheymayresultinthecompletelossofamarshecosystem.Suchaneventcouldbeaccompaniedbyareductionoraneliminationofthepopulationsofmarinebirdsthatusethehabitatfornestingorfeeding.26

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Similareffectsarelikelywhenonshoreareasandoffshorebarrierislandsareaffectedbydevelopmentofoilandgasresources.Changesinwaterflowduetodredgingcouldeasilychangetidalandcurrentpatterns,resultingintheeliminationofislandsusedfornesting.Terrestrialaccesstolargerislandsmayresultintheintroductionofpredators(e.g.,foxes,raccoons)thatcouldeliminateanentirebirdcolonyinaseasonortwo.Disturbanceengenderedbyconstructioncouldresultinthemassdesertionofatraditionalbreedingareabysomespecies.Geologicalcharacterizationsofthesensitivityofshorelinestooilspillsarenotsufficienttodeterminetheoverallsensitivityoftheseareas(OwensandRobilliard1981).OwensandRobilliardpointedoutthatsometimeandsite-specificfeatures(e.g.,presenceofendangeredspecies,migrationsoffish,seabirds,waterfowl,andshorebirds)arelargelyoverlookedinsuchconsiderationsandmay make asitehighlysensitiveforonlyshortperiodsoftime.Theystatedthat"thevulnerabilityofmobile(whetherresidentormigratory)marinebirdspeciesisprobablymoresignificantthantheimpactofoilonthenearbyintertidalhabitats."TheyfurtherpointedoutthatthethreatthattheIXTOCIblowoutpresentedtotheegglayingoftheendangeredKemp'sRidleyseaturtlealonganisolatedTexasbeachwouldnothavebeenidentifiedbytheusualprocessesofdeterminingsensitiveareas.Severalrecentreportsreviewedhumanactivitiesrelevanttodevelopmentofonshoreoilfacilities.ThesereportsincludeMulvihilletal.'s(1980)detailedreviewoftheeffectsofshorelinestructuresonthecoastalenvironment,Morton's(1976)reviewoftheecologicaleffectsofdredging,andBuckleyandBuckley's(1976a,1977a)andBurger's(1981a)reviewsoftheeffectsofhumandisturbanceoncoloniallynestingbirds.Onshorehabitatchangeorlossresultingfromthedevelopmentoffacilitiesrelatedtooffshoreoilproductioninthelongrun,probablywillmoreadverselyaffectwaterbirdsinthesoutheasternUnitedStatesthanwilltheproductionofoilitself.Thespeciestreatedinthisvolumecomposethegroupmostvulnerabletopetroleumdevelopmentinthesoutheast.Developmentofonshorefacilitiestoproduce,transport,andholdoilwillprobablybeamajorsourceofenvironmentaldisruptionanddisturbanceforthesebirds.Otherformsofcoastaldevelopment(e.g.,recreationandrealestate)alsomaycauseseriousdeclinesinpopulationsofthespeciesincludedhere,andmayhavemoreseriousconsequencesthandevelopmentofpetroleumresources.Furtherresearchintotheflexibilityintheuseofbreedinghabitatisneededtodeterminetheextenttowhichdevelopmentmay damagepopulationsbut,asBuckleyandBuckley(1980a)pointedout,sucheffortswillhavelittlevalueunlessmanagementeffortsincludemassivehabitatprotection.Asecondaryhazardtobreeding ofthesoutheastistheingestionofoil-contaminatedfood.Theimpactmaynotbeimmediate,butdeleteriouseffectsonthehealthofadultmarinebirdsandontheproductionofyoungcanbesevere.27

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RECOMMENDATIONSFORFUTURERESEARCHThisreportrevealslargegapsinthebodyofknowledgenecessarytodealeffectivelywithproblemsrelatingtomarinebirdsandOCSdevelopment.Belowaresomeoftheproblemsthatwebelieveneedmostattention.STATUSANDBIOLOGYOFBREEDING SPECIES Thesizesofbreedingpopulationsofmarinebirdsinthesoutheastarestillpoorlyknown,yetlimiteddatasuggestthatthisareacontainssignificantnumbersofsomespecies(Table6).Recentsurveys(SootsandParnell1975a;Portnoy1977,1978ms;Blacklocketal.1978,1978ms;ParnellandSoots1979ms;Portnoyetal.1981)providevaluableinformationonthesizesofpopulations,butsurveysareincompleteformanyareasandhavebeentakenovertooshortaperiodtoallowfortheannualvariationusuallyfoundinpopulationsofmarinebirds.InformationfromcoastalareasofGeorgiaandSouthCarolinaisinadequate,andinformationfromFloridaisavailableonlyforportionsofthestate.Inaddition,thelocationsandsizesofbreedingcoloniesofconspicuousbreeders(e.g.,SandwichandRoyalTerns)thatoccupyrelativelyfewnestingsitesarebetterknownthanthoseofotherspecies(e.g.,LaughingGulls,Forster'sTerns)whosecoloniesaremorenumerousandwidelydispersed,butarefoundinlesseasilysurveyedhabitats.Speciesthattendtonestinassociationwithotherspecies(e.g.,Gull-billedandCaspianTerns)areoftenoverlookedonsurveysandtheactualnumbersbreedinginthesoutheastarenotwellknown. Manyrecentpapershavedealtwiththedifficultiesofcensusingwaterbirds(Nettleship1976;HarrisandLloyd1977;Buckleyetal.1978b;BirkheadandNett1eship1980;Drury1980;Erwin1980a,1981;ErwinandOgden1980;Hutchinson1980;Portnoy1980).Perhapstoomuchemphasishasbeenplacedonobtaininghighlyaccuratecensuseswhenobtainingconsistentresultsovertimeismoreimportant.Erwin(1980a)pointedoutthat"themostimportantfactorinobtainingsystematicsamplingresultsisprobablyobserverconsistency"andthatusingthesameobserversoveraperiodofyearsallowsacomparisonthatemphazizesrelativeratherthanabsolutechangesinnumbers.IncommentingoncensusesconductedinAlaskaandNewEngland,Drury(1980)statedthat"oneshouldnotdrawconclusionsbasedondatagatheredinthecourseofscatteredsurveysmadeoverafewyears,nomatterhowpreciseandthoroughthecountingofeachsample,"apointwithwhichweagreethoroughly.Druryalsoremarkedthatusefulconclusionscanbedrawnfromvariedsurveysif(a)estimatesweremadesystematically,(b)iftheoverallchangeinnumbershasbeenconsiderable,and(c)ifthebiologicalreasonsforperiodicchangesareknownandunderstood.Hepointedoutthatvariationsinnumbersbetweensurveyscanbedealtwithbycalibratingfordifferencesbetweenobservers,countingtechniques,andcountstakenatdifferenttimesofdayandindifferentpartsofthebreedingseason.28

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Table6.EstimatedpopulationsofcharadriiformmarinebirdsbreedinginthesoutheasternUnitedStatescomparedwiththeirstatusinotherareasadministeredbytheU.S.SpeciesLaughingGullHerringGullGreatBlackbackedGullGull-billedTernCaspianTernRoyalTernNumberbreedinginthesoutheastca.263,000ca.500ca.20ca.3,000ca.3,000ca.99,000WorldbreedingrangeMostlyNorthAmericaWidespreadintheHolarcticregionsAtlantic,NorthAmericaandeasternEurasiaNearlycosmopolitanWidespreadbutpatchyMostlyNorthAmerica29SignificanceofpopulationsinthesoutheasternU.S.comparedwithotherareasunderU.S.jurisdictionAmajority(ca.65%)oftheU.S.populationandprobablymostoftheworldpopulationbreedsinthesoutheast;mostoftheremainderbreedalongtheNorthAtlanticcoast.TheyalsobreedintheCaribbean,includtheVirginIslands,butthesizeofpopulationsbreedingthereispoorlyknown.Negligible.Wellover100,000birdsbreedalongtheNorthAtlanticcoastofNorthAmericaandintheGreatLakesregion.Negligible.MostoftheNorthAmericanpopulationbreedsalongtheNorthAtlanticcoastwiththemajoritybreedinginMaineandMassachusettsThesoutheastcontainsabout95%ofallGull-billedTernsbreedingintheU.S.MorethanhalfofthesoutheasternbirdsarefoundinTexas.OthersbreedintheCaribbeanbutwhetheranybreedinareasadministeredbytheU.S.isnotknown. ThesoutheastholdsoverafifthoftheU.S.population.MostofthesoutheasternbirdsbreedinTexas.Over90%oftheU.S.populationbreedsinthesoutheast.

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Table6.Continued.NumberbreedingintheSpeciessoutheastSandwichTernca.75,000RoseateTernca.400(a)CommonTernca.9,000Forster'sTernca.24,000LeastTernca.26,000 Horld breedingrangeMostlyNorthAmericaandEurope Harm andtemperatewatersoftheworldMostlyineasternNorthAmericaandthepalearcticoftheOld Horld NorthAmericaLargelyinthewesternandeasternU.S.30SignificanceofpopulationsinthesoutheasternU.S.comparedwithotherareasunderU.S.jurisdictionOver95%oftheU.S.populationbreedsinthesoutheast.SomecoloniesinLouisianaareamongthelargestknown.AnunknownnumberalsobreedsintheU.S.VirginIslands.Negligible.MostoftheRoseateTernsbreedinginareasadministeredbytheU.S.areintheAtlanticnortheastandintheU.S.VirginIslands.Slight.TheonlysignficantpopulationbreedsinNorthCarolinaandrepresentsperhapsabout10%ofthebirdsbreedingintheeasternU.S.BreedingstatusalongtheGulfcoastandintheCaribbeanisuncertain.MostoftheU.S.populationbreedsaroundtheGreatLakesandintheAtlanticnortheast.Nearly70%oftheU.S.populationandprobablymorethanhalftheworldpopulationbreedsinthesoutheast.About80%ofthesoutheasternpopulationbreedsinLouisiana.About60%oftheU.S.populationoccursinthesoutheastwherethelargestknowncoloniesofthespeciesarefound.ConsiderablenumbersalsonestalongtheU.S.NorthAtlanticcoast.OthersbreedintheCaribbeanandintheU.S.VirginIslands,butpopulationstherearepoorlyknown.

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Table6.Concluded.SpeciesBridledTernSootyTernBrown NoddyBlackSkimmer NumberbreedinginthesoutheastNoneca.80,000ca.3,000ca.59,000WorldbreedingrangePantropicalexceptmuchofPacificPantropicalPantropicalNorthandSouthAmericaSignificanceofpopulationsinthesoutheasternU.S.comparedwithotherareasunderU.S.jurisdictionBreedsintheU.S.VirginIslandsandelsewhereintheCaribbeanbutlittleisknownoftheirpopulationsFloridaholdstheonlysizeablepopulationbreedinginthecontiguousUnitedStates.ThousandsbreedintheCaribbean,includingtheU.S.VirginIslands,andintheMarianaIslands.HundredsofthousandsbreedinHawaiiandAmericanSamoa. TheonlysizeablepopulationbreedinginthecontiguousUnitedStatesisinFlorida.UnknownnumbersalsobreedintheU.S.VirginIslandsandmanythousandsbreedinHawaii,AmericanSamoa,andtheMarianaIslands.Over85%oftheskimmersbreedingintheU.S.arefoundinthesoutheastandthisareaundoubtedlycontainsamajorityoftheNorthAmericanpopulation.About70%ofthepopulationintheeasternU.S.breedsinLouisianaandTexas.(a)Recentcomments byW. B. Robertson,Jr.(pers.corom.)suggestthatthenumbernowbreedinginFloridaisevenlessthanthefiguregiveninthespeciesaccount.31

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The datacurrentlyavailable,whileprovidingsomeideaofbreedingpopulationsofsoutheasternmarinebirds,donotadequatelyassessthetotalnumberofbirdsusingtheareanordotheyprovideadequateinformationaboutthesizeofnonbreedingpopulations,theseasonalandannualvariationsinpopulations,orthehabitatstheyuse.Managementdecisionsbasedonpresentinformationmayerrbecauseavailabledatadonotreflectconditionsapplicablewhenthedecisionsneedtobemade.Acontinuingcommitmenttothemonitoringofmarinebirdpopulationsinthesoutheast(andelsewhere)isneeded.Nothinglesswillprovidetheinformationnecessarytomakejudiciousdecisionsthatwillbenefitboththemarinebirdandhumanpopulationsthatcompetefortheresourcesofthecoastalzone.Dataonregionaldifferencesinnestingchronology,degreeofannualvariationinreproductivesuccess,factorsinfluencingnest-siteselection,determinantsofcolonylocation(particularlyinrelationtofoodresources),andotherdemographicparameters,arenecessaryforsatisfactoryevaluationoftheeffectsofmanagerialdecisionsonthewell-beingofpopulations.Studiesshouldbeundertakenoveraperiodofseveralyears;thoseconductedduringasingleseasondonotprovideenoughinformationformostmanagerialpurposes.AstheNationalEnvironmentResearchCouncil(NERC)(1977)pointedout,"such[long-termstudiesare]essentialasabaselineagainstwhichtheresultsoffuturestudiesorenvironmentalimpactscanbemeasured."ThebreedingbiologyofsomeofthespeciesinthisreporthasbeenstudiedinNorthAmerica(e.g.,Ring-billedGull),intheOldWorld(e.g.,Gull-billedandBlackTerns),orinboth(e.g.,HerringGull,CommonandRoseateTerns),butfewofthecharadriiformsbreedinginthesoutheasthavebeenstudiedthere.OnlytheSootyTernandtheLaughingGullhavebeenrelativelywellstudiedinthesoutheast,andmuchremainstobelearnedofthebiologyandnumbersofthelatter.SomeinformationisavailableonthebreedingbiologyofLeast Terns andBlackSkimmersinthesoutheast.Royal,Sandwich,Forster's,andGull-billedTernsarelargelyunstudiedinthesoutheastandthelatterthreehavebeenlittlestudiedanywhereintheUnitedStates.VirtuallynothingisknownaboutthebreedingbiologyofCaspianTernsinthesoutheastorofthestatusandreproductivebiologyoftheCommonandRoseateTerns.Thelatterisdeclininginmanyportionsofitsrangeandhasbeenrecommendedforendangeredspeciesstatus(BuckleyandBuckley1981).Onewayofpredictingtheeffectsofoilspillsonlocaltodevelopmathematicalmodelsthattakeintoaccountvariousmetersthatdeterminetheresponseofpopulationstospills.todothisbyFordetal.(1982)indicatedthatthefollowingthosemostneededforsuchmodels:populationsisbiologicalparaArecentattemptparametersare1)thesizeofthenonbreedingpopulation,2)themovementpatternsofforagingindividuals,3)thespatialandtemporaldistributionandavailabilityoffoodnearbreedingcolonies,4)therelationshipbetweenfeedingratesandtheageofyounginrelationtogrowthratesandsurvivalprobabilities,32

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5)thedegreeofdensitydependenceinvariouspopulationparameters,6)theprobabilitythatagivenbirdwilldieasthedirectresultofanoilspill,7)theage-specificmortalityschedulesoflocalpopulationsundernormalconditions,8)therateatwhichpopulationsrespondtoperturbationsandregainanequilibriumdistributionatsea,and9)theeffectofanoilspillontheshort-termavailabilityoffood.Fordetal.(1982)thoughtthatthelastfourparametersarethemostimportantfordevelopingthemodel,butnoneareeasytomeasureandsomerequiremassivelogisticalsupport.Futureresearchonspeciesbreedinginthesoutheastshouldbedirectedtowardsansweringsomeofthesequestions.DISTRIBUTION OFTRANSIENT,WINTERING,ANDPELAGICPOPULATIONSOurknowledgeofthedistribution,numbers,andbiologyofcharadriiformmarinebirdsoccurringoffshoreorduringnonbreedingperiodsisevenmorelimitedthanourknowledgeofthestatusofbreedingspecies.TheBridledTernisoneofthecommonestofoffshorespecies,butlittleisknownofitsareasoforiginorbiology.LargeportionsoftheNorthAmericanpopulationsofBlackTernsandFranklin'sGullsmigratethroughthestudyarea.TheareaisalsoamajorwinteringgroundforLaughing,Bonaparte's,Ring-billed,andHerringGulls;Caspian,Royal,andForster'sTerns;andBlackSkimmers.Weknowlittleabouthabitatusebythesewinteringpopulations,nordoweknow muchabouttotalpopulations. With theexceptionoftheSootyTern,verylittlebandinghasbeendoneintheenormousmarinebirdcoloniesofthesoutheast;inconsequencescientistsusuallyhavenoideaoftheroutesbywhichtheydispersefromthesebreedingcolonies.Amassbandingprogram,judiciouslyundertaken,shouldaddgreatlytoourknowledgeoftheirdispersal.Suchworkshouldbeconductedoveraperiodofyearsandbeintegratedwithamarkingprogramthatwouldallowtherecognitionofindividuals.Such aprogramcouldalsoprovideusefuldataontimingofmigrationthroughthearea,aswellasprovideinsightintoforagingranges,localmovements,andvariousaspectsofbreedingbiology.Amongthecharadriiformspeciesnestinginthesoutheast,LaughingGulls;Caspian,Royal,Sandwich,andForster'sTerns;andBlackSkimmerswouldprobablybethespeciesmosteasilytreatedbythisapproach.Presentknowledgeofpelagicbirdsinthesoutheastispoor.Onlyafewpelagicsurveyshavebeenconductedinthisareawithanythoroughnessandregularity.StudiesbyDavidLeeinNorthCarolina,CharlesDuncanand Ralph HavardinAlabama,andbyJohnJohnsoninFlorida,havebeenonlypartiallyreportedandtendtoconcentrateonrarebirdsandthoseseenwelloffshore.DuncanandHavard'srecent(1980)studyrevealedthatspeciesliketheBluefacedBooby(Suladactylatra)andBridledTern,formerlythoughtrareinthenorthern infactmajorcomponentsofthepelagicavifauna.33

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Comprehensiveanddetailedoffshoresurveysofmarinebirds,conductedbyboatatleastmonthlyandoveracarefullychosengrid,wouldhelpimmenselyinplanningcontingenciestobetakenfollowingoilspills.Suchsurveysshouldlastatleast2yearsbecausedatafromanygivenyearmaynotrepresenttypicalconditions. aerialstudiesofthisnature(e.g.,Frittsetal.1982ms)helpdelineatetheextenttowhichoffshoreareasareusedbymarinebirdsandatleastshouldresultinusefulinformationabouttheamountofbiomassinagivenareaatagiventime.Onelimitationofthisapproachisthelowreliabilityofidentificationtospecies.Frittsetal.(1982ms)werenotabletodistinguishfromtheairbetweenfourspeciesofSterna(Forster's,Common,Arctic,andRoseateTerns),norcouldtheyusuallydistinguishbetweenspeciesofstorm-petrels,tropicbirds,jaegers,orphalaropes.TheirreportmentionsneitherCaspiannorGull-billedTernsalthoughbothmusthavebeenpresentininshoreareas.Aninexpensivewaytodeterminethepotentialeffectofoildevelopmentwouldbetomakeobservationsfromoilrigsonspeciescompositionandbehaviorofpassingtransients.Somespecies,suchastheboobies,areprobablyattractedtotheseplatforms,becausetherigsmayconcentratelocalfoodresources.Internationalbordersarebiologicallyimaginarylinesthattendtodistortourknowledgeofthedistributionofbirds."Foreign"birdsmayformalargeproportionofthebiomassofmarinebirdsinsoutheasternwatersforpartoftheyear.Oneofthelargestoilspillsinhistoryfollowedtheblowouton 3June1979oftheIXTOCIwellofPetroleosMexicanosinthesouthernGulfofMexico.Eventuallyover3millionbarrelsofpetroleumwerelostandanoilslickthatspreadacrossthesoutherngulfbyAugustwashedashorealong80km(50mi)oftheTexascoast(Waldichuk1980).Althoughonlyabout20birdswerefounddeadfromoilinginthisarea(Garmon1980),severalofthesewerepelagicspeciesseldomseenalongtheTexascoastbutcommoninthewatersoftheU.S.GulfofMexico(Clappetal.1982b).ThesedeadbirdsincludedfiveBlue-facedBoobiesandoneBrown Noddy(B.Chapman,pers.comm.),neitherofwhichiscommonofftheTexascoast.Thelowtollofmarineseabirds,oneofthegroupsknowntobemostseverelyaffectedbyoiling,isnotsurprisingsincetheTexascoastissome 800km(ca.500mi)northoftheblowoutsiteintheBayofCampeche(atca.19020'N,92025'W).Whatissurprising,however,isthatnoneofthefollow-upreportsaddressedtheproblemofwhathappenedtoseabirdcoloniesnearertheblowout.ThreeareasneartheIXTOCblowout(CayosArcas[200l3'N,9lo58'W],CayosArenas[22007'N,91024'W]andArrecifesAlacran[AlacranReef][22023'N,89040'W]),presentlyorformerlysupportedpopulationsofbreedingorroostingseabirds.Theclosestofthese,CayosAreas,isonlysome 45 mi(75km)fromthewellheadoftheIXTOCIspill.Itheldsome5,000Blue-facedBoobiesinlate1952.Another400wereseenonCayoArenas.Thisspecies,theMagnificentFrigatebird(Fregatamagnificens)andtheRoyalTernareallknowntobreedon CayoArcas(Paynter1955).AlacranReefsupportsbreedingpopulationsofBlue-facedBoobies,BrownBoobies(Sulaleucogaster),andMagnificentFrigatebirds.AlacranReefmayalsosupportnestingpopulationsofRoyalandSandwichTerns(Boswall1978).Boswall(1978)reportedabout2,000Blue-facedBoobiesand2,000BrownNoddiesin34

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September1975.HenotedthatthousandsofSootyTernsapparentlynestonthereef.Intheearly1950'ssome 800 BrownBoobieswerepresentandasmanyas5,000MagnificentFrigatebLrdsbredthere(Paynter1955).ThenumberssuggestthisareaisanimportantsourceforthepelagicavifaunaoftheGulfofMexico.SeveralofthemarinebirdstreatedhereandinClappetal.(1982b)breedintheCaribbean.Audubon'sShearwater(Puffinuslhermineri)andtheBridledTern,periodicallycommontoabundantoffoursoutheasterncoasts,breedintheU.S.VirginIslandsandelsewhereintheCaribbean.TheCaribbeanmaybeanimportantsourcefortheseaswellasforotherspecies(e.g.,Blue-facedBooby,MagnificentFrigatebird,SootyTern)commonintheoffshorewatersofthesoutheast.SpeciesofmarinebirdsthatbreedorhavebredintheU.S.VirginIslandsincludeBlue-facedBoobies;BrownBoobies;Red-footedBoobies(Sulasula);MagnificentFrigatebirds;White-tailedTropicbirds(Phaethon lepturus); Red-billedTropicbirds(Phaethonaethereus);Sandwich,Sooty,andBridledTerns;andLaughingGulls(DeweyandNellis1980).TheRoseateTernpopulationisoneofthelargestremainingintheCaribbean.TheU.S.VirginIslandsareoneofthefew(perhapstheonly)areasunderU.S.jurisdictionwherebreedingAudubon'sShearwaters,Red-billedTropicbirds,andBridledTernsmaybefound.ColoniesofmarinebirdsintheVirginIslandsaremuchindangerfromintroducedandferalanimalsandfrompoachers(DeweyandNellis1980).WhatinformationisavailablemakesclearthattheunoccupiedislandsoftheCaribbeanareoneofthemainrefugiaformarinebirdsinthewarmerwatersoftheworld.WerecommendthateffortsbemadetoinitiatecooperativeinternationalsurveysofmarinebirdpopulationsinthesouthernGulfofMexicoandintheCaribbean.Internationalsurveyswouldsupplynotonlya muchbetterunderstandingoftheoverallstatusofthespeciesinvolved,butalsowouldpermitfarbetterinsightintotheconsequencesoflocalmanagerialdecisionsonaspeciesthroughoutitsrange.Previouseffortsalongtheselines,particularlywithregardtowaterfowl,havebeenhighlyeffectiveinproducingtheinformationneededtomanagepopulations.EFFECTSOFOILONSOUTHEASTERN MARINE BIRDSWebelievethatattemptedrehabilitationofoiledbirdsfollowingamajoroilspillislargelyawasteoftime,money,andotherresources.Agroupofmarinebirdexperts(NERC1977)statedthat"sincetheresultsofattemptstorehabilitateoiledbirdsaresopoor,itmaybe moreprofitabletoexpendeffortsatpreventingbirdsfrombecomingpolluted."Itisdesirable,however,tosalvageoiledbirdstofindoutwhatspecieswereaffectedandtoobtaininformationthatwillallowformoreprudentresponsestofuturespills.SatisfactoryeffortstoadequatelydocumentlossestooilspillsintheUnitedStatesarealmostnil.Thisinformationisoftendifficulttoobtainbecausecountsofdeadorcontaminatedbirdsmaynotindicatethenumbersthatwereaffectedorwilldie.Seldomisthereinformationonthenumberofbirdspresentinanareabefore,during,andaftercontamination.Whileitmaybenext35

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toimpossibletodo morethanestimatenumberspresentbeforeaspill,rapidresponsestooilingincidentsshouldallowsomeideaofthenumberspresentduringandafteraspill.Detailedobservationsmadeduringandafterthespillmayallowbetterestimatesoftheseverityofthespill.Otherfactorsmakeestimationofthedamagecausedbyspillsmoredifficult.Forexample,thetimeofanoilslick'spassagethroughanareamaybecrucialindeterminingthedegreeofcontaminationandmortalityexperiencedbyeachspecies.DuringthecontaminationoftheFirthofForthinFebruary1978,oilpassednearthemainfeedingareaforwaterbirdsatnight;consequently,therewas aproportionatelygreaterlossofnight-feedingGreaterScaup(Aythyamarila)andPochards(Aythyaferina)thantherewasofCommonGoldeneye clangula)andCommonEiders(Somateriamollissima),mostofwhichhadmovedelsewheretoroost(Campbelletal.1978).Theproportionofbirdsoiledfollowingapollutionincidentmayvarywidelybetweenspecies,dependingonthehabitatsusedandthehabitsofthebirds.Theprobabilityoffindingmostoiledbirdsthatroostorloafonshoreneartheiroffshorefeedingareasisgreaterthanitisforfindingbirdsthatspendallormostoftheirtimeoffshoreandthat,followingoiling,sinkfromsightnevertobeseenagain.Further,wind,offshorecurrents,andmovementsbythebirdsthemselvesmaytakemostofthevictimsofanoilspillfarfromwheretheywereoiledbeforeanyonenoticestheirplight.InsomepartsofEuropeandonthewestcoastoftheUnitedStates,prevailingwindsbringvictimsofoilingtoshore.Incontrast,ontheAtlanticseaboardwindstakeoiledbirdsouttosea.Consequently,comparisonsofdamagefromoilpollutionincidentsbetweentheseareasarenotvalid.Likewise,wecannotuseestimatesofmortalityfrombeachedbirdsurveysinEuropetopredicttheincidenceofmortalityinthewesternAtlantic.Atbest,theEuropeanreportssuggestthatdamagetowildbirdsfromoilontheU.S.eastcoastmaybegreatlyunderestimated.PowersandRumage(1978)documentedonespilloffMassachusettswellenoughtodemonstatedifferencesbetweenonshoreandoffshoreobservations.FollowingtheARGOMERCHANTspill,prevailingwindsandtidesmadeitunlikelythatoiledbirdswouldwashashoreandledPowersandRumagetosuspectthatthefewbirds(181)foundoiledonthebeachesofNantucketIslandandMartha'sVineyardhadmadeanactiveefforttogetashore.HundredsofoiledGreatBlack-backedandHerringGullswereseenoffshoreduringat-seasurveysfollowingthisspill(Powers,pers.comm.).OiledNorthernFulmars,aspeciesnotfoundalongthebeaches,werealsoseen.Mostoiledbirdswereseennearthetankeroroilslick,butoiledgullsandBlack-leggedKittiwakeswerewidelydispersedduetotheirpenchantforfollowingfishingtrawlers.Levy(1980)analyzedtheoilfoundondeadormoribundbirdsinAtlanticCanadaandsuggestedthatHerringandGreatBlack-backedGullsobtainednearSableIsland,NovaScotia,hadbeencontaminatedbyoilfromtheARGOMERCHANTspillthathadoccurredsome 840km(520mi)away.36

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Despitethedifficultiesinobtainingunbiaseddataontheeffectsofoilspills,westillrecommendthatbettereffortsbemadetomonitorandpublishreportsoftheeffectsofthesespillsonmarinebirds.MuchoftheinformationneededtoanswerquestionsrelatingtooilpollutionandmarinebirdsinthesoutheasternUnitedStateswouldbeavailableifsucheffortshadbeenmadeinthepast.Wealsorecommendthatmoreattentionbepaidtomonitoringlong-termeffectsofoilpollutioninthesoutheast.Oneofthebetterandlessexpensivewaysinwhichthismaybeaccomplishedisaperiodiccensusofbirdsfounddeadalongthebeaches.Thislendssomeobjectivebasistospeculationsabouttheeffectsofoilpollution,andalsoprovidesinformationaboutunusualorincreasingmortalityfromothercauses(e.g.,pesticides).Overtime,thismayserveasanearlywarningindicatorofseriousproblems.SuchsurveysareconductedintheeasternUnitedStatesbytheAtlanticandGulfCoastBeachedBirdSurveyProject,buttheareacoveredinsomeregionsisverysmall(e.g.,2 mioftheTexascoast[M.Simons,pers.comm.]).ACKNOWLEDGMENTSManypeoplecontributedtothisreportinavarietyofways.GeneW.BlacklockallowedustousehisunpublishedmanuscriptontheoccurrenceandstatusofthebirdsofPadreandMustangIslandsandsupplieduswithunpublishedmaterialontheTexasColonialWaterbirdCensus.WilliamB.Robertson,Jr.andHerbertW.Kale,II,provideduswithunpublishedsummarymaterialonFloridabirds.MalcolmM.Simons,Jr.,DirectoroftheAtlanticandGulfCoastBeachedBirdSurveyProject,suppliedunpublisheddataonincidenceofoilingandmortalityofbirdsalongtheAtlanticandGulfcoasts.ThelibrarystaffoftheSmithsonianNaturalHistoryMuseum wasespeciallyhelpfulinobtainingcopiesofmostofthethesesandmanyofthepapersthatmake upthefilesuponwhichthisandthecompanionreportsarebased;CarolynS.Hahn,AmyE.Levin,andJackF.Marquardtwereextremelyhelpfulinthisregard.Agnes C.NalleyalsoaidedusinfindingandobtainingliteratureintheBirdLibraryoftheGabrielsonLaboratoryofPatuxentWildlifeResearchCenter.LindaA.Hollenberghelpedassemblespeciesbibliographiesandtheliteraturefiles.InitialversionsofthemapswerepreparedbyMartha B. HaysandcompletedbythestaffoftheNationalCoastalEcosystemsTeam,Slidell,Louisiana.WethankHelenL.Harbett,TracyChevalierandLindaM.Wolfewhotypedsomeofthepreliminaryandfinaldraft.LindaM.Wolfewasalsoveryhelpinproofingfinaldraft.WealsothankCharlotteI.AdamsonwhopreparedtheillustrationsthatinterspersethetextandWilliamC.Whitewhosetechnicalassistanceinkeepingourfalteringwordprocessorsfunctionalwentfarbeyondtherequirementsofhisjobandismuchappreciated.JeffreyA. Bpendelow, 37

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JillParkerandCherryKellerprovided some much needededitorial assistance and we thankthelllfortheirefforts.Wealso thenk FrancineG.and PsulA. Buckley,R. KicheelErwin, MalcolllC.Coulter, Martha B.Hays, W .IS. Robertson,Jr., kalph W.Schreiber,and Richard L.Zusi whoco_nted on vsrious speciesaccountsaDdoftenpointedout infOl:lll3.tionthst wehad missed. TheseindividualsoftenwerethesourceoflIuchvaluableunpublisheddata.Early versions ofseveralspeciesaccountswerepreparedby Wayne A.Hoff man, whoalsosupplied infot1llation onunpublishedsightingsofmarinebirdsintheGulfof Mexico. CharlesD. Duncan andThomaa H. Fritts ptovidedunpublishedinformationforthisarea.Othersalsosuppliedunpublishedmanuscriptsorcopiesof pal)ers, journals,Otrepottsthatwouldhaveothetwisebeenverydifficulttoobtain,andprovidedguidanceastoareaswhereadditionalinforlllationcould be acquired.Fortheseandotherservices,wethankDavidG. Ainley, HansBlokpoel, &irik A.T.Blom,W.R.P.Bourne,DannyBystrak,BrianR. Chapman, W.FrankCobb,Jr.,MercedesS.Foster,PatrickJ.Gould,CraigS.Harrison, Jerome A.Jackson, M. Kathleen Klilllc.1ewicz,koxie C.Laybourne, Kary K.LeCroy,DavidS.Lee,KartinK. Mc.Nicholl, IanC.T.Nisbet,StorrsL.Olson,SalllPatten,Jr.,JohnW.Portnoy,KevinD.Powers,ChandlerS. kobbins, LarryR. Shanks, C.Sibley,David H. Smith,JeffreyA.Spendelow,Judith A. Toups,JacobValentine, George1:;. Watson,Ill;JohnS. Weske, ClaudiaP.Wilds,andDonaldW.Woodard.ThisprojectwassponsoredbytheBureauofLand Management '8OuterContinentalShelfOffice(nowcalledtheMineralsManagementService).--AnoiledadultLaughingGullatFlamingo,Florida.PhotographbyRoger8.Clapp.38

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KED-uECKEDPHALAKOPE (Phalaropuslobatus) lOA: Odinshane, DU: Grauwefranjepoot, EN: Ked-neckedPhalarope,FI:Isovesipaasky, FK: Phalaropeabecetroit,Phalaropehyperbore, GE: Odinshuhnchen,Odinswassertreter,SchmalschnabligerWassertreter,IC:Odinshani,IT:Falaropobeccosottile,JA:Aka-erihire-ashi, NW: Svommesnipe,po:Platkonogrdzawoszyi,PK:Falarodo,KU:(Kound-nosedPhalarope),SP:Falaropopicofino,Chorlillonorteno, SW: Smalnabbadsimsnappa,US:NorthernPhalarope] GENEKALDISTKIBUTIONHorth America Ked -neckedPhalaropesbreedinlowArcticandSubarcticNorthAmericafromthePribilofandAleutianislandseastandnorththroughnorthernAlaska,thenceeastalongthenorthernpartofthecontinentthroughsouthernVictoriaIslandandcentralKeewatintosouthern Baffin Island.TheybreedsouthtosouthernAlaska,southwesternYukon,andnorthwesternbritishColumbia,andfromnorthwesternMackenziesoutheasttonorthernSaskatchewan,Manitoba,andOntario.TheyalsobreedeastonislandsandthecoastofJames Hay throughnorthernQuebectotheeastcoastofLabrador(AOU1957,Godfrey1966).Ked-neckedPhalaropesmigrateinconsiderablenumbersalongandoffthecoastsofNorthAmerica(AOU1957,Godfrey1966)andarewidespreadmigrantsintropicalAtlanticwaters(Watson1966).Theyalsomigratethroughtheinteriorofwestern North America(AOU1957)andregularly,butinmuchsmallernumbers,through the interiorfarthereast.TransientshavebeenrecordedinCentralAmericainGuatemala,Honduras,CostaKica,andPanama (Blake 1977). DistributionRed-neckedPhalaropesbreedintheOld WorldinsouthernGreenland,Iceland(Holm1965),theFaeroes,ScotlandandIreland(rarely) (BOU 1971),inNorwayandSwedensouthtoabout62 N, andinSpitsbergenandFinland,locallysouthinthelattertoabout64N.ThisspeciesalsobreedsintheU.S.S.R.innorthern Estonia andinnorthernKussiaonthenorthernKolaPeninsulaandKandalaksha Bay easttotheTaimyrPeninsula(toabout72N)andeastalongthecoasttotheChukotskiPeninsula.FromnorthernSiberiaitbreedsasfarsouthasabout64-66N,andeasttotheCommanderIslands(Vaurie1965).EuropeanpopulationsfromsoutheasternScandinaviaandnorthernKussiamigratesoutheasttothePersianGulf;birdsfromOldWorldpopulationsfartherwestmigratesouthalongthewestcoastof Europe towinteroffAfrica(Hohn1966,HildenandVuolanto1972).Taxonomicnote:ThisspecieshasoftenbeenplacedinthegenusLobipes."NorthernPhalarope"isthecommonnameusedinmostoftherecent Amercanliterature,butweprefertousethename"Red-neckedPhalarope",followingtheusageadoptedbytheforthcomingAOUchecklist.39

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Ked-neckedPhalarope Both Oldand New Worldpopulationswinteratsea,largelyintropicalandsubtropicalwaters.Inthe l,ew World,thespecieswintersinlargenumbersoffthePeruviancoast (Nurphy 1936),whereitoccurschieflyontheouterfringes-oftheHumboldtCurrent (Blake 1977). Blake (1977)considereditcasualinthecoastalwatersofChilesouthtoSantiago,butothers(AOU1957,Vaurie1965)thoughttheWinteringrangeoffPacificSouthAmericawasfromtheGalapagosandEcuadorsouthtoChileandsouthernArgentina.ThesephalaropesalsowintercommonlyoffthesouthAtlanticcoastofSouthAmerica (Hohn 1965,1969).Ked-neckedPhalaropeswinterintheOld WorldoffwestAfrica,Arabia,andinthewatersofsoutheastAsia (BOU 1971);inthewesternPacifictheyoccurinlargenumbersoffJapan(OSJ1974),andwinterfromtheKyukyussouthandeasttonorthern [,ew Guinea(Vaurie1965),wheretheyarenotuncommon (Condon1975).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATES North CarolinaPearsonetal.(1942)consideredKed-neckedPhalaropesraretransientsalongthecoast,butrecentobservationsshowtheyareactuallynUmerousoffshoreduringmigration.Thisspecieshasbeenrecordedmorethan70timesin Horth Carolina(Leeand llooth 1979)withmostsightingsfrom10Augustthrough25Octoberandfrom3 Hay through10June.Weknowofapproximately50publishedrecordsoftheKed-neckedPhalaropefrom Horth Carolina,butwelistbelowonlythose-thatinvolvefourormorebirds.190923Sep.5seen(3coll.)at\IhiteLake,BladenPearsonetal.County1942 196213Sep.30seen22-25mi EHatteras InletGrant 19631966 4Sep.5seenoffHatterasInletParnell1967a197227 Hay 30seenatPeaIslandTeulings1972c1972 8Oct.45seenoffHatteras 'l'eulings 1973a1973 26 Hay 30seenoff Norehead CityTeulings1973e 1973 2Sep.4seenoffHatterasTeulings1974d 1973 16Sep.15seenoffHatterasTeulings1974d 1974Sep.6seenoffHatterasTeulings1975a1975 12Oct.70seenoffCapeHatterasTeulings1976a1976 18May4seenfeedingatsandflats,OregonTeulings1976dInlet40

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Red-necked Phalarope197611Aug.4seenoffOregonInletTeulings1977a1976 5Sep.26seenoffOregonInletTeulings1977a1976 6Sep.27seenoffHatterasTeulings1977a197621Sep.20seenoffOregonInletTeulings1977a1977 9June3-6seenoffOregonInletLeGrand 1977b 1978 10Oct.13seenoffOregonInletLeGrand1979a1979 10May91seenoffOregonInletLeGrand 1979dSouthCarolinaThestatusoftheked-neckedPhalaropeinSouthCarolinaisprobablythesameasinNorthCarolina.AlthoughregardedbySpruntandChamberlain(1949)asararetransient,mostlyalongthecoast,theyarealmostcertainlycommonmigrantsoffshore.Wayne(1910)andSpruntandChamberlain(1949)agreedthatthespecies was abundantoffshoredespitethepaucityofrecords.Wefoundonly11recordsforthestate,themajority(7)fromthespringmigration.Springrecordsextendfrom17Maythrough3June,andfallrecordsfrom11Septemberthrough25October.188017 Hay colI.inChesterCo. 150 miinlandLoomis18801885 25Sep.seenatFrogmore, lleaufort Co.SpruntandCham-berlain1949 1885? 25Oct.seenat"SeaIslands"SpruntandCham-berlain1949 1903 3JunecapturedatCharlestonWayne 1905 191328 Nay seenoffHt.PleasantSpruntandCham-berlain1949 1933 30 Nay femaleseenatCape Romain Sprunt1933 1934 29 Mayad.femalecolI.at tlull's IslandChamberlain1934 195011Sep.seeninlandinColumbiaarea tlurton 1970 1956 2May2seeninlandatAiken tlurton 1970 1961 22 1IayfemaleseeninlandatLake Hurray Smith1961 1972 '1.7Hay seenatHuntington l>eachJ.'eulings 1972cGeorgia Red-necked PhalaropesarethoughttoberareinshoreandoffshoremigrantsinGeorgiaandaccidentalinland(Dentonetal.1977).Weknowofonly41

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Red-neckedPhalaropeabout18recordsforthestate.Recordsfromthespringmigrationextendfrom5MaythroughearlyJune;recordsfromthefallmigrationfrom24Julythrough3October.Thereisonewinterrecord.1933 24MayfemalecolI.atmouthofSavannah River Burleigh1958 1937 1950 1950 3Oct.9May9Sep.1malecolI.inGradyCountymalecolI.inlandnearAugustaseenatSavannahBurleigh1958DentonandChamberlain1950b Tomkins 1958 195125Sep.2seenonpond 3 mi E SavannahChamberlain1952a19585MaycolI.atHutchinson'sIsland,SavannahTomkins 1958 1958 16 Aug. 1958 13Aug.13Sep.1959 19Sep.seenatSavannah4-11seenatHutchinson'sIsland2seenatHutchinson'sIslandDentonetal.1977Chamberlain1959a,Tomkins 1958Chamberlain1960a 195923,24Sep.4,11964 10Sep.seenatHutchinson'sIslandseenatTybeeIslandChamberlain1960aParnell1965a 1968 12Sep.197227May197623Hay2+2seenatJekyllIslandseenatSt.SimonsIslandseen(1)inlandnearGainesville 1968Teulings1972cTeulings1976c 1976 23 Hay 2(?)seen(1)inlandatPendergrassDentonetal.1977 1976earlyJune2 197724-26July2seen(1)nearPendergrassseeninlandatEufaulaNWRLeGrand 1976 LeGrand 1977b 197911Feb.3seenoffJekyllIslandLeGrand 1979bFlorida-AtlanticCoastTheRed-neckedPhalaropeisararemigrantalongthecoastandisoccasionallyfoundinland(Kale1979msa).TherearefewearlyreportsofthisspeciesfromtheAtlanticcoastofFlorida;Howell(1932) three1976recordsmaybeofthesamebirds.42

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Ked-neckedPhalaropelistedonerecord,andanotherfiveweresummarizedbySprunt(1954).Sincethentherehavebeenatleastanother22recordsfromthecoast,aswellassevenfrominlandlocalities. We listbelowonlycoastalrecordssinceSprunt's(1954)publication,butincludeallcoastalobservationsinTable7. NumbersofKed-neckedPhalaropesseeninrecentyearsclearlyindicatethatthespeciesismuch moreabundantoffshorethanpreviousrecordssuggested,andismostabundantasatransientinApril-May,andinSeptember-October (Table 7).NumbersseeninDecemberandJanuaryalsosuggestthatsomemaywinterinoffshorewaters. 195/l /l-1O !'lay seenatCanaveralStevenson1958c1962 24Apr.seenatFortPiercePaulsonandStevenson1962 19626,8Mayatleast32seen(somecolI.)insmallflocks E ofPalmBeachPaulsonandStevenson1962seenatGuano LakeStevenson1966aseenatPlayalinda Heach Cunningham1964aseenatSawpitSanctuary,DuvalCountyKobertsonandOgden 196/l counted40ydsoffshorebetweenStevenson1962bJunoBeachandJupiterOgden 1974Edscorn1975Kobertson1970Stevenson1972aOgden 1972Stevenson1973Stevenson1973Woolfenden1973Kale1973Edscorn1974Edscorn1974seenoff Hoynton InletseeninIndian Ki vernearSebastionseen18 mi E CapeCanaveralseen24 mioff Nayport seendailyatPortCanaveralseen20 mioffCocoafounddeadnearStuartoffCocoaseenatPortCanaveralseeninIndianKiveratRockledgeseenatLittleTalbotIsland1962 16Dec.16 1963 24Sep.2 196510-11Sep.196721Oct.1969 9Oct. 22197221 Mar. 12 1972 28May6 1972 26 Augseen1972 28Oct.4 1972 16Dec.1973 4 Hay 65+ 1973 9Sep. /l 197313-29Oct.1-2197429June 2 1974 25Sep.43

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]{ed -neckedPhalarope1975 7Sep.19seenoffHayport1977 23Jan.36seen10-15mioffCanaveral197829Apr.upto30seenoffPonceInlet1978 30Dec.seenatPortCanaveralEdscorn1976Stevenson1977Kale1978aStevenson1979aFloridaKeysWeknowofonlyfiverecordsfromtheKeys;allbutoneareofeitherfallmigrantsorwinteringbirds.1954 15 Aug. 1seenatKeyWestStimson1955 195414-15 Hov. seenatLittleDuck KeyStevenson1955a1954 23 Nov. 1seenatKeyVacaStevenson1955a1967 15 Oct. founddeadatNorrisCut,KobertsonandVirginiaKeyOgden 1968 1972 20Maymaleseenoff Duck Key,FloridaKale1972StraitsFlorida-GulfCoast ]{ed-necked PhalaropesoccurovertheopenwatersoftheGulfofMexicobutareseldomrecordedfromtheFloridaGulfcoast(Kale1979msb).liefoundonly16recordsforthisarea,predominantlyofspringmigrants(14Harch-25May).MorethanhalfofallKed-neckedPhalaropesreportedalongtheFloridaGulfcoasthavebeenseenbetween13and25 May.Twosummerrecordscouldhavebeeneitherofearlyfallmigrantsorofbirdsthathadlingeredinthesouthduringthebreedingseason. Few Ked-neckedPhalaropeshavebeenrecordedalongthiscoastinfallandwinter.191814 11ar. seen175mi WofTampaHowell1932 1929 14MaycolI.nearPlantCityHowell1932 195429Apr.seen,colI.atSnakeBight(CapeStevenson1954aSablearea)195625 l'lay colI.atLakeJacksonStevenson1956c1959 22 Nov. 2seenatCapeSanBIasNewman1960a196314May4seenca.1 mioffAlligatorPointStevenson1963b 1963 6JulycolI.nearTampaStevenson1963c196527JuneseenatSanibelIslandOgdenandStev-enson1965 44

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Ked -neckedPhalarope196919-20 Hay seenonupperSt.Marks Kiver, PantelidisandWakullaCountyStevenson1969 197012,16May1,2seenonfreshwaterpond,TampaEdscorn1971 197211-15 Hay seenat HcKay BayKale1972 197213 Hay seenatNavarreBeachImhof 1972 197312Oct.seenatTampaEdscorn1974 1975 27 Nov.foundatNavarrePurrington1976 197621Jan.seenatSanibelIslandStevenson1976 1977 25 Hay ad.femaleseenatToy townDumpKale1977 AlabamaKed-neckedPhalaropesoccuronlyoccasionallyinAlabama,predominantlyduringthefallmigration(Imhof1976b).Weknowofonly12recordsforthestate;eightarefrominlandlocalities,therestfromthecoast.Springrecordsarefrom29Aprilto17May.Fallrecordsoccurfrom6Augustto13November,andthereisonewinterrecord.Imhof(1976b)listednineoftherecords;welistbelowthosefromthecoastandthosepublishedsubsequenttoImhof(1976b).197727-29Aug.1(7)seeninLimestoneCo. 196427Aug.seen1973 10Sep.6 197313-14Oct.seen1975 6 Aug. 197729Apr.atDauphinIslandseen9 mi S ofDauphinIslandatBlakelyIsland,MobileseenatWheeler NWK seenatBlakelyIsland,MobileImhof 1976bImhof1976b Imhof 1976bPurrington1976 Imhof 1977Purrington1978MississippiKed-neckedPhalaropesarerarelyseeninMississippi.Thereareonlythreerecords,alloffallmigrants.Thefirsttwophalaropeswererecordedinlandatsewagelagoonsin1976.Onewasseen11SeptemberatKosciusko,AttalaCounty(Sanders1976)andanotherwasseen21SeptemberatHattiesburg(Gates1976). the thirdsightingwas madealongthecoastatPascagoulaon10September1977(Purrington1978).LouisianaKed-neckedPhalaropesarerarelyseeninLouisiana.Weknowofonlysixrecordsforthestate,fiveofthemlistedbyLowery(1974).Ason the Texascoast,thisspeciesisevidentlymorefrequentduringspringmigrationthaninthefall.45

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Red-neckedPhalarope1966 8 Hay ad.femalecolI.10 mi WofJohnson'sLowery 1974Bayou,nearSabinePass1970May6seenatCameron Lowery 1974 1971 30Apr.-4,3seenatCameron Lowery 1974 1May197112Sep.seen10 mi WJohnson'sBayou Lowery 1974 1978 14MayseenatHollyBeachOilField,Cameron Imhof 1978TexasOberholser(1974)reportedthatRed-neckedPhalaropesareregularbutscarceintheinteriorduringfallmigration;theyarerareduringspringmigration.Alongthecoasttheyareraretocasualinfalland(alongthecentralcoast)casualinspring.SpringmigrationoccursmainlyfromlateApriltolateMay,andmostfallmigrationoccursfrommid-Augusttomid-October.Extremedatesofoccurrencerangefrom4Aprilto6June,andfrom12Julyto22 December(Oberholser1974).Weknowofapproximately55recordsofRed-neckedPhalaropesfromTexas.Aboutthree-quartersoftheserecordsarefrominlandlocalities;nearlytwofifthsofthemweremadeatAustin,wherethespeciesisregularlyrecordedduringthefallmigration.SeptemberisthemonthofpeakoccurrenceintheinteriorbutalongthecoastmostRed-neckedPhalaropeshavebeenseenfromthelatterhalfofAprilthroughearlyMay.Welistbelow13recordsobtainedonornearthecoast.193812JulyseenatJonesLake,Aransas NWR Oberholser1963ms1940 4Apr.2seenatAransas NWR Oberholser1963mS194025-29Apr.7seenonflatsWofMullet Bay, OberholserAransasNWR1963ms1952 26Apr.seenatKockportWatsonandGold-man1952,Ober-holser1963ms1959 4Oct.seenatBaytownTunnelWebster1960a,Oberholser1963ms1959 15Sep.seenatLagunaAtascosaWebster1960a196118July 2. seenatAransas NWR Webster1962a,Oberholser1963ms46

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Ked -neckedPhalarope196217Apr.ca.35 19639-10l1ay 2 1968 18 Oct. 19739-10l1ay 2 197531 Oct. 1979 29Apr.seeneatinginsectsonseaweed,HustangIslandbeachseenonOsoCreek,WofCorpusChristiseen10 mi E of tlrownsville seenatCorpusChristiphotogr.atPadreIslandNatl.Seashorephotogr.in tlrazoria CountyWebster1962bWebster1963Webster1969a tllacklock 1978ms ulacklock 1978msWebster1979c breeding Ked-neckedPhalaropesbreedcircumpolarlyinthenorthernHolarctic,largelybetween52and74N.Theyarefoundmostlyinthetundra,butmayalsobreedinthe zone.Winter Ked -neckedPhalaropeswinteratsea,primarilyintropicalandsubtropicaloceans.TheymaybefoundoffthecoastsofSouthAmerica,northwestAfrica,Arabia,andsoutheastAsia.MigrationTheKed-neckedPhalaropeoccursinthesoutheasternUnitedStatesprimarilyasatransientinspringandfall(Tables7,8).Mostoccuroffshore,andtheirstatusoffmanystatesislargelyunknownbecausethesewatershavebeenpoorlyinvestigated.In Uorth CarolinaandFloridawhereoffshoresurveyshavebeenconductedmostfrequently,thesephalaropesarecommonmigrants.Althoughconsiderablylessinformationisavailableforthenorthern Gulf ofMexico,itseemSevidentthatKed-neckedPhalaropesarelessabundanttherethanofftheAtlanticcoastandareapparentlyrelativelymorecommonduringspringmigration.Peakperiodsofoccurrenceinbothtne Gulf andAtlanticareasareApril-MayandSeptember-October(Table7).OnthebasisofringingrecoveriesobtainedintheOldWorld,Hohn(1966)thoughtthatthebirdsofSweden,Finland,the baltic area,andadjacentnorthernKussiamigrateoverlandtowardtheCaspianSeaandprobablyfromtheretothePersian Gulf. HohnalsosuggestedthatmigrantsoffwesternEuropeprobablycornefromtheeastcoastofGreenland,Iceland,theFaeroes,thenorthern british Isles,andNorway. Hilden andVoulanto(1972),aftera morerecentexaminationofFinnishringingrecoveries,agreedwithHohn;theybelievedthatthemigrationfromFinlandtakesplacetothesoutheastovertheEuropeancontinent.AdditionalinformationonmigrationinotherareasisgivenbyMeinertzhagen(1925), Bent (1927),Hohn(1965),Schiemann(1972),and Glutz von blotzheim etal.(1977).47

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Ked-necked Phalarope Table7.ApproximatenumberofKed-necked Phalaropes recordedbymonthforthecoastalsoutheasternUnitedStates(a).State/regionNorthCarolinaSouthCarolinaGeorgiaFlorida-AtlanticCoastSubtotal-ATLANTIC COAST Florida-KeysFlorida-GulfCoastAlabamaMississippiLouisianaTexas-CoastSubtotal-GULFCOASTTOTAL-ALL AKEASJAHFEJ:!MAl{AP1{ MAY JUN JULAUG SEPOCTNOVDEC -17174 7 144 130 81 2 1 3 622 5 16 1 361231104 22 834423236 4123128912614245 176 232------------------------2121 3 1 3 642 3 41114644 1 35228 53118 5 1----------------------3741383 318131118256 185 933------------------------(a)Ifthesourcedidnotmakeitclearwhetheroneormorebirdswereseen,weassumedthatonlyonewasseen.Whenbirdswereseeninmorethanonemonthwecountedthemseparatelyineachmonth.HABITATNestingBent(1927)describedthepreferrednestinghabitatofKed-neckedPhalaropesinthewesternAleutianIslandsasthewetterportionsofflatalluvialplains.Nestswereplacedinsmallhollowsinmoundsortussocksinwetmeadowsneartheedgesofmarshypondsornearthemouthsofsmallstreams.At Scammon liay, Alaska,thesephalaropesnestedonsedge-grassmarshlands,inclearingsinalderandwillowscrub-coveredslopes,andonheath-coveredslopesjustabovetheseslopes(Hohn1968). Preferred nestinghabitatwastheloweredgeofthealder/willowscrubarea.Kistchinski(1975)notedthatRed-neckedPhalaropesineasternSiberianestedmorefrequentlyalongpermanentbodiesofwaterthandidtheclosely-relatedKedPhalarope.InmostArcticareasthesephalaropeshavebeenreportedtonestinmarsheswithsmallponds(HildenandVuolanto1972andauthorscitedtherein).Nestsaremade on mossinwetplacesoramongsedgesneartheedgeofthewater;thesenestsareoftenplacedonsmallhummockssurroundedbywater.HildenandVuolanto(1972)foundnestsitesatthesouthernmostpermanentbreedingareainFinlandtobeinconsiderablydifferenthabitat.Almosthalf(47.3%,n=74)48

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Ked-necked PhalaropeTable8.Datesofoccurrencefor Red-necked PhalaropesinthecoastalsoutheasternUnitedStates(a).StateNumberofoccurrencesSpringDatesofoccurrenceFallNorthCarolina70+ 3May-l0June10Aug.-25Oct.(10July-4Aug.)(5Feb.)SouthCarolina1117May-3June11Sep.-25Oct.Georgia185l1ay-earlyJune24July-3Oct.Florida2821Mar.-28May23Aug.-2Nov.-Atlanticcoast(29June)(20Nov.-23Jan.)Florida-Keys5 20May15Aug.-15Oct.(14Nov.-23Nov. )Florida16 14Mar.-25May12Oct. -Gulf coast(27June-6July)(22Nov.-21Jan.)Alabama1229Apr.-17May6Aug.-14Oct.(13Nov.-6Dec.)Mississippi3 10Sep.-21Sep.Louisiana5 30Apr.-14 May 12Sep.Texas-coast134Apr.-l0May15Sep.-31Oct.(2July-18July)(a)Kecordsinparenthesesmayrepresentsummeringbirdsorearlyfallmigrants,orlatefallmigrantsorwinteringbirds.ofthenestswere20 m(66ft)ormorefromwater,andmost(78.6%,n=70)wereondrysites(sand,gravel,drymeadow)ratherthanonwetones.Allnestsfoundwereinpatchesofloworsparsevegetation,usuallygrasses(Festuca,Deschampsia,Puccinellia;41.9%,n=74),spikerushes(Eleocharis;28.4%),orsedges(Carex;21.6%).Hohn(1969)notedthatthisspeciestendstonestfartherinlandthanthe Red Phalarope.Kistchinski(1975)reportedadensityineasternSiberiaof0.3-0.5pairs/ha(0.7-1.2/ac)forbirdsnestingintheIndigirkaDelta,comparedto1-2pairs/ha(2.5-4.9/ac)forthe Ked Phalarope.InanareasouthoftheKolymskayaChannelthatKitchinskiregardedaslessdesirablenestinghabitatforRedPhalaropes,comparablefigureswere0.5pairs/ha(1.2/ac)forthe Ked-necked Phalaropeand0.9pairs/ha(2.2/ac)forthe Red Phalarope.AllnestsobservedbyHildenandVoulanto(1972)werefoundinornearcoloniesofArcticTerns.Theseauthorssuggestedthatthisassociationwaschosenbythephalaropes,pointingoutthatthephalaropesreacttoternalarmcallsasdootherspeciesshowingstrongassociationwithlarids,andthatthehighestnestingdensitiesofthesephalaropesinFinnishLaplandalwaysoccurwheretherearenestingArcticTerns.49

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Red-neckedPhalaropeFeedingBianki(1967)remarkedthatRed-neckedPhalaropesusuallyfeedinopenwaterwherebitsofdetrituscollect.Theyarefoundlesscommonlyneartheshorelineathightide.Bent(1927)commentedthatboththisspeciesandthe Red Phalarope,whenatsea,preferredtofeedintiderips,onornearfloatingseaweed,andoftennearwhalesorschoolsoffish.Scott(1959)notedmigrantRed-neckedPhalaropesfeedingamongrowsofdriftingseaweedoffsouthwesternNovaScotia.DuringfallmigrationalongtheYukoncoast,thesephalaropesfeedalongbeaches,intheleeoficeflows,andontheopenocean;theypreferredtofeedinshelteredwatersonwindydays(VermeerandAnweiler1975).Migrantsinlandarefrequentlyseenonsewageponds,rainpools,impoundments,andlakes.Cox(1973)suggestedthatRed-neckedPhalaropesawayfrompelagichabitatsprefershallowpools,incontrasttoRedPhalaropes,whichapparentlypreferdeeperwater.NonbreedingandOffshoreDuringmigrationinthenorthernChesapeakeBight,justnorthofNorthCarolina,Red-neckedPhalaropesoccursomewhatcloserinshorethando RedPhalaropes;theformerusuallyoccur20km(12mi)ormoreoffshore,thelatter70km(44mi)ormore(Rowlett1980).MigrantsarrivinginGreenlandandIcelandinthespringremaininlargeflocksincoastalbaysanddeltasuntilinlandnestingpondsarefreeofice(authorscitedbyHildenandVoulanto1972).FOODANDFEEDINGBEHAVIORRed-neckedPhalaropesfeedprimarilybysurface-seizing(AinleyandSanger1979),i.e.,sittingonthewaterandpickingtheirpreyfromthesurfacewiththeirbills.Phalaropesmayalsoturnrapidlyincircles,abehaviorreferredtoas"spinning".Thismaystirupedibleparticlesfromthebottominshallowwater,butitalsomayservetoactivatechilledinvertebratesindeeperwater(Hohn1971).Hohnsuggestedthatthisbehavioroccurswherethereisadenseconcentrationoffooditems,becausethespinningmotionallowsthebirdstofeedrapidly.Spinningmayalsoplayaroleincourtshipdisplay(Hohn1971,Everett1976).Phalaropesoccasionally"tip-up"likedabblingducks,immersetheirheadsandnecks,andseizepreyunderwater(Hohn1971).Divinghasbeenrecordedonceforthisspecies(Selous1915inHohn1971),andsomefeedbyusingsideto-sidesweepsofthebillinshallowwater(Hohn1968).Benning(1971)notedaninstanceinwhichthisspecieshawkedinsectsuptoafootorsoabovethewater.Red-neckedPhalaropesaregregariousandoftenformfeedingflocks,particularlyattheconclusionofspringmigrationwhenweatherpreventsentrytothebreedinggrounds.Thisgregariousnessextendsintothebreedingseason.DuringthisperiodatScammonBay,Alaska,theyoftencongregateintoflocksofabout20birds(Hohn1968).50

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Ked-neckedPhalarope onlycomprehensivestudyofthefoodhabitsoftheKed-neckedPhalaropein Horth AmericawasconductedbyWetmore (1925), whoexamined155stomachscollectedfromMaytoOctober.Hefoundthattheseshorebirdswerealmostentirelycarnivorous,withanimalmatterforming 97.2% byvolume. Small insectswerethemostimportantitemsofdiet;flies(Diptera)constituted32.8%ofthebulk,buttruebugs -31.8%)andbeetles(Coleoptera-16.5%)werealsoimportant.Oftheflies,earlydevelopmentalandadultstagesofmosquitoes(Culicidae)andgnats(Chironomidae)wereespeciallyimportant;thelarvaeofmosquitoesmadeup6.3%ofthesample.MostofthestomachsinwhichthemosquitolarvaewerefoundevidentlywerecollectedonthebreedinggroundsinAlaska.Amongtheotherinsectgroupsformingmostofthedietafewtaxapredominated.i,ater-boatmen(Corixidae)werethemostsignificantfoodamongtheHemipteraeaten,anddiving-beetles(Dytiscidae)andwater-scavengerbeetles(Hydrophilidae)weremostimportantamongthecoleopterids.Crustaceanswereeatenthroughouttheyearbutmadeuponlyasmallproportion(9.3%)ofthediet;amphipodswerefrequentlytaken,andbrineshrimps(Artemia)wereeatenextensivelyatGreatSaltLake,Utah,duringmigration.Wetmore(1925)suggestedthatcrustaceansprobablyformaconsiderablyhigherproportionofthedietwhenthesebirdsareatsea.OtherkindsoffoodthatWetmore(1925)recordedasoccasionallyorrarelyeatenbyKed-neckedPhalaropesincludedmolluscs(snailsorothergastropods),otherinsects(largelythosefoundinaquatichabitats),marinewormS,spiders,mites,andanunidentifiedfish.Plantscomprisedonlyasmallportionofthefood(2.8%)butwereregularlyfoundinthestomachs;thismaterialconsistedmostlyoftheseedsofplantscharacteristicofwetareas(e.g.,Ruppia,Potamogeton,Scirpus).baker(1977)examinedthecontentsof24stomachscollectedatChurchill,Manitoba,butdidnotreportspecificitemsofdietindetail,notingonlythatthesephalaropespreyedheavilyuponadultchironomids.StomachsofbirdscollectedonKandalakshabay, U.S.S.K., containedonlyterrestrialinsects(bianki1967)thatevidentlyhadbeengleanedfromthesurfaceofthewater.Dement'evandGladkov(1951),inanearliersummaryoffoodhabitsofKed-neckedPhalaropesintheU.S.S.K.,foundthatterrestrialinsectswereeatenonlycasually.Onthewhole,preytakenintheU.S.S.K.wassimilartothattakeninNorthAmerica,i.e.,aquaticinsectsandtheirlarvae.birdsalongtheCaspianSeainfallmainlyfedonamphipods,whilethosefromtheHurmanCoasthadeatenmolluscs,insects,vegetablematter,andpebbles(Dement'evandGladkov1951).PhalaropesoncoastalislandsintheGulfofbothniafedprimarilyonlarvalandadultChironomidaeinsalinesituations,butatemostlyTrichopteralarvae,waterfleas,tadpoles,waterspiders,andcollembolansinfreshwaterponds(HildenandVuolanto1972).51

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Red-neckedPhalaropeSUSCEPTIBILITYTOOILPOLLUTIONWefoundnoreportsofoiledRed-neckedPhalaropes.Althoughtheyfeedonthesurfaceinflocks,theiroffshoredistribution,theirshortstayinmostsoutheasternwaters,andtheirscarcityinpartsofthesoutheastmakethemofrelativelylowconcernwithregardtothedevelopmentofpetroleumresources.Thisisprobablynottrueinotherareas.TheirfeedinghabitsandflockingbehavioronfallmigrationalongtheYukoncoastledVermeerandAnweiler(1975)toconsiderthemperhapsthemostvulnerabletooilpollutionofallshorebirdsoccurringthere.KingandSanger(1979)agreedbygivingtheRed-neckedPhalaropeoneofthehighestratingsforvulnerabilityforanyshorebirdoccurringinthenortheasternPacific.Theyindicatedthatthisspeciesshouldbeofspecialconcernvisavisthepotentialeffectsofoilpollution.BIBLIOGRAPHY1982Orr,C.D.,R.M.P.Ward,N.A.WilliamsandR.G.B. Brown.1982.MigrationpatternsofRedandNorthernPhalaropesinsouthwestDavisStraitandinthenorthernLabradorSea.WilsonBull.94:303-312.Ruttledge,R.F.1982.Red-neckedPhalarope--ahithertounrecordedcaseofbreedinginCo. Mayo.IrishBirds2:196-197.1981 Hohn, E. O.1981.NorthernPhalaropeflocksatMiquelonLake,Alberta.BlueJay39:41-43.Pigeon,P.1981.Observationd'unPhalaropeabecetroitPhalaropuslobatusetd'unPhalaropeabeclargePhalaropusfulicariusaAulnoissous-Laon(Aisne).Alauda49:232.1980Becker,P.1980.ZumVorkommendesOdinswassertreters(Phalaropuslobatus)andenKusteSudwestafrikasundzurfeldornithologischenUnterscheidungderWassertreter.Ornithol.Mitt.32:40-44.Guex, C.1980.Observationhivernaled'unPhalaropeabecetroitPhalaropuslobatussurIeLeman.[WinterobservationsoftheRed-neckedPhalaropePhalaropuslobatusonLakeGeneva.]NosOiseaux35:372.[InFrench.]Hohn, E. andD.J.Mussell.1980.NorthernPhalaropebreedinginAlberta.Can.Field-Nat.94:189-190.King,F.1980.Red-neckedPhalaropesbreedingatAkeraghLoughinCountyKerry.IrishBirds1:540-541.52

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Red-neckedPhalaropeOrsini,P.1980.PremiereobservationhivernaleduPhalaropeabeeetroitPhalaropuslobatusdansIeMoyen-Atlas(Maroc).[FirstwinterobservationofRed-neckedPhalaropePhalaropuslobatusintheMiddleAtlas(Morocco).]Alauda48:258-259.1979Holmes,G.1979.FirstrecordofRed-neckedPhalaropeinNewSouthWales.Austr.Birds13:75-76.Roudybush,T.E.,C.R.Grau,M.R.Petersen,D.G.Ainley,K.V.Hirsch,A.p.GilmanandS.M.Patten.1979.Yolkformationinsomecharadriiformbirds.Condor81:293-298.Toups,J.A.,M. B. HodgesandP.V.Donaldson.1979.NorthernPhalaropeatPascagoula--firstphotographicrecordforMississippi.MississippiKite9:9-10.1978Dunnett,G.M.andS.W.Kotogama.1978.Red-neckedPhalaropeinSriLanka.J.BombayNat.Hist.Soc.75:215-216.Ruttledge,R.F.1978.Red-neckedPhalaropesbreedingsouthoflatitude53037'inIreland.IrishBirds1:229-231.Taylor,W.B.1978.Seasonalstatus,populationsize,andhabitatutilizationofRedandNorthernPhalaropesintheSouthernCaliforniaBight.(Abstractonly.)Pac.SeabirdGroupBull.5:74.1977Ash,J.S.andO.M.Ashford.1977.GreatBlack-headedGullsLarusichthyaetusandRed-neckedPhalaropesPhalaropuslobatusinlandinEthiopia.J.EastAfr.Nat.Hist.Soc.31:13.Baker,M.C.1977.ShorebirdfoodhabitsintheeasternCanadianArctic.Condor79:56-62.Brichetti,P.Italia.1977.IIfalaropobeccosottile,Phalaropuslobatus(L.)inUccellid'Italia2:129-131.Gianella,C.1977.Observazionediunfalaropobeccosottile-Phalaropuslobatus(L.)-nellaLagunadiPonentediOrbetello.Riv.Ital.Ornitol.47:293-294.Muller,G.andH.-P.Keller.1977.BeobachtungenvonWeissflugelseeschwalben(Chlidoniasleucopterus),steinwalzer(Arenariainterpres)undOdinswassertreter(Phalaropuslobatus)inderOberlausitz.Beitr.Vogelkd.23:60-61.53

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Red-neckedPhalaropeRuthke,P.1977.FuttertragenundDunenkleiddesOdins-wassertreters(Phalaropuslobatus).[FoodcarryingandjuvenileplumageoftheRed-neckedPhalaropePhalaropuslobatus.]Vogelwelt98:229-231.[InGermanwithEnglishsummary.]Schiemann,H.1977a.UberdasVorkommenderWassertreter(Phalaropodidae)indenbrandenburgischensachsischenundthuringischenBezirken,souieinBerlin.Beitr.Vogelkd.23:49-56.1977b.BeringungsergebnisseNordeuropaischerOdinshunchen.[BandingresultsofNorthEuropeanRed-neckedPhalaropes.]Sterna16:73-80.[InGermanwithNorwegiansummary.]Sinclair,J.C.1977.Thephalaropes.Bokmakierie29:90-91.1976Crawford,P.andD.Crawford.1976.SecondrecordofNorthernPhalaropeinNashvillearea.Migrant47:94.Everett,J.J.1976.MutualspinningbyRed-neckedPhalaropes.Brit.Birds69:219-220.Gates,L.J.1976.NorthernPhalaropeinHattiesburg,Mississippi.MississippiKite6:37.McNamara,J.A.1976.TwofurtherrecordsoftheRed-neckedPhalarope.S.Austr.Ornithol.27:142.Sanders,J.A.1976.ANorthernPhalaropeinAttalaCounty,Mississippi.MississippiKite6:36.Vanhercke,L.1976.Degrauwefranjepoot(Phalaropuslobatus).Wielewaal42:342-343.1975Dontschev,S.1975.Neue AngabenuberdasAuffindenvonPhalaropuslobatus(L.),GlareolanordmanniFisch.-Waldh.undArenariainterpres(L.)amderbulgarischenSchwarzmeerkuste.Larus26-28:183-187.Kistchinski,A. A.1975.BreedingbiologyandbehaviouroftheGreyPhalaropePhalaropusfulicariusinEastSiberia.Ibis117:285-301.Menon,S.S.1975.OccurrenceoftheRed-neckedPhalarope(Phalaropuslobatus)oninlandwatersinBangalore.J.BombayNat.Hist.Soc.72:846-847.Nakamura,K.1975.[OnaRed-neckedPhalaropeencountered[ina]typhoonduringmigration.]Tori23:95-96.[InJapanesewithEnglishsummary.]54

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Red-neckedPhalaropeSchmidt,G.1975.BeobachtungenuberdenWegzugdesOdinswassertretersausnordnorwegischenBrutgebietenanderBarents-See.Beitr.Vogelkd.21: 233 244.[InGerman.]Vermeer,K.andG.G.Anweiler.1975.OilthreattoaquaticbirdsalongtheYukoncoast.WilsonBull.87:467-480.1974Eller,G.D.1974.NorthernPhalaropeinuppereastTennessee.Migrant45:96.Kopke,G.1974.Odinswassertreter(Phalaropuslobatus)imJuniaufBaltrum.Vogelkd.Ber.Nieders.6:18.Seeger,J.-J.1974.OdinswassertreteramGu1perSee. Falke 21:284.Steinbach,R.1974.VorkommerderwassertreterPhalaropuslobatus(L.)undPhalaropusfu1icarius(L.)imGebietderHaselbacharundEschefe1derTeichesowieamSpeicherbechtenWindischleuba.Abh.Ber.Naturkundemus.'Mauritanium'8:333-338.1973Cox,J.B.1973.AfurtheroccurrenceoftheRed-neckedPhalaropeinSouthAustralia.S.Austr.Ornithol.26:116-117.Fevold,H.R.,E. W. PfeifferandJ.S.Rice.1973.Steroidbiosynthesisbyphalarope(SteganopustricolorandLobipeslobatus)adrenaltissue.Gen.Comp.Endocrinol.21:353-357.Smith,F. To H.lobatus).1973a.Red-neckedPhalaropeatAltonasa1tworks(PhalaropusBirdObs.473:12.1973b.FurthertotheAltonaRed-neckedPhalarope.BirdObs.477:4-5.Stelzer,M.1973.UberdasVorkommenderWassertreter(Pha1aropodidae)inderSchweijundihrenRandgebieten.Ornithol.Beob.70:157-170.[InGermanwithFrenchsummary.]vanAvermaet,G.1973.Franjepoten.Wielewaa139:216-220.1972Hilden,O.andS.Vuo1anto.1972.Pha1aropus lobatus inFinland.BreedingbiologyoftheRed-neckedPhalaropeOrnis Fenn. 49:57-85.Jovanovic,V.1972.Novipodaci0seobiliskonogetankokljune,Phalaropus10batus,uVojvodini.Larus24:164.55

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Red-neckedPhalaropeRaner,L.1972.Forekommerpolyandrihossmalnabodsimsnappaochsvartsnappa?[PolyandryintheNorthernPhalaropeandtheSpottedRedshank.]FaunaFlora67:135-138.[InSwedish.]Schiemann,H.1972.UberWinterquartierenordeuropaischerOdinshuhnchen.[OnwinteringareasofnorthernRed-neckedPhalaropes(Phalaropuslobatus).]Vogelwarte26:329-336.[InGermanwithEnglishsummary.]Vaisanen,R.A.,O.Hilden,M.SoikkeliandS.Vuolanto.1972.Eggdimensionvariationinfivewaderspecies:theroleofheredity.OrnisFenn.49:25-44.1971Benning,W.E.1971.PhalaropeshawkinginsectsoverthewateratMontezuma.Kingbird21:11.Edscorn,J.B.1971.NorthernPhalarope(Lobipeslobatus)occurrencesincentralFlorida.Fla.Nat.44:94.Everett,M.J.Ireland.1971.BreedingstatusofRed-neckedPhalaropesinBritainandBrit.Birds64:293-302.Hohn,E.O.1971.ObservationsonthebreedingbehaviourofGreyandRedneckedphalaropes.Ibis113:335-348.McGregor,A.1971.Rarebirds:Red-neckedPhalaropes.BirdLife7:2-5.Pringle,J.S.andW.S.Pringle.1971.Red-neckedPhalaropePhalaropuslobatusatStrandfontein.Ostrich42:228-229.Scholz,J.Artern.1971.Odinswassertreter(Phalaropuslobatus)amSchlammteichbeiBeitr.Vogelkd.17:174-175.Verheyen,J.A.1971.Red-neckedPhalaropePhalaropuslobatusintheIslandofFlores,Indonesia.Ardea59:64.1970Alsop,F.G.,III.NationalPark.1970.NorthernPhalaropeintheGreatSmokyMountainsMigrant41:39-40.Coffey,J.w.1970.NorthernPhalaropeinWashingtonCounty.Migrant41:66-67.Haight,T.1970.NorthernPhalaropeinDutchessCounty.Kingbird20:179-180.Hohn,E.o.1970.GonadalhormoneconcentrationsinNorthernPhalaropesinrelationtonuptialplumage.Can.J.Zool.48:400-401.56

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Red-neckedPhalaropeLowery,G.H.,Jr.1970.TheLouisianastatelist.La.Ornithol.Soc.News56:1-7.1969Bates,A.K.andM.L.Bierly.1969.White-rumpedSandpipers,NorthernPhalaropes,RuddyTurnstones,Gull-billedTernsandStiltSandpipersatMarionFishHatchery.Ala.Birdlife17:52-54.Eckert,J.1969.Red-neckedPhalaropeinsouthAustralia.Emu69:184-186.Eriksson,K.1969.WeibchenschwarmedesOdinswassertreters(Phalaropuslobatus)unddieDatierungihrerGelegeinFinnischLappland.[OntheflockingoffemaleRed-neckedPhalaropes(Phalaropuslobatus)duringthebreedingseasonandonthetimeofegg-layinginFinnishLappland.]Ornithol.Mitt.21:157-160.[InGermanwithEnglishsummary.]Hohn,E.O.1969.The Sci.Am.220:104-111.Schoennagel,E.1969.GehauftesAuftretendesOdinswassertreters(Phalaropuslobatus)aufderlnselBorkum(Nordsee).Ornithol.Mitt.21:18.Smith,F.T.H.1969.TheRed-neckedPhalaropenearMelbourne.Austr. Bird Watcher3:166-167.1968Gerasimov,N.N.1968.[BreedingoftheRed-neckedPhalaropeinKamchatka.]Ornitologiya9:344-345.[InRussian.]Glayre,D.1968.UnPhalaropeabecetroitsejournedixjoursaChavornay.NosOiseaux29:275-276.Hohn,E.O.1968.SomeobservationsonthebreedingofNorthernPhalaropesatScammon Bay, Alaska.Auk85:316-317.Masters,G.1968.Black-neckedStiltsandNorthernPhalaropesatJekyllIsland,Ga.Oriole33:50-51.Plucinski,A.1968.Odinswassertreter(Phalaropuslobatus)beiGoslar.Ornithol.Mitt.20:239.1967Kumerloeve,H.andP.A.D.Hollum.1967.ZumdurchzugvonPhalaropuslobatus(L.)inKleinasien.Vogelwarte24:64-65.57

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Red-neckedPhalarope1966 Hohn,E.O.1966.Ringing(banding)andrecoveriesofphalaropes--asummaryofpresentlyavailableinformation.Bird-Banding37:197-200.Warfield,R.w.1966.VagrantNorthernPhalaropeatLilypons.Md.Birdlife22:14-15.1965Dittberner,H.andW.Dittberner.1965. Sturmmowe,Laruscanus,brutendundOdinswassertreter,PhalaropuslobatusrastendamGrossenSchwerin(Muritz).Beitr.Vogelkd.11:103-105.Hohn,E.O.1965.DieWassertreter.NeueBrehn-Bucheri359.ZiemsenVerlag,Wittenberg.60pp.Turk,M.1965.NorthernPhalaropeatDauphinIsland.Ala.Birdlife13:42.1964Clark,W.S.1964.Anunusualvisitor(NorthernPhalarope).Ala.Birdlife12:12.Rittinghaus,H.1964.Phalaropuslobatus(Phalaropodidae)Nahrungserwerb.Pp.309-311inG.Wolf(Ed.)EncyclopediaCinematographica,Inst.furdenWissenschaftlichenFilm,Gottingen.[InGerman.] 1963Smith,F.T. H.1963.AnAustraliansightrecordoftheRed-neckedPhalarope(Phalaropuslobatus).Austr.BirdWatcher2:1-4.1962Longstreet,R.J.1962.NorthernPhalarope(Lobipeslobatus)on Lake Monroe.Fla.Nat.35:92.Pocock,T.N.1962.Red-neckedPhalaropePhalaropuslobatusandotherbirdsatIscorDams,VanderbijlPark.Ostrich33:41-44.1961Smith,E.D.1961.NorthernPhalaropeon LakeMurray,SouthCarolina.Chat 25:68.1959Hall,G.A.1959.AlaterecordforNorthernPhalaropeinWestVirginia.WilsonBull.71:194.58

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Red-neckedPhalaropeOgden,J.1959.NorthernPhalaropesatNashville.Migrant30:55.1958Bellier,L.and R. Leveque.1958.PhalaropeabecetroitenCamargue.Alauda26:230.[InFrench.]1957Haftorn,S.1957.Omforeldrenesomsorgforeggogungerhossvommesnipe,Phalaropuslobatus(L.)andtemmincksnipe,Calidristemminckii(Leisl.).[On parentaY-Care inPhalaropuslobatus(L.)andCalidristemminckii(Leisl.).]K.Nor.Vidensk.Selsk.Mus.Arbok1957:12-144.Harrisson,T.1957.Red-neckedPhalarope,Lobipeslobatus(Linnaeus)insoutheastAsia.J.BombayNat.Hist.Soc.54:947.MorzerBruijns,W.F.J.andM.F.MorzerBruijns.1957.Waarnemingenvande GrauweFranjepoot,Phalaropuslobatus(L.),indeIndischmOceaan.[ObservationsoftheRed-neckedPhalarope,Phalaropuslobatus(L.),intheIndianOcean.]Ardea45:72-84.[InDutchwithEnglishsummary.]1956Christensen,N.H.1956.Odinshanens(PhalaropuslobatusL.)oglnorshanens(PhalaropusfulicariusL.)forekomsti Danmark. [TheoccurrenceinDenmarkoftheRed-neckedPhalarope(PhalaropuslobatusL.)andtheGreyPhalarope(PhalaropusfulicariusL.]Dan.Ornithol.Foren.Tidsskr.50:191-206.[InDanishwithEnglishsummary.]Mester,H.1956.Odinshuhnchen(Phalaropuslobatus)beiFrondenbergjRuhr.Ornithol.Mitt.8:53.1955Geroudet,P.1955.LepassagedesphalaropesenSuisseromande.NosOiseaux23:42-47.[InFrench.]Stimson,L.A.1955.NorthernPhalarope.Fla.Nat.28:29.1954Maluquer,S.M.1954.PhalaropuslobatusenEspana.Ardeola1:121-1221953Alexander,H.G.1953.RedneckedPhalaropenearDelhi.J.BombayNat.Hist.Soc.51:507.59

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Red-neckedPhalaropeBerndt, R. and R. Reinecke.1953.ErstnachweisdesOdinswassertreters,Phalaropuslobatus(L.)furdasBraunschweigerHugelland.Ornithol. Meier,H.1953.SchmalschnabligerWassertreterimUrnerReussdelta.Ornithol.Beob.50:32.Mouchet,E.1953.LePhalaropeabecetroitauxGrangettes.NosOiseaux22:72.1952Eberhardt,R.L.withfishes.1952.NorthernPhalaropesandXanthusMurreletassociatedCondor54:314.Simon, H.1952.Observationd'unPhalaropehyperboredansIe OiseauRev.Fr.Ornithol.22:322.1951Hanstrom, R. 1951.Densmalnabbadesimsnappen,enoriginellsvensksmavadare.FaunaFlora1/2:57-66.[Wood,J.D.].1951.StudiesofsomespeciesrarelyphotographedXXXI.The Grey[and]Red-neckedPhalarope.Brit.Birds44:235-236.1950Nevin,W.S.1950.Red-neckedPhalaropeontheKent-Sussesborder.Brit.Birds43:191-192.1949Longstreet, R. J.1949.FlockofNorthernPhalaropesatDaytonaBeach,Florida.Auk66:204.vanIjzendoorn,A.L.J.1949.Apreeningphalarope(Lobipeslobatus).Auk66:89-90.1948Walkinshaw,L.H.1948.NestingsofsomeshorebirdsinwesternAlaska.Condor50:220-223.1945Vaughan,R.1945.Ararevisitor.CountryLife97:1001.60

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Red-neckedPhalarope1943Cole,L.J.1943.BehaviorofNorthernPhalaropewithyoung.Condor45:39.1938Michael,C.W.1938.BehaviorofNorthernPhalaropes.Condor40:85.1937 Low,G.c.1937.GreyandRed-neckedPhalaropesintheArabianSea.Ibis(14thSer.)1:866.Meinertzhagen,R.1937.Grey andRed-neckedPhalaropesinArabianSea.Ibis(14thSer.)1:667.Sits,E.1937.BeobachtungenuberPhalaropuslobatusanderMatsuluBucht.Ann.Soc.Nat.Dniv.Tartu43:16-24.Wust,w.1937.SchmalschnabligerWassertreter,Phalaropuslobatus(L.)beiIsmaningerlegt.Anz.Ornithol.Ges.Bayern2:449450.1935Tinbergen,N.1935.FieldobservationsofEastGreenlandbirds.I.ThebehaviouroftheRed-neckedPhalarope(PhalaropuslobatusL.)inspring.Ardea24:1-42.1934 Bonami,L.[L.])1934.CatturadiunFalaropoabeccosottile(PhalaropuslobatusStudiTrentini15:196-197.Chamberlain,E.B.1934.NorthernPhalaropeinSouthCarolina.Auk51:519520.1933Sprunt,A.,Jr.1933.SecondoccurrenceofNorthernPhalaropeinSouthCarolina.Auk50:358-359.Stone,W.1933.NorthernPhalaropesontheNewJerseycoast.Auk50:475-476.Taning,A.V.1933.Thewinterquartersofthephalaropes.Ibis(13thSer.)3:132-133.Young, C.G.1933.Thewinterquartersofthephalaropes.Ibis(13thSer.)3:548.61

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Red-neckedPhalarope1932Urner,C.A.1932.PhalaropesinNewJerseyinspring.Auk49:475-476.1931Glegg,W.E.1931.OnthenestingoftheRed-neckedPhalaropeinShetland.Oologists'Rec.9:10-14.1929Poole,E.L.1929.NorthernPhalarope(Lobipeslobatus)inPennsylvania.Auk46:108.Townsend,C.w.1929.BreedingrangeoftheNorthernPhalarope(Lobipeslobatus).Auk46:108-109.1928Schorger,A.w.1928.ThewinteringareaoftheRedandNorthernPhalaropes.Auk45:206.1927Momiyama,T.1927.[OnthespecimensofLobipeslobatusobtainedatPrefect.Saiitama,Hondo.]Tori5:267-271.[InJapanese.]1926Grote,H.1926.ZurVerbreitungvonPhalaropuslobatus(L.)undPhalaropusfulicarius(L.).Ornithol.Monatsber.34:122.Spingarn,E.D.W.1926.NorthernPhalaropeinDutchessCounty,N.Y.Auk43:90-91.1925Meinertzhagen,R.1925.Thedistributionofthephalaropes.Ibis(12thSer.)1:325-344.Rapine,J.1925.LesPhalaropes.Rev.Fr.Ornithol.17:54-56.Wetmore,A.1925.FoodofAmericanphalaropes,avocets,andstilts.U.S.Dep.Agric.Bull.1359.20pp.1923TschusizuSchmidhoffen,V.1923.Phalaropusfulicarius,P.lobatusundAcanthisflavirostrisinOberosterreich.Ornithol.Monatsber.31:62.62

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Red-neckedPhalarope1921Gordon,A.1921.AnoteonthenestingoftheRed-neckedPhalarope.Brit.Birds15:90-91.1915Selous,E.1915.AdiaryofornithologicalobservationsmadeinIcelandduringJuneandJuly,1912.Zoologist19(Ser.4):54-56,169-174,303-307,20:54-68,139-152,267-272.1914Best,M.G.S.andM.D.Haviland.1914.NotesontheRed-neckedPhalaropeintheOuterHebrides.Brit.Birds8:9-12.1910Brimley,H.H.1910.NorthernPhalaropeinBladenCounty,NorthCarolina.Auk27:206.Morris,R.O.1910.CaptureoftheNorthernPhalaropenearSpringfield,Mass.Auk27:79.1909Smith,C.B.1909."Sundhani".Avic.Mag.7:309-311.1908Bahr,P.H.1908.SomeobservationsonthebreedinghabitsoftheRed-neckedPhalarope.Brit.Birds1:202-207.Service,R.1908.Red-neckedPhalaropeinSolwayArea.Ann.Scott.Nat.Hist.1908:253.1907Gladstone,H.S.1907.TheRed-neckedPhalaropeinIreland.Brit.Birds1:174-1771905Wayne,A.T.1905.SouthCarolina.NotesoncertainbirdstakenorseennearCharleston,Auk22:395-400.1903Williams,E.1903.BreedingoftheRed-neckedPhalaropeinIreland.IrishNat.12:41-45.63

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Red-neckedPhalaropeForrest.J.1901.Red-necked Phalarope (Phalaropushyperboreus)in North Wales.Zoologist1901:428.Loomis. L.11. 1880.TheNorthernPhalaropeinChesterCounty.SouthCarolina.8ull. Nuttall Ornitho1.Club5:242.Coues. E. 1878. The NorthernPhalaropein North Carolina.Ornitho1.Club3:40-41.Red-neckedPhalaropeinwinter plumage. PhotographbyClaytonTaylor.64

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RED PHALAROPE(Phalaropusfulicaria)[DA:Thorshane,DU:Rossefranjepoot,EN:GreyPhalarope,FI:Vesipaasky,FR:Phalaropeabeclarge,GE:Thorshuhnchen,Thorswassertreter;IC:Thorshani,IT:Falaropoabeccolargo,JA:Hai-irohire-ashishigi,NW:Polarsvommesnipe,PO:Platkonogplaskodzioby,PR:Falarodo,RU:(Flat-nosedPhalarope),SP:Falaropopicogrueso,Chorlillonorteno;SW:Brednabbadsimsnappa] GENERAL DISTRIBUTIONNorthAmericaThebreedingrangeoftheRedPhalaropeinNorthAmericaoverlapsthatoftheRed-neckedPhalaropebutextendsconsiderablyfarthernorth.ThisspeciesbreedsfromwesternandnorthernAlaskaeastacrossCanadianArcticislandstoBaffinIslandandnorthtonorthernEllesmereIsland.AlongtheCanadianmainlanditbreedsinnbrthernMackenzie,easternKeewatin,northernQuebec(AOU1957,Godfrey1966),andprobablynorthernLabrador(Godfrey1966).NorthAmericanpopulationsmigratesouthalongbothcoastsandinthePacificandAtlanticOceanstowinteroffSouthAmerica(AOU1957).WorldDistributionIntheOldWorld,RedPhalaropesbreedfromGreenlandandIcelandeastthroughtheArcticislandsofSpitsbergenandsouthernNovaya ZemlyatotheNovosibirskiyeIslands.AlongthemainlandtheybreedacrossnorthernSiberiafromtheTaimyrPeninsulaeasttotheChukotskiPeninsulaandthecoastofAnadyrland(AOU1957,Vaurie1965).PrimaryWinteringgroundsareatseaoffbothcoastsofSouthAmericasouthtoPatagoniaandtheFalklands,offwestAfricasouthtosouthernArabia,andprobablyinthePacificofftheBoninIslandsandSevenIslandsofIzu(Vaurie1965)southtoNewZealand(AOU1957).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESNorthCarolinaRedPhalaropesoccasionallyoccuralongthecoastofNorthCarolina(Pearsonetal.1942)andareseeninlandinfrequently.Throughtheearly1960'stherewerefewrecords,butwiththerecentincreaseinoffshoreobservations,ithasbecomeclearthattheRedPhalaropeisaregularmigrantthatissometimesencounteredinlargenumbers(LeeandBooth1979).Mostmigrationoccursbetween30Septemberand29December,andagainbetween8Marchand4MayaccordingtoLeeandBooth(1979).TheysuggestedthatthespeciesmaywinterinNorthCarolinawatersbecauselargenumbersareseeninlateDecemberandearlyApril.LeeandBooth(1979)impliedtherewereatleast43recordsofRedPhalaropesoccurringinNorthCarolina.Wefoundabout40publishedrecords,but65

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RedPhalaropelistbelowonlycoastalrecordsthatinvolvefiveormorebirds.Sevenoreightrecordsareforinlandlocalitieswherethesephalaropesoccur from 30Septemberthrough21March.TherearerecordsforallinterveningmonthsexceptFebruary;allotherrecordsarefromthecoastoroffshore.1896 2or3ca.12founddeadatCape LookoutLighthouseApr.1907 8Apr.6colI.atBodieIsland190717Apr.6call.atBodieIsland19371,3May2,8seenatPeaIslandNWR1956 14Apr.23seen(3call.)atPeaIslandNWR1965 24Apr.72seen30 miESEWrightsvilleBeach 1966 9Apr.149seenSofBeaufortPearsonetal.1942Pearsonetal.1942Pearsonetal.1942Pearsonetal.1942Chamberlain1956bJones1965Parnell1966b 1966 23Apr.1,000+seen6 mioffWrightsvilleBeachParnell1966b 1966 4Sep.1973 6 Dec. 197414Nov. 1975 26Jan.88 200 15028seenoffHatterasInletseen5 mioffCharlestonseenin3flocks,40-50miNECharlestonseeninshore,WrightsvilleBeachParnell1967aTeulings1974bTeulings1975aTeulings1975b 1975Aug.-Oct."afew"seeninCapeHatterasareaTeulings1976a 197729Dec. 1978 4Apr.197817Apr.100's-1,000'sseenoffNorthCarolina100's-1,000'sseenoffNorthCarolina1,100seeninGulfStreamoffOregonInletLeeandBooth1979 LeeandBooth1979Teulings1978a 197814Nov.8seenoffOregonInletLeGrand 1979a 1978 5 Dec.100's-1,000'sseenoffNorthCarolina66LeeandBooth1979

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1978 30Dec.1979 23Apr.100's-1,000'sseenoffNorthCarolina1,000+seenoffOregonInletRedPhalaropeLeeandBooth1979 LeGrand 1979cSouthCarolinaBurton(1970)consideredtheRedPhalaropetherarestofthethreephalaropesinSouthCarolina.HeaddedtworecordstothetwocertainonesreportedbySpruntandChamberlain(1949).Therearefouradditionalrecords,allfromoffshore.ThenumbersseensuggestthatthestatusofthisspeciesinSouthCarolinaissimilartoitsstatusinNorthCarolina,i.e.,commontoabundantinlatefallandearlywinter,aswellasinearlyspring.Thebirdsseenin1898 werenotpositivelyidentifiedasRedPhalaropes,butconsideringthedifferenceinseasonalabundancebetweenthisspeciesandtheRed-neckedPhalaropeofftheAtlanticcoast,thisidentificationisthemostprobable.189817Mar."enormousseenca.50 mioffnortherncoastSpruntandCham-flocks"berlain1949 1900 4Dec.malecaughtatMt.PleasantSpruntandCham-berlain1949 1934 22Apr.seennearCharlestonLightshipSprunt1934 196011JuneseenoffCapeIslandnearMcClellans-Baldwin1960ville196315Sep.seenon pondnearWallaceRiver,Shuler1964Rantowles,CharlestonCo. 196412Mar.21seen26-30miSECharlestonHarborSykes1966 1972 30May2seenoffshore35 miESECharlestonTeulings1972c 1973 6Dec.ca.200seen5 mioffCharlestonTeulings1974b 1974 14 Nov. 125seen40-50miNECharlestonTeulings1975aGeorgiaDentonetal.(1977)consideredtheRedPhalaropeaccidentalinGeorgiaandlistedonlytworecords.Onephalaropewascollected29October(Dentonetal.1977)or5 November1970(Teulings1971a)inlandatThomaston,50 miNEofColumbus.Anotherwasseen28 mioffSt.SimonsIsland8 March1975(Teulings1975b).Weknowofnorecordssincethenbutbelievethatthesephalaropesarealmostcertainlyabundantoffshore;theygounrecordedforlackofobservation.Florida-AtlanticCoastKale(1979msa)reportedthatthisspeciesisrareoffshoreduringmigrationandinwinter.The numberofreportsfortheFloridaAtlanticcoastarefew(ca.37),butthenumbersseenoffshoreduringrecentsurveysmakeitclearthatRedPhalaropesarecommonandattimesabundant.Toofewdataareavailabletodelineateperiodsofpeakabundanceorto67

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RedPhalaropeestimatetotalpopulationsoccurringoffshore.Howell(1932)andSprunt(1954)togetherlistedeightobservationsfortheAtlanticcoastofFlorida;wefoundanother29,almosthalfofthemmadesince1970.WelistbelowonlythosepublishedsubsequenttoSprunt(1954)andthoseknowntoinvolvemorethanonebird.Extremedatesofoccurrencerangefrom23Julyto27April,butapreponderance(ca.85%)oftherecordsarefrom14Augustto27March.1956 2Jan.smallnumbersseen10-20mioffCocoa Beach 195627Mar.smallnumbersseen10-20mioffCocoaBeachStevenson1956bStevenson1956b 1959 7Feb.7seen10mioffMiamiStevenson1959b 1960Mar.ca.500seenoffCanaveralStevenson1960b,1961 1964 31July1967 16Feb.197127Apr.1971 14 Aug. 2022231seen18 mioffCocoaseenatJacksonvilleBeachseen10-30mi EPortCanaveralseen15-28mioffCocoaStevenson1964bStevenson1967bKale1971 Ogden 1971 1971 5Sep.500seenoffPortCanaveralRobertson1972 1972 22Jan.1972 23July197321Jan.1973 2 Aug. 1973 9Sep.1974 28July1975 7Sep.197723Jan.197711Sep.20-100seenoffJacksonville2seenoffCocoa 19seenoffMayport8seenoffCanaveral120+seen24mioffMayport4seen20 mioffCocoa67seenonpeakcountoffMayport400+seen10-15mioffCanaveral20seenoffBoyntonInletStevenson1972a Ogden 1972Woolfenden1973Edscorn1974Edscorn1974 Ogden 1974Edscorn1976Stevenson1977Edscorn1978Florida-GulfCoast Rale (1979msb)consideredthisspeciesextremelyrareontheFloridaGulfcoastandinland.Sprunt(1954)listedthreerecordsfromthecoast,andwefoundonlysevensince,listedbelow.Thetemporalspan68

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RedPhalaropeoftheserecords,allbutoneinvolvingsinglebirds,is29Octoberto8 May;halfofthesesightingsoccurredinNovemberandDecember. Thestatusofthisspeciesoffshoremaybequitedifferent.RedPhararopeswereregularlyreportedoffPensacola(mostlybycommercialfishermen)from1946through1954,inlargenumbers.ReportspublishedinAmericanBirdsandbyWeston(1953)revealthatbirdswereseenonatleast26individualdatesduringthisperiod.MostoftheseweresummarizedbyWeston(1953),whoconcludedthattheRedPhalaropecouldbe"consideredaregular,andsometimescommon,winterresidentofthemiddlenorthernGulfofMexico."Headdedthatallbuta fewobservationsweremade morethan30milesoutintheGulfandthatonlyoneobservationwas madeasnearas5miles.Theseobservationsspannedaperiodfrom13October(Weston 1953)through11April(Newman1954);thelargestnumbersofbirdswereseenfromlateDecemberthroughlateFebruary.WedonotlisttheseoffshorerecordsorthethreerecordsgivenbySprunt(1954)here,butincludetheminTable9.195721Apr.seenoninlandpondatPensacolaNewman1957b1959 7May3seen5mioffManasota ky Stevenson1959c 1962 30 Nov.foundinlandatWCTVTowerStevenson1967a 1965 8MayseennearAlligatorPointCunningham 1965b 196621Nov.foundinlandatWCTVTowerStevenson1967a 1967 6 Dec.seeninlandatLakeTalquinStevenson1968a 1975 27 Nov.foundinjuredatNavarrePurrington1976 Alabama Imhof(1976b)consideredtheRedPhalaroperegularandsometimescommoninwinterofftheAlabamacoast,primarilyonthebasisofobservationsmadeoutofPensacolaBay. Imhof(1976b)listedfiverecordsoffPensacolafrom29October1946 (Weston1947)through23 December 1954(Newman1955).HepointedoutthattheseobservationscouldhavebeenmadeoffeithertheAlabamaorFloridacoasts.WehavechosentoincludetheseobservationswiththeFloridaGulfcoastrecordsforthepurposesofTable9.TheonlyotherobservationsavailabletoImhof(1976b)werefourfrominland;weknowofonlytwosubsequentreports,one frominland,onefromoffshore.NumberspresentinAlabamawatersremainamystery.Duncan and Havard(1979ms) sawonlyonebirdoffAlabamainseveralyearsofobservation,buttheymentionedthatcommericalfishermeninAlabamareportlargenumbersofphalarope-likebirdsinwinter.1924lasthalfJan.1968 20Oct.19725-11Jan.colI.inlandatPickettSpringsseeninlandinShelbyCo.seeninlandatLakePurdy69Holt1924Purrington1969 Imhof1976b,James 1972

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RedPhalarope197210-15Sep.1photogr.inlandatLakePurdyImhof 1976b, 1973 1976Mayfemaleseen100kmoffAlabamacoastDuncanandHavard 1979ms1978 5 Nov.seeninlandatEufaulaRefugePurrington1979MississippiWeknowof only threerecordsfortheRedPhalaropefromMississippi.OnewasseeninlandontheHattiesburgsewagelagoonson9October1977(GatesandRunzo1978)andanotherwasseenthere12to15October1980(Mooreetal.1981).TheonlyrecordfromthecoastisofoneseenoffBiloxion 30September1978(Purrington1979).Itislikelythatthespeciesismorecommonoffshore.LouisianaWeknowofonlythreerecordsofRedPhalaropesinLouisiana:one wascollected12October1950ontheBatonRouge CampusofLouisianaStateUniversity;anotherwascollected16September1961 1 mi WofHollyBeach,CameronParish;andone wasseen29 November-lO December1970attheNatchitochesFishHatchery(Lowery1974).Lowery commentedthatthesephalaropesshouldoccurinoffshorewatersinwinter.TexasAccordingtoOberholser(1974),RedPhalaropesarecasualinTexasinfallandaccidentalinspring.Weknowofonly18recordsofthisspeciesinTexas,11ofwhicharelistedbyOberholser(1974).WelistbelowonlyrecordsfromcoastalareasandthosepublishedsubsequenttoOberholser(1974).Two-thirdsoftherecordswere madeinlandandallbutfourrecordsweremadeinthefallorearlywinter(8September-24November).Therearethreespringrecords(1April-31May), and one summerrecord(15July);thelatterwasprobablyanearlyfallmigrant.Thesefourrecordsareallfromthecoast,suggestingthatRedPhalaropesaremorecommontherethaninlandinspringandSummer.1935Apr.seenatCoveOberholser1974 19525-6Oct.seenatRockportOberhols-er1974 1952 9Oct.seenon WshoreofLagunaAtascosaOberholser1974 197319-28Sep.seeninlandatAustinsewage pondWebster1974a 197513Sep.-2seeninlandatAustinsewagepondsWebster1976a 1 Nov. 197515Nov.seeninlandatMitchellLake,SanWebster1976aAntonio197631MayfemaleseenatBolivarFlats,Webster1976cGalveston70

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1977 8Mar.11seen(incl.4females)offPortAransasRedPhalaropeWebster1977c 197815July19784Nov.photogr.atLagunaAtascosaNWRseeninlandatAustinsewagepondsWebster1979aWebster1979aSYNOPSISOFPRESENTDISTRIBUTIONANDABUNDANCEBreedingThe RedPhalaropehasthemostnortherlydistributionofthethreephalaropes,breedingcircumpolarlyinthenorthernHolarcticbetween60and82 Nlatitude(BOU1971).TheybreedfromnorthernandwesternAlaskathroughnorthernQuebec,Greenland,andIceland,alongtheArcticEurasiancoast,and onislandsfromSpitsbergeneasttoSiberia.WinterThewinterrangeoftheRedPhalaropeisatseaandbelievedtobeprimarilyoffthecoastsofsouthernSouthAmerica,westernAfrica,andsouthernArabia.MigrationRedPhalaropesmigratealongcontinentalcoastlinesandoverthebroadexpansesoftheAtlantic,Pacific,andIndianOceans.TheyarecommonmigrantsoffthesoutheasternAtlanticcoastoftheUnitedStatesthatoccasionallywinter.TheyoccurlaterinthefallandearlierinthespringalongtheAtlanticcoastthandoRed-neckedPhalaropes(LeeandBooth1979,Tables9,10).AlongthePacificcoastoftheUnitedStatesRedPhalaropesmigrateaboutone month later thandoRed-neckedPhalaropes.Thespringmigrationthereisshortandusuallynearthecoast;fallmigationislongerandbirdsarefoundoverawiderareaandfartherouttosea(Taylor1978).Migrantsusuallyoccursinglyorinsmallflocks.Rowlett(1980)reportedthatRedPhalaropesinthenorthernChesapeakeBightareseeninflocksof20to80birdsinApriland December,Scott(1959)observedthatmigrantsintheBayofFundyinAugustgenerallyoccurinflocksof15-30birds,butnotedtwoflocksthatcontainedatleast500birds.ThestatusoftheRedPhalaropeinthenorthernGulfofMexicoislargelyunresolvedandtherearefewrecentrecordsfromanystate,leadingDuncanandHavard(1979ms)toregarditascasualinoccurrence.Ontheotherhand,scatteredobservations(Weston1953,DuncanandHavard1979 ms)suggestthatitmaybeperiodically,perhapsregularly,commoninsomeoffshoreareas.Moreextensivepelagicsurveysareneededtodefineitstemporal,geographical,andnumericaldistributioninthenorthernGulf.AlthoughRedPhalaropesoccurcommonlyandareattimesnumericallyabundantoffthesoutheasterncoast,theproportionoftheworldorcontinentalpopulationofthespeciesthatoccurstheremustbeverysmall.71

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RedPhalarope_Table9.ApproximatenumberofRedPhalaropesrecordedby monthforthecoastalsoutheasternUnitedStates(a).State/regionJANFEBMARAPRMAYJUNJULAUGSEPOCTNOV DEC NorthCarolinaSouthCarolinaGeorgiaFlorida-AtlanticCoast28 114211447 30 503 3456141102142641961 16895160 125 503 2005Subtotal-ATLANTICCOAST47531539 34981226 45 787 6 286 708Florida-GulfCoast227 6678736 4 24 35 1033 Alabama 2 1 1 11Mississippi12Louisiana1 1 1 Texas-Coast112Subtotal-GULFCOAST229 667 98376 13 30 37 1034-------------------TOTAL-ALLAREAS704698637 3535 182745 790 36 323 1742-------------------(a)Birdsfounddeadinthefirst10daysofa montharearbitrarilyassignedtothepreviousmonth.Weassumedonlyonewasseenifthesourcedidnotspecifywhetheroneormore wasseen.Ifanindefinitenumber wasseen(e.g.,"afew","numbers","hundredstothousands"),orifarangewasgiven,wehaveassumedthesmallestnumberimpliedbythestatement.Table10.DatesofoccurrenceforRedPhalaropesinthecoastalsoutheasternUnitedStates.StateApproximatenumberofoccurrencesDatesofoccurrence(a)NorthCarolina43+ 4September4May(29May,19-30August)SouthCarolina9 15September22April(30May-11June)Georgia229October8 MarchFlorida-AtlanticCoast3714August?-27April(28July-2August)Florida-GulfCoast3613October-21April(7-8May) Alabama 6 10September15+January(May)Mississippi2 30September9OctoberLouisiana3 16September10 December Texas 6 5October1April(31 May,15July)(a)Exceptionaldatesofoccurrencearelistedinparentheses.72

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RedPhalaropeHABITATNestingRedPhalaropesnestprimarilyinlowcoastalArctictundra.Mayfield(1979)describedthenestinghabitatatBathurstIslandintheCanadianArcticassedge-mosstundrainterspersedwithfreshwaterponds.SimilarhabitatineasternSiberiawasdescribedbyKistchinski(1975)aspolygonalandtussockymoss-sedgetundrarichinswamps,lakes,andtemporaryponds.Thesebirdsusuallynestnearwater,butthe"ponds"maybeverysmall,sometimesonlya fewsquaremetersinextent(Portenko1921).Hohn(1969)notedthatRedPhalaropenestsareneverfoundmorethana fewmilesinland,pointingoutthatthisspeciesismorecoastalinitsnestingdistributionthanistheRed-neckedPhalarope.RedPhalaropesonBathurstIslandintheCanadianArcticusuallyplacetheirnestsamongsedgestallenoughtocoverthembutnotthickenoughtocompletelyconcealtheeggs(Mayfield1979).Mayfielddiscernednorelationshipbetweennest-siteandprominentfeaturesofthehabitat(e.g.,ponds,marshedge,boulders)exceptthatnestswerealwaysonthewetterportionsofflats.IneasternSiberia,Red Phalarope nestswerefoundmostly(84.9%,n=93)amongsedgesneartheedgesoftemporarypondsorontussocksinfloodedswamps. Theynestedlessfrequentlyalongpermanentlakes(7.5%),onisletsinsmalllakes(7.5%)orondrytussocksandridgesinthepolygonaltundra(7.5%)(Kistchinski1975).SitesreportedelsewhereincludethemossyrimofanIvoryGull(Pagophilaeburnea)nestonbareSeymourIslandnearBathurstIsland(MacDonaldinMayfield1979),pilesofseaweedintheopeninSpitsbergen(Munsterhjelmand LeRoiinLovenskiold1964),and (Lovenskiold1964).RedPhalaropes,likeotherphalaropes,tendtonestnearoneanother.Someobservers(Kistchinski1975,Mayfield1979)havereferredtothisasclusteringofnests;othersconsiderthespeciestobecolonial(Lovenskiold1964,Kozlova1961inMayfield1979).Hohn(1965 in Mayfield1979)statedthatallthreephalaropesnested.....mostlyinloosecoloniesoffourtoeightpairs;solitarypairsarerarer."Mayfield(1979)foundgroupsofthreetofivenestsatBathurstIsland,whilethelargestcolonyfoundbyLovenskiold(1964)inSpitsbergencontained25pairs.Nestdensity,inwhatMayfield(1979)consideredoptimumhabitatfrom1970to1976atBathurstIsland,averaged4.9nests/sqkm(12.9nests/sqmi),butthenumberofnestspresentinthe1sqkm(0.39sqmi)studyplotvariedfrom0to14(0-36.3nests/sqmi)fromyeartoyear.Mayfieldpointedoutthatnestdensitiesreportedinotherareasweremuchgreater.Citingotherauthors,hementioneddensitiesof7 and15pairs/sqkm(18.1and38.9pairs/sqmi)onsouthwesternBaffinIsland,andestimatesof20and12pairs/sqkm(51.8and31.2pairs/sqmi)forCambridgeBay,VictoriaIsland,andJennyLindIsland,respectively.At a0.67sqkm(0.26sqmi)studyareanearBarrow,Alaska,in1974 and1975,densitiesof9.0and24.0nests/sqkm(23.3and62.2nests/sqmi)werefoundbySchamel andTracy(1977).Oneparticularlyfavorablemarshheld90%ofallnestsatadensityof44.6nests/sqkm(115.5nests/sqmi).Mayfield(1979)alsonotedearlierpapersthatsuggestedconcentrationsof50-73

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RedPhalarope100males/sqkm(130-260males/sqmi)inSiberiannestingareas,butheremarkedthat"theunitsofareamaybemisleadingifthebestnestingareasareattenuatedanddiscontinous."FeedingOnthebreedinggrounds,RedPhalaropesfeedinorneartheirnestinghabitats.PhalaropesnearBarrow,Alaska,fedextensivelyattheedgesofshallow(1-2cm)ponds(SchamelandTracy1977).IneasternSiberia,incubatingmalesfedbothontemporarypondsandpermanentlakes;thelatterareusedlaterintheseason,afterthetemporarypondshavedriedup,andmaybefarfromthenests(Kistchinski1975).RedPhalaropesbreedingnearlagoonsandtheseaontheChukchiiPeninsulafednearthesurf,intheestuaries,and"whereverelsetheycanfindmuchfood."Whentheseawasroughthesebirdsfedbetweenthesurfandtheshorerun-off,butpreferredcalmerwaterssuchasbaysandlagoons(Portenko1972).NonbreedingandOffshorePrebreedingphalaropesarrlvlngatthebreedinggroundsontheChukchiiPeninsularemainaroundlagoonswheretheiceopenssoonerthanonthesea(Portenko1972).PostlayingfemalesandnonbreedingmalesineasternSiberiadepartfromthenestinggroundsandmovetolakesandcoastallagoons;bymid-Julyvirtuallyallphalaropesexceptincubatingmalesarefoundalongtheseashore(Kistchinski1975).Somefemalesoccurwelloffshore,particularlyamongicefloes(Portenko1972).Duringmigration,theRedPhalaropeisthemostpelagicofthephalaropes(Mayfield1979)andisseldomseeninshore.MigrantRedPhalaropesinthenorthernChesapeakeBight,northofNorthCarolina,occasionallyfedaroundsmallclumpsofsargassoweed(Sargassumsp.)andalongriptidescontainingdetritus(Rowlett1980).RedPhalaropesseenherewereusuallymorethan70km(44mi)fromshore.FOODANDFEEDINGBEHAVIORRedPhalaropes,likeRed-neckedPhalaropes,feedprimarilybysurfaceseizing,andlikethatspeciesoftenexhibitthecharacteristic"spinning"behavior.Ridley(1980)recentlydescribeddifferencesinfeedingtechniquesforthreesituations:(1)birdsonthewaterandfeedingonpreybelowthesurfacemayimmersetheheadandneckwiththebillpointingverticallydownwardwhiletippingup (muchinthemannerofadabblingduck);(2)birdsfeedingonlandandattheedgeofapoolprobeandpeckverticallydownward; and(3)birdsfeedingonorganismsonthesurfaceoronvegetationseizetheirpreywiththebeakheldhorizontally.Ridleytermedthesethreetechniques"deepfeeding","edge-feeding"and"surface-feeding",andconcludedthatfemalesonthebreedinggroundsinSvalbardconsistentlypreferdeep-feedingandthatincubatingmalesprefersurface-feeding.Ridleyalsopointedoutthat80%ofallpecksweresuccessfulinobtainingpreyandnotedthathighratesofpeckingwereobservedinsurface-feedingbirdswhileconsiderablyloweroneswereseeninedge-anddeep-feeders.LikeRed-neckedPhalaropes,RedPhalaropesaregregariousthroughouttheyear.Theyoftengatherinfeedingflocksoffourorfivebirdsduringthe74

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Red Phalaropebreedingseason,butincubatingmalesmayalsofeedsolitarily(Kistchinski1975).Mostfeedingprobablyoccursbyday,butthesephalaropeshavealsobeenseenfeedingonbrightnights(Portenko1972).Wetmore(1925)providedtheonlydetailedaccountofthedietofNorthAmericanRedPhalaropes,onthebasisof36stomachscollectedfromMaytoNovember,mostlyfromthePribilofIslands.Thesephalaropesconsumedalmostentirelyanimalfood.Insectsmade upmostofthediet;themostimportantfooditemswerebeetles(Coleoptera),whichmade up27.3%ofthebulk,andflies,whichaccountedfor22.7%.Aconsiderablevarietyofspecieswaseaten,butonlydung-flies(Scatophaga)werefoundinalargeproportion(38.8%)ofthestomachs.Crustaceawerealsofrequentlytakenandrepresented33.5%ofthebulkeaten.Amphipods werefoundin14stomachsandwater-fleas(Daphniidae)werefoundinthree.A numberofverysmallfisheswerealsoeaten;thosethatcouldbeidentifiedweresculpins(Cottidae).Afewspiders,molluscsandotherinsectswereeateninfrequently.LittlerecentinformationfromNorthAmericaisavailable.Hohn(1971)examinedthreestomachsfrombirdscollectedinJuneinwesternHudsonBay.Onecontainedlarvalchironomidsandsmalleumelibranchs;theotherscontainedseedsorwillowcatkinsandflowers.Wander(1981)observedaphalaropefeedingonthecarcassesofa Red Knot(Calidriscanutus)andaHerringGullatJamaicaBayWildlifeRefugeinNewYorkCity. LeRoi (inLovenskiold1964)remarkedonthedietof Red PhalaropesonSvalbard,basinghiscomments onanexaminationofthecontentsof58stomachs.Ofthese,67.2%containedCrustacea,primarilyeithergammaridsorostracods,dependingonwherethebirdswerecollected.Alsofoundinasignificantproportionofthestomachsweremolluscs(46.6%),dipteridlarvae(15.5%),andalgae(22.4%);afewarachnids,annelids,andonebeetlewerealsotaken. Rid ley(1980)baseda morerecentreportoffoodeatenonSvalbardonananalysisof21faecalsamplesobtainedduringthe1978breedingseason.Thedietconsistedprimarilyofsmallflies;chironomidsweremostimportant,butmycetophilidsalsowereeatencommonly.Theseandafewotherflyinginsectswerepresentin95%ofthesamplesandwereabundant(i.e.,morethan10individualremains)innearlyhalfofthem.Spiderswerealsoimportantandwerefoundin63.6%ofthesamples;theremainsofunidentifiedCrustaceawerefoundin22%.Ridley(1980)notedthatdifferentfoodsweretakenindifferenthabitats.PhalaropesfeedingbelowthesurfaceofthewatertookflylarvaeandCrustacea,surface-feedingbirdstookflies,andthosefeedingonlandorattheedgeofthewatertookspiders,flies,and(perhaps)Collembola.Inexperimentalstudies,Dodson andEgger(1980)foundthat Red Phalaropesprefertofeedonlargerzooplankton(particularlythosemorethanabout1mmlong).TheychosecladoceransovercopepodsandchoseDaphniamiddendorffiana,D.pulex,Eurycercuslamellatus,andDiaptomusbacillifermostfrequently.Twospeciesofsmallcalanoidcopepods(DiaptomusbacilliferandEurytemoracanadensis)weretakenmorefrequentlythanwaspredictedonthebasisoftheirsize.DodsonandEggersuggestedthatthismayhaveoccurred(1)becausephalaropesaresomehowspecializedforcatchingcopepods;(2)becausethesmaller,slower75

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RedPhalaropefooditemsaremoreeasilycaught;and(3)becausethesmallercopepodsaremorevisibletothephalaropesthanthelarger,moretransparentcladocerans.SUSCEPTIBILITYTOOILPOLLUTIONWefoundlittlerecordofRedPhalaropesbeingaffectedbyoil.Theyareagregarious,surface-feedingbirdthatoccursinlargenumbersofftheAtlanticcoast,however,andarepotentiallymoderatelysusceptibletooiling,atleastduringcoldweather.Thestronglypelagicdistributionofthisspeciessuggeststhatitwould bevulnerableonlytolargeoffshorespillsandnottoinshoredevelopment.King andSanger(1979)gavetheRedPhalaropeahighoilvulnerabilityindexvalueindicatingthattheyareamongthemostvulnerableshorebirdsoccurringinthenortheasternPacificregion.Ingeneral,phalaropesaremuchlesssusceptibletooilingthanothermarinebirds.Connorsand Gelman(1979)reportedthatnaiveyoungRedPhalaropesinitiallywereequallylikelytochoosepansoffoodthatdidordidnothaveathinfilmofoilonthesurfaceofthewater.Experiencedbirdspreferredclearwaterandfedforlongerperiods.Asaresultoftheseobservations,Connors and Gelmansuggestedthatphalaropescaneasilylearntoavoidoiledsurfaces.BIBLIOGRAPHY1982Conners,P.G.andK.G.Smith.1982.Oceanicplasticparticlepollution:suspectedeffectonfatdepositioninRedPhalaropes.Mar.Pollut.Bull.13:18-20.Orr,C.D.,R.M.P.Ward,N.A.Williamsand R. G.B. Brown.1982.MigrationpatternsofRed andNorthernPhalaropesinsouthwestDavisStraitandinthenorthernLabradorSea.WilsonBull.94:303-312.1981Green,J.C.1981.FirstRedPhalaropeforMinnesotaPoint.Loon53:52-53.Moore,J.R.,T.Fairley,D.HamiltonandP.Rodriguez.1981.Observationsofa RedPhalaropeinHattiesburg,Mississippi.MississippiKite11:15-17.Pigeon,P.1981.Observationd'unPhalaropeabecetroitPhalaropuslobatusetd'unPhalaropeabeclargePhalaropusfulicariusaAulnois-sous-Laon(Aisne).Alauda49:432.Wander,W.1981.RedPhalaropeeatingcarrion.WilsonBull.93:557.76

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RedPhalarope1980Chappell,M.A.1980.ThermalenergeticsofchicksofArcticbreedingshorebirds.CompoBiochem.Physiol.A.CompoPhysiol.65:311-317.Dodson,S.I.and D.L.Egger.1980.SelectivefeedingofRedPhalaropesonzooplanktonofarcticponds.Ecology61:755-763.Nistico,P.1980.RedPhalaropeintheMarylandPiedmontatHughesHollow.Md.Birdlife36:139.Ridley,M.W.1980.ThebreedingbehaviorandfeedingecologyofGreyPhalaropesPhalaropusfulicariusinSvalbard.Ibis122:210-226.1979Connors,P.G.andS.Gelman.inforagingexperiments.1979.RedPhalaroperespondstothinoilfilms only.)Pac.SeabirdGroupBull.6:43.Gustafson,D.1979.ARedPhalaropeinMilwaukee.PassengerPigeon41:136.Maurizio,R.1979.ThorshuehnchenaufdemSilserseebeiMaloja.[GreyPhalaropeatSilserLakenearMaloja.]Ornithol.Beob.76:134-135.Mayfield,H.F.1979.RedPhalaropesbreedingonBathurstIsland.LivingBird17:7-39.Roudybush,T.E.,C.R.Grau,M.R.Petersen,D. G.Ainley,K.V.Hirsch,A.P.GilmanandS.M.Patten.1979.Yolkformationinsomecharadriiformbirds.Condor81:293-298.1978 Brown, A.1978.ReactionofGreyPhalaropetoSparrowhawk.Scott.Birds10:22.Brown, B. andP.C.M.Latham.1978.GreyPhalaropeintheBayofPlenty.Notornis25:198-202.Gates,L.andJ.Runzo.1978.ARedPhalaropeinHattiesburg,Mississippi.MississippiKite8:2.Huber,H.1978.RedPhalarope--MilleLacsLake.Loon50:45-46.Mayfield,H.F.1978.UndependablebreedingconditionsintheRedPhalarope.Auk95:590-592.Taylor,W.B.1978.Seasonalstatus,populationsize,andhabitatutilizationofRed andNorthernphalaropesintheSouthernCaliforniaBight.(Abstractonly.)Pac.SeabirdGroupBull.5:74.77

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RedPhalarope1977Anderson,E.1977.RedPhalaropeatMoorhead. Loon 49:173.Savaloja,T.1977.Minnesota'ssecondRedPhalarope.Loon49:44-45.Schamel,D.andD.Tracy.1977.Polyandry,replacementclutchesandsitetenacityintheRedPhalarope(Phalaropusfulicarius)atBarrow,Alaska.Bird-Banding48:314-324.Schiemann,H.1977a.UberdasVorkommenderWassertreter(Phalaropodidae)indenbrandenburgischensachsischenundthuringischenBezirken,souieinBerlin.Beitr.Vogelkd.23:49-56.Sinclair,J.C.1977.Thephalaropes.Bokmakierie29:90-91.Wachtler,D.andG.Wachtler.1977.RedPhalaropeatMoorhead.Loon49:172-173.Watson,G. E.1977.AsecondspecimenofRedPhalaropefromDelaware.DelmarvaOrnithol.12:49.1976Brinkschroder,W.1976.BeobachtungeinesThorswassertreters(Phalaropusfulicarius)imReg.-Bez.Osnabruck.Vogelkd.Ber.Nieders.8:24.Risebrough,R. W.,G.E. Watson andJ.P.Angle.1976.A RedPhalarope(Phalaropusfulicarius)inbreedingplumage onAnversIsland.Antarct.J.U.S.11:226.Smith,F.T.H.1976.AnAustralianrecordoftheGreyPhalarope.Austr.BirdWatcher6:292-299.Wheeler,J.1976.A newwaderforAustralia--GreyPhalaropePhalaropusfulicarius.GeelongNat.13:52.1975Buhler,P.1975.deutschland.Thorshuhnchen(Phalaropusfulicarius)imPrachtkleid,inSudJahresh.Ges.Naturkd.Wuerttemb.129:78-80.Heise,G.1975.Thorwassertreter(Phalaropusfulicarius)inderUckermark.Beitr.Vogelkd.21:362-364.Kazama,T.1975.[AnewrecordofPhalaropusfulicariusfromJapanSea.]Tori23:29-30.[InJapanesewithEnglishsummary.]Kistchinski,A.A.1975.BreedingbiologyandbehaviouroftheGreyPhalaropePhalaropusfulicariusinEastSiberia.Ibis117:285-301.78

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RedPhalaropevanSwelm,N.D.1975.ForagingbehaviourofGreyPhalarope.BirdStudy22:52.1974Adams, R.J.,Jr.andD.E.Wood.1974.Firstspringrecordofa RedPhalaropeinMichigan.Jack-PineWarbler52:44.Boyle,G.L.1974.GreyPhalaropefeedingonearthworms.Brit.Birds67:352.Brown,J. F. 1974.GreyPhalaropes(Pha1aropusfulicarius)nearLagos.Bull.NigerianOrnithol.Soc.10:53.Gi11andt,L.1974.BeobachtungenaneinerThorshuhnchen-Popu1ation(Phalaropusfu1icarius)inSudwest-Island(Aves:Charadriiformes:Phalaropodidae).Abh.Verh.Naturwiss.17:55-83.[InGerman.J Lauerrnann, H.1974.EinThorshuhnchen(Pha1aropusfu1icarius)amMeise1dorferTeich.Egretta17:23-27.Marsh,C. andR.J.Hoder.1974.RedPhalaropeinWakeCounty,N.C.Chat38:71-72.Noer,H.and B.M.Sorensen.1974.Forekomenafstormfug1eProce11ariae,ThorshanePha1aropusfu1icariusogSabinernage,XemasabinivedBlavandshuk1963-71.Dan.Ornitho1.Foren.Tidsskr. Steinbach,R.1974.VorkommerderwassertreterPha1aropuslobatus(L.)undPhalaropusfu1icarius(L.)irnGebietderHaselbacharundEschefe1derTeichesowieamSpeicherbechtenWindisch1euba.Abh.Ber.Naturkundemus.'Mauritanium'8:333-338.1973But'ev,V.T.1973.[OnthenestingoftheGreyPhalaropeinthenorthofNovaya Zem1aIs.]Ornito1ogiya10:329-330.[InRussian.]Fitzpatrick,J.1973.GreatCrestedGrebesattackingGreyPhalaropeandStormPetrel.Brit.Birds66:365.Kistchinskii,A.A. andY.I.Chernov.1973.NotesonthefoodsoftheGreyPhalaropeinthetundraofeastSiberia.MoscowSocietyofNaturalists.Fauna andEcologyofWaders.Issue1:61-64.Kocian,L.1973.Novyvyskytlyskonohap1oskozobehoPhalaropusfu1icarius(L.,1758)naSlovenska.[NewrecordofPhalaropusfu1icarius(L.,1758)inSlovakia.]Zb.Slov.Nar.Muz.Prir.Vedy19:197.[InCzechwithGermansummary.]79

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RedPhalaropeLok,C.M.1973.SomeobservationsontheGreyPhalarope.Sterna12:284-285.Rettig,K.1973.Thorswassertreter(Phalaropusfulicarius)amRysumer Nacken [RedPhalaropeatRysumNeck.]Ornithol.Mitt.25: 222.[InGerman.]Sannazzaro,A.1973.IIPhalaropusfulicariusinPiemonte. inPiedmont.]Riv.Ital.Ornitol.43:462-466.[Phalaropusfuli[InItalian.]----Schmitz,J.-P.andJ.Weis.1973.Premiereobservationetcaptureduphalaropeabeclarge(Phalaropusfulicarius)dansIeGrand-DuchedeLuxembourg.Regulus11:35.Stelzer,H.1973.UberdasVorkommenderWassertreter(Phalaropodidae)inderSchweizundihrenRandgebieten.Ornithol.Beob.70:157-170.[InGermanwithFrenchsummary.]vanAvermaet,G.1973.Franjepoten.Wielwaal39:216-220.Webster,R.E.1973.Shortnotes.GreyPhalarope.Distribution.Lammergeyer15:76.1972Portenko,L.A.1972.PtisyChukotskogopoluostrovaiostrovaVrangelya.[BirdsoftheChukchiiPeninsulaandWrangelIsland.]Volume1.xvand446pp.[1981translation.AmerindPubl.Co.,NewDelhi.]Prigge, R. 1972.HeobachtungeinesThorwassertreters(Phalaropusfulicarius)amTreuel/ElbeimBezirkMagdeburg.Beitr.Vogelkd.18: 434.Sehl,R.H.1972.RedPhalaropeinBucksCounty.Cassina53:46-47.Summerfield,D.1972. Red PhalaropeinDecember.KentuckyWarbler48:18.1971Bond,S.I.1971. Red PhalaropemortalityinsouthernCalifornia.Calif.Birds2:97.Danks,H.V.1971.AnoteontheearlyseasonfoodofArcticmigrants.Can.Field-Nat.85:71-72.Fantin,G.1971.NotiziedalVeneto:ilFalaropobeccolargo.[ReportsfromVenice:theGreyPhalarope.]Riv.Ital.Ornitol.41:17-24.[InItalian.]Hohn, E. O.1971.ObservationsonthebreedingbehaviourofGreyandRed-neckedphalaropes.Ibis113:335-348.80

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RedPhalaropeKing,B.1971.GreyPhalaropeon autumnpassageplungingintowaves.Brit.Birds64:29-30.1970Rettig,K.1970.ThorswassertreterbeiEmden.Ornithol.Mitt.22:47.1969 Hohn,E.O.1969.Thephalarope.Sci.Am.220:104-111.Kittinger,H.H.andJ.N.Carusos.1969.SecondinlandRedPhalaroperecordinAlabama.Ala.Birdlife17:50-51.Schiemann,H.1969.Suddeutschland.UberdasVorkommendenWassertreter(Phalaropodidae)inVogelwelt90:184-188.Schmidt,G.D.1969.Polymorphuspetrochenkoisp.n.(Acanthocephala)fromtheRedPhalarope,PhalaropusfulicariusL.,inAlaska.J.Parasitol.55:335-336.Sutton,G.M.1969.TheRedPhalaropeinOklahoma.Bull.Okla.Ornithol.Soc.2:26-28.1968Bengtson,S.-A.1968.BreedingbehaviouroftheGreyPhalaropeinWestSpitsbergen.VarFagelvarld27:1-13.Ruppell,G.1968.Phalaropusfulicarius(Phalaropodidae)Bruten.EncyclopaediaCinematographica.E1381. AngabenzumFilmandFilmenhalt,pp.1-6.[InGerman. ] 1967Smith,W.andR.Klauke.1967.AsightrecordoftheRedPhalaropeinAlberta.BlueJay25:25.Spasskij,A.A.andJ.N.Konovalov.1967.Anomotaeniareticulatasp.n.Cestoidea,Dilepididae,aparasiteofPhalaropusfulicarius.Vestn.Zool.1967:43-48.1966 Hohn, E.o.1966.Ringing(banding)andrecoveriesofphalaropes--asummaryofpresentlyavailableinformation.Bird-Banding37:197-200.81

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Cooch,F.G.1965.fulicarius).RedPhalarope1965AnexampleofsinistralisminRedPhalaropes(PhalaropusAuk82:276-277.Bohn,E.o.1965.DieWassertreter.NeueBrehm-Bucheri359.ZiemsenVerlag,Wittenberg.60pp.Woodward,I.D.1965.ThestatusanddistributionoftheGreyPhalaropeinHertfordshire.BirdsIllust.11:150.1964Acton,J.R.1964.RedPhalaroperecordedfromeasternWashington.Murrelet45:11.Lovenskiold,H.L.1964.Avifaunasvalbardensis,withadiscussiononthegeographicaldistributionofthebirdsinSpitsbergenandadjacentislands.Skr.Nor.Polarinst.No.129.460pp.Prozesky,o.P.M.1964.NotesontheGreyPhalarope(Phalaropusfulicarius(LinnaeusfoundatVoortrekkerhoogte,Pretoria,Transvaal.Ostrich35:70.Shuler,J.B.1964.SouthCarolina.SightrecordofaRedPhalaropeinCharlestonCounty,Chat28:30.1963 Lakeman,M.1963.RedPhalaropeatHuntingCreek?Atl.Nat.18:250.1962Breiding,G.H.1962.RedPhalaropeinWestVirginia.WilsonBull.74:288.1961Alderson,G.1961.InlandoccurrencesoftheRedPhalaropeinOregon.Condor63:97-98.Cohen,E.andJ.H.Taverner.1961.ColourationofsoftpartsofGreyPhalaropes.Brit.Birds54:116-117.1960Baldwin,W.P.1960.RedPhalaropeatCape Romain,S.C.Chat24:76.Cowcill,L.andA.Smith.1960. GreyPhalaropeblowingbubbles.Brit.Birds53:574.Milne,B.S.1960.HugeflocksofGreyPhalaropesintheIslesofScilly.Brit.Birds53:403.82

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RedPhalaropeOlier,A.1960.28:69-70.PhalaropePhalaropusfulicariusauMarocoriental.[InFrench.11959AlaudaGeorge,U.1959.Thorwassertreter(Phalaropusfulicarius)beiHamburg.Ornithol.Mitt.11:235.Gilmore, R. M.1959. Red Phalaropeinmid-easternPacific.Condor 61:227.Lee,H.J.1959.GreyPhalaropeatLakeNakuru.J.EastAfr.Nat.Hist.Soc.23:175.Wilcox,L.1959.LargeflightsofRedPhalaropeson LongIsland.Kingbird9:24-25.1956Christensen,N.H.1956.Odinshanens(PhalaropuslobatusL.)ogThorshanens(PhalaropusfulicariusL.)forekomsti Danmark. [TheoccurrenceinDenmarkoftheRed-necked Phalarope andtheGreyPhalarope.]Dan.Ornithol.Foren.Tidsskr.50:191-206.[InDanishwithEnglishsummary.]1955 Andrew,D.G.1955.Note onthedisplayoftheGreyPhalarope.Brit.Birds48:546.Geroudet,p.1955.LepassagedesPhalaropesenSuisseromande.NosOiseaux23:42-47.[InFrench.]1954Sage,B.L.1954.Onthebill-colouroftheadultGreyPhalaropeinwinterplumage.Bull.Br.Ornithol.Club74:12.Young, C.G.1954.Phalaropes,stormpetrelsandBarnOwlatsea.Ibis96:311.1953Adler,o.1953.ThorshuhnchenimBinnenlandOsterreichs.Ornithol.Mitt.5:55.Musacchia,X.J.1953.Astudy 0f thelipidsinArcticmigratorybirds.Condor55:305-312.Seiple,G.W.1953.RedPhalaropeinUtah.WilsonBulL65:205.Stanford,W.P.fulicarius.1953.WinterdistributionoftheGreyPhalaropePhalaropusIbis95:483-491.83

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RedPhalaropeWeston,F.M.1953. Red Phalarope(Phalaropusfulicarius)winteringnearPensacola,Florida.Auk70:491-492.1952Kennedy,J.1952.AnunusualrecordoftheGreyPhalaropeinSouthernRhodesia.Ostrich23:122.1951 [Wood,J.D.].1951.Studiesofsomespeciesrarelyphotographed.XXXI.The Grey[and]Red-neckedPhalarope.Brit.Birds44:235-236.1947Weston,F.M.andM.M.Wernicke.1947.TheRedPhalaropeinFlorida.Auk64:473.1946Walker,W.M.1946.ARedPhalaropeinTennessee.Auk63:102.1944Wilson,W.1944.The GreyPhalaropeonDeeside.Proc.LiverpoolNat.FieldClub83:19-20.1943 Lowe,W.P.1943.GreyPhalaropesoffthewestcoastofAfrica.Ibis85:103.Wust,W.1943.lborshuhnchen,Phalaropusfulicarius(L.)imIsmaningerTeichgebiet.Ornithol.Monatsber.51:43.1942Stanford,J.K.1942.GreyPhalaropesoffthewestcoastofAfrica.Ibis84:520.1941Einarsen,A.S.1941.Theeffectsofstormonshorebirds.Murrelet22:36.Slipp,J.W.1941.NotesontheRedPhalaropeinthe Murrelet22:38.1939Griscom,L.1939.MigrationoftheRedPhalaropeoffMassachusetts.Auk56:185.84

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Holmes,P.F.1939.ofphalaropes.RedPhalaropeSomeoceanicrecordsandnotesonthewinterdistributionIbis(14thSer.)3:329-342.1938Michael,C.W.1938.FeedingbehaviorofaRedPhalarope.Condor40:181.Stoner,E.1938.RedPhalaropeatBenicia,California.Condor40:182.Wilcox,L.1938.Island,N.Y. AflightofRedPhalaropes(Phalaropusfulicarius)onLongProc.Linn.Soc.N.Y.49:6063.1937 Low,G.C.1937.Grey andRed-neckedPhalaropesintheArabianSea.Ibis(14thSer.)1:866.Meinertzhagen,R.1937.Grey andRed-neckedPhalaropesinArabianSea.Ibis(14thSer.)1:667.1936Chislett,R.1936.GreyPhalaropeinYorkshire.Naturalist(Lond.)1936:7.1935Brooks,A.1935.OnPhalaropusfulicariusjourdainiIredale.Ibis(13thSer.)5: 887-888. Gabrielson,I.N.1935.DestructionofRedPhalaropes.Murrelet16:27-28.Jewett,S.G.1935.ARedPhalaropedisaster.Murrelet16:15-16.Pearse,T.1935.RedPhalaropeson VancouverIsland.Murrelet16:16-17.1934Jourdain,F.C.R.1934.ThestatusofPhalaropusfulicariusjourdainiIredale.Condor36:220.Peters,J.L.1934a.ThestatusofPhalaropusfulicariusjourdainiIredale.Condor36:85.1934b.Furthercomment onPhalaropusfulicariusjourdaini. ------Condor 36:220-221.Sprunt,A.,Jr.1934.TheRedPhalaropeoffSouthCarolina.Auk51:374.85

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RedPhalarope1933Ingram,G.C.S.andH.M.Salmon.1933.Ruffand GreyPhalaropeinCarmarthenshire.Brit.Birds27:206-207.Taning,A.V.1933.Thewinterquartersofthephalaropes.Ibis(13thSer.)3:132-133.Wilson,W.andF.C.R.Jourdain.1933.GreyPhalaropesinCheshire,Hampshire,Sussexand Devon.Brit.Birds27:207-208.Young, C.G.1933.Thewinterquartersofthephalaropes.Ibis(13thSer.)3:548.1932Urner,C. A.1932.PhalaropesinNewJerseyinspring.Auk49:475-476.1930Trautman,M.B.andC.F.Walker.1930.AnotherrecordoftheRedPhalaropeinOhio.Auk47:249-250.1928Hine,J.S.1928.TheRedPhalarope(Phalaropusfulicarius)inOhio.Auk45:94-95.Schorger,A.W.1928.ThewinteringareaoftheRed andNorthernPhalaropes.Auk45:206.1926Friedmann,H.1926.RedPhalaropeatsea100mileswestofGibralter.Auk43:234.Grote,H.1926.ZurVerbreitungvonPhalaropuslobatus(L.)undPhalaropusfulicarius(L.).Ornithol.Monatsber.34:122.1925Coles,R.J.1925.Sea-birdsatCapeLookout,NorthCarolina.Auk42:123-124.Meinertzhagen,R.1925.Thedistributionofthephalaropes.Ibis(12thSer.)1:325-344.Rapine,J.1925.Lesphalaropes.Rev.Fr.Ornithol.17:54-56.Wetmore, A.1925.FoodofAmericanphalaropes,avocets,andstilts.U.S.Dep.Agric.Bull.1359.20pp.86

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RedPhalarope1924Holt,E.G.1924.Phalaropusfulicarius:a newbirdforAlabama.Auk41:601.Roberts,A.1924.RedPhalarope(Phalaropusfulicarius)inAfrica.Auk41:600-601.1923TschusizuSchmidhoffen,V.1923.Phalaropusfulicarius, I. lobatusundAcanthisflavirostrisinOberosterreich.Ornithol.Monatsber.31:62.1922Jourdain,F.C.R.1922.[Remarks ontheSpitsbergenphalarope.]Bull.Br.Ornithol.Club.42:55-56.Ross,R.C.1922.RedPhalaropeinsouthernCalifornia.Condor 24:66-67.Welch,L.W.1922.VermillionFlycatcherandRedPhalaropeatLongBeach,California.Condor 24:62.1919 Laubmann, A.1919.ZurnomenklaturderGattungPhalaropusBrisson1760.Ornithol.Monatsber.27:75-76.Stone,W.1919.OccurrenceoftheRedPhalaropeinPennsylvania.Auk36:419-420.1915Haviland,M.D.1915.NotesonthebreedinghabitsoftheGreyPhalarope.Brit.Birds9:11-16.1914Bridge,L.E.1914.RedPhalarope(Phalaropusfulicarius)offBostonHarborinsummer.Auk31:536.1913 Lowe,P.R.1913.[GreyPhalaroperecordedfromneartheCape VerdeIslands.]Bull.Br.Ornithol.Club33:40-41.1910Willett,G.1910.RedPhalaropeinsouthernCaliforniainwinter.Condor12:175.87

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Ked PhalaropeHaag,F.1909.PlattschnabligerWasse.rtreter(PhalaropusfulicariusL.).Oologie19:102-104.------Banks, R. C.1908.GreyPhalaropeinCo. Wexford.Brit. Birds2:240-241. Champernowne.. A.w.1908.GreyPhalaropeinsUlIIIIlerinDevonshire. Brit. Birds2:204-205.Dyche,L.L.1906a.The Red Phalarope(Crymophllusful1cariusLinn.).birdforthe Kansas list.Trans. Kans. Acad. 1906b.The RedPhalarope(Crymophllusfulicarius),a new birdforthe --Kansas list. Auk 23:220.------Thayer,G. H.1902.The Red PhalaropeinNorthCarol1na.Auk19;285-286.Patten, J.1901.ThenaturalhistoryoftheGreyPhalarope(Phalaropusful1carius). Irish Nat.10:53-67.---Wayne,A.S.1901.The Red Phalarope(Crymophilusfulicarius)onthecoastofSouthCarolina. Auk 18:271.------AdultLaughingGullin-breedingplumage.PhotographbyRogerB.Clapp.88

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LAUGHINGGULL(Larusatricilla)[FR:Goelandatricille,SP:Gaviotarisuena,SW:Amerikanskskrattmas]GENERALDISTRIBUTIONNorthAmericaLaughingGullsareamongthemostabundantcoastalbreedingbirdsofthesoutheasternUnitedStates.TheyarelargelyrestrictedintheirbreedingdistributiontoNorthAmericaandtheCaribbean.InCanadathesegullsbreedverylocallyinNovaScotiaandsouthernNewBrunswick(Godfrey1966).WithintheUnitedStates,thegreatmajorityofbreedingbirdsarefoundalongtheAtlanticandGulfcoastsfromMaineandMassachusettssouthtoTexas(AOU1957).WithinthisareanonebreedinRhodeIsland,Connecticut,Georgia,Alabama,andMississippi,but,for the firsttimeinmanyyears,thespecieshasreturnedtobreedonLongIsland,NewYork(Buckleyetal.1978a,PostandRiepe1980).InthePacificstatesLaughingGullsformerlybredonislandsatthesouthendoftheSaltonSeainsoutheasternCalifornia(Small1974).InMexicotheybreedatleastonAlacranReefandCayosArcasinthesouthernGulf(Paynter1955),andalongthenorthwesterncoastinSinaloaandSonora(AOU1957).IntheWestIndies,theybreed"onmanyofthesmallerislandsorcays" (Bond1956),andDeweyandNellis(1980)reportedthattheybreedat12of27seabirdcoloniesintheU.S.VirginIslands.Detailsofdistributionandpopulationsizesintheseareasarepoorlyknown.LaughingGullswintersouthonthePacificcoastfromsouthernMexicotoLima,Peru(ca.12S).OntheAtlanticsideofthecontinenttheywinterfromNorthCarolinasouthalongtheAtlanticandGulfcoaststothemouthofthe funazon neartheequatorinBrazil(AOU1957,Blake1977).SomealsowinterintheCaribbean,butthesegullsarenotcommonfromOctobertoMarchintheWestIndies(Bondi956)andfromNovembertoMarchinTrinidadandTobago(ffrench1973),suggestingthatthesearenotmajorwinteringareas.WorldDistributionLaughingGullsbreedonislandsoffthecoastofVenezuela(LaOrchila,LosRoques,LasAves)(Blake1977),onLittleTobago,andprobablyonSt.GilesIsland,and onthenorthcoastofTobago(ffrench1973).BreedingwasalsorecentlyreportedonGrandConnetableIslandandtheBatturesdeMalmanoury,FrenchGuiana(Condamin1978).Nonbreedingbirdsoftenremaininwinteringareasduringthenorthernbreedingseason,andwanderingbirdsmaystragglewelloutsidethebreedingorwinteringranges.LaughingGullshavestrayedfrequentlytoGreatBritain(BOU1971,Preston1975,Verrall1977,Rogersetal.1982),andalsohavewanderedtoSweden,France(AlbrektssonandLindberg1967),andMorocco(Kennerley1979).IntheNewWorldthesegullsalsohavewanderedtonorthernOntario(Godfrey1966),southernSaskatchewan(SavileandSavile1976),andtoGreenland(AOU89

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LaughingGull1957).TheyhavealsobeenfoundinthePacificOcean onHawaii 1972),onJohnstonAtoll(AmersonandShelton1976),thePhoenixandLineIslands(King1967),andWesternSamoa (Museetal.1980,MuseandMuse1981).OnebirdbandedasaflightlessjuvenilenearBarnegatLight,NewJersey,wasrecoveredinKauai,Hawaii,lessthanfivemonthslater(TelferandShisler1981).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESNorthCarolinaLaughingGullsoccurinNorthCarolinathroughouttheyear.Theynestinlargenumbersbutareuncommoninwinter.Pearsonetal.(1942)believedthatasmanyas15,000bredtherein1939.About26,790bredin20coloniesinNorthCarolinain1973(SootsandParnell1975a),andabout25,030bredin20coloniesin1977.Figuresfor1976(Table11)aresimilartothosereportedforotheryears.LaughingGullsnowbreedinNorthCarolinasouthtotheCapeFearRiver(Map1)andthebreedingpopulationisbelievedtobestable(ParnellandSoots1979ms).Thelargestcoloniesreportedin1977wereasfollows:oneon adredgespoilislandinRoanokeSound,DareCounty,thathadabout1,850breedingbirds;oneonGullIsland,DareCounty,withabout4,790;oneonBeaconIsland,HydeCounty,withabout2,850;onewithabout3,740onMorganIsland,CarteretCounty;andonewithjustover7,000birdson adredgedislandinCarteretCounty.Thesefivecoloniesbetweenthemcontainedoverfour-fifthsoftheLaughingGullnestscensusedin1977,andthelastislandlistedwasthelargestcolonyinthestatein1976and1977(ParnellandSoots1979ms).MostLaughingGullsreturntoNorthCarolinabylateMarchandmostbreedingoccursfromMaythroughlateJuly(ParnellandSoots1979ms).WinteringpopulationsaresmallcomparedtothoseoccurringinFloridaandthecoastofTexas,butreachatleastthelowthousands(Map2).SouthCarolinaLaughingGullsareabundantalongtheSouthCarolinacoastthroughouttheyear.EggsareusuallylaidinlateMayorearlyJune(SpruntandChamberlain1949).Littleisknownofbreeding,transient,orwinteringpopulations.In1976,about10,940breedingbirdswerefoundnestingatthreelocalities:White (ca.2,940birds),Deveaux Bank(ca.5,000),andBirdIsland(ca3,000)(Portnoyetal.1981).Thisfigureisprobablyagrossunderestimateofthetotalnumberbreedinginthestate,however,becauseLaughingGullsbreedthereatmorethanthreelocalities.Blusand Lamont(1979)reportedthatLaughingGulls"nestonalmostalloftheSouthCarolinabarrierislands",andstatedthatroughestimatesin1975suggestedmorethan10,000pairsbredthere--abreedingpopulationmorethatdoublethatreportedbyPortnoyetal.(1981).RelativelyfewbirdshavebeenreportedonrecentChristmasBirdCounts(Map2).GeorgiaDentonetal.(1977)regardedLaughingGullsasabundantalongtheGeorgiacoastinspringandfall,butasuncommoninwinter.Burleigh(1958)consideredthespeciesmostabundantasaspringandfallmigrant.LaughingGullsarenotknowntobreedinGeorgiaandlittleattentionhasbeen90

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3040-7660-76120-76400-76to7870-76120-76 DRY TORTUGASBIRD NAME,,)I ''----\\ .., .--..,--(..,.., -.-" '-8"" -.J'88" MEXICO JACKSON GULFOF '"". I ).,,)r-.J --.s2228-76'A1250-774000-734384-75 9 000-77 12000-7610000-764000-738648-751026-7316308-771950-739601400-77940xE TBreedingRangeMapforSoutheasternUnitedStatesDALLASNumbersinboxesdenotemaximum estimates of breeding birds at coloniesinrecent years.Firstfigure indicates maximum number of birds. Second figure indicates yearinwhich estimatewasobtained. North Carolina colony sites of less than 150 breeding adults not censused. Texas colony sitesofless than 1,000 breeding adults not censused. G:;XAMPlE320-77, MAlNO.YEAR( -0-0/\S'/ 2030-7520000-731i __ It5780-773744-777672-74Map1

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26" LAUGHINGGULLBIRD NAME'88".&rF3FAsh,bo-----r. ... -.--,------"'--\, ,\ ., <, ., 32I ,I ., II _TGO_., .""....CKSON-V 30 -0 I\ ."" GULFOFMEXICO ....... I , r-.J--. ,"'.,)'...r-? ).
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LaughingGullTable11.RecentestimatesofLaughingGullpopulationsnestinginthesoutheasternUnitedStates(a).StateNorthCarolinaSouthcarolinaGeorgiaFlorida-AtlanticCoastFlorida-Keys(c)Florida-GulfCoast(d)AlabamaMississippiLouisianaTexasNumberfoundbreedingin1976(b)24,71010,940noneca.4,640ca.2,800ca.58,450none none56,090105,620263,250Percentofsoutheasternbreedingpopulation9.44.21.81.122.221.340.1(a)Insimilartablesinotherspeciesaccountswecomparedestimatesmadein1976withthosemadeotheryearstosuggestthemaximumbreedingpopulationsthatmaybepresentduringanygivennestingseason.LaughingGullpopulationsaresopoorlyknown,however,thatweareunabletodosointhisinstance.TheTexasCoastalWaterbirdSurveyrecordedbreedingpopulationsin1973and1977thatwerealmosthalfagainaslargeasasthatrecordedin1976andlargerpopulationshavedoubtlessoccurredinothersoutheasternstates.Consequently,maximumbreedingpopulationsinthesoutheastmaybeasmuchasseveralhundredthousandlargerthanisimpliedbythistable.(b)Totalsroundedtothenearest10.(c)About2,250breedingbirdswerepresentat13coloniesinsouthernFlorida(Portnoyelal.1981).Anearlierreportofthissurvey(KushlanandWhite1977)listed2additionalcoloniesandanestimatedtotalbreedingpopulationofabout2,800birds.The twoadditionalcolonies,UpperArsnickerKey(4breedingbirds)andNestKey(ca.500breedingbirds),evidentlywereaccidentallyomittedduringthepreparationoftheatlasbyPortnoyetal.(1981).(d)Thisfigureishighlyspeculative.Thetruenumbermaybeconsiderablymore(R.W.Schrieber,pers.comm.).paidtothesizeofmigrantandwinteringpopulations.Florida-AtlanticCoastandKeysLaughingGullsarecommontoabundantinthisareathroughouttheyear.Formerlyrareasabreederalongthecoast,severalthousandsnownestthere(Kale1979msa).A1976surveyofthe93

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LaughingGullFloridaAtlanticcoastsouthtoKeyWest(Portnoyetal.1981)revealedabout2,250nestingbirdsin13colonies.AllofthesecolonieswereeitherinFloridaBayatthesoutherntipofFloridaorintheKeys.Informationaboutnestingpopulationsfurthernorthalongthecoastisincomplete.Varioussources(Map1,Kale1979msa)indicatethattheprimarybreedinggroundfarthernorthisonspoilislandsintheBananaRiver,MerrittIslandNWR,andsuggestthatthebreedingpopulationisconsiderablylargerthanthatreportedforsouthernFlorida.BecauseFloridaisoneoftheprimarywinteringgroundsforLaughingGullsfromcoloniestothenorth,winteringpopulationsareimmenseandarefarlargerthanbreedingpopulations.ConcentrationsreportedonChristmasCountsfromboththeAtlanticandGulfcoastsarelargerthanhavebeenreportedfromanyotherpartofthe range(Map2).PopulationswinteringinFloridamorethantripledbetween1966and1972(SchreiberandSchreiber1977)withpeakpopulationspresentfromNovembertoMarch(Kale1979msa);Kalenotedthatover18,000LaughingGullswererecordedononeChristmasCountinDadeCounty.Florida-GulfCoastLaughingGullsareabundantontheFloridaGulfcoastyearround.Thelargestknownbreedingandwinteringpopulationsofthespeciesoccurthere(Maps1,2),butthetotalnumberspresentarepoorlyknown. ThebreedingpopulationofthisspeciesintheTampaBay-CharlotteHarborareashowed ahugeincreasefromthemid-1960'stothemid-1970'sthatSchreiberandSchreiber(1977)suggestwasduetoincreasedamountsoffoodavailableatgarbagedumpsandtoincreasedexploitationoftheseareasbyLaughingGulls.Populationscontinuetoincrease(R.W.Schreiber,pers.comm.).MostnestingoccursbetweenAprilandSeptember.PeakegglayingusuallyoccursduringthefirstweekofMay. MostyoungfledgeinJuneandleavethebreedingareasbyearlyAugust(SchreiberandSchreiber1978).AlabamaLaughingGullsarecommontoabundantalongtheAlabamacoastallyearbutarenotknowntobreedthere.Numbersaremuchsmallerinmidwinterandmid-summerthaninmigration(Imhof1976b).Concentrationsofmorethan10,000havebeenseen(Table12),butfewdataareavailableonnumberspresentinthestate.BandrecoveriesfromNewJersey,Virginia,andNorthCarolina(Imhof1976b)showthatatleastsomeproportionofthefallandwinterbirdscomefromAtlanticcoastcolonies.MississippiAlthoughfrequentlyabundantinMississippiwaters,LaughingGullshavebeenknowntobreedthereonlyonce(Jacksonetal.1980).Williams(1962a)foundacolonyofabout6,000pairsnestingonthewesternendofPetitBoisIslandon24June1962.Youngwereseenduringthisvisit,someofwhichhadfledgedby18July,and somefresheggswerestillpresentonthislaterdate.InsomeyearsLaughingGullsarenearlyabsentfromlateNovembertoearlyMarch(Burleigh1944).Mostcommonduringmigration,theyarelessabundantfromearlyMaythroughJune,aperiodwhenmostbirdsareprobablyconcentrated94

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LaughingGullneartheirbreedingcoloniesinLouisiana.LouisianaLaughingGullsareabundantalongthecoastthroughouttheyearbutarerarelyseeninland.Smallernumbersarepresentduringthewinterthanduringthebreedingseason(Lowery1974).A1976surveyofthenorthernGulfcoastrevealedabout56,000breedingLaughingGullsin19coloniesinLouisiana(Portnoy1977).Morethanhalf(ca.34,650birds)thetotalbreedingpopulation was foundinasinglecolonyonRaccoonPoint(#602031).Anecdotalreportsofformercoloniessuggestthatlargerpopulationsmayhaveoccurredthereinthepast.About47,000breedingadultswerepresentontheislandsoftheBretonandLouisianaAudubonreservationsin1908(Kopman1908a)andanother35,000birds,mostlyyoung,werepresentontheEastTimbalierreservation(Kopman1908b).Lowery(1974)statedthatthemainnestinggroundsareintheChandeleurIslandsandonthemudlumpsatthemouthoftheMississippiRiver.ThebreedingseasonisfromAprilthroughJulywitheggsrecordedfromearlyApril(Oberholser(1938)throughmid-July(Portnoy1977).TexasLaughingGullsarethemostabundantbreedingmarinebirdincoastal havebeenrecordedfrom8Aprilto7July(Oberholser1974)andyoungbirdswerestillpresentinmid-September1977incoloniesinandnearCorpusChristiandthenorthernLagunaMadre(Chaneyetal.1978).LaughingGullsbreedallalongtheTexascoastbutaremostabundantontheupperGulfcoast(Blacklocketal.1978ms).Thelargestcoloniesinrecentyearshavebeenonethatcontainedabout12,000birdsonSouthBirdIsland,KlebergCounty,in1976;onewithabout10,000onislandssouthofthislocalityin1976;onewithabout20,000onDangerIsland,NeucesCounty,in1973;onewithabout24,000onNorthDeerIslandandnearbyspoilislandsinGalvestonCounty,in1976;anotheronSouthDeerIslandthatcontainedabout20,000in1973;andoneonPelicanIsland,GalvestonCounty,thatheldabout50,000breedingbirdsin1973andabout10,000in1976(Blacklocketal.1978).Thefourlargestcoloniesin1976betweenthemcontainedabout56,000breedingbirds,ormorethanhalfofallthosebreeding in thestate.AlthoughbreedingLaughingGullsaremoreabundantinTexasthananywhereelseinthesoutheast,Blacklocketal.(1978ms)believedthepopulationtobedecliningandfeltthatthespeciesnumbersshouldbecarefullymonitored.LaughingGullsarealsoabundantmigrantsandwinterresidentsontheTexascoast(Table12).Five-yearaverages(Map2)forrecentChristmascountsinsevencoastalareashavetotaledabout16,000birds.Consideringtheextremelysmallareathatthesecountscovered,totalwinteringpopulationsmustbeatleastanorderofmagnitudelarger.SYNOPSISOF PRESENT DISTRIBUTIONANDABUNDANCEBreedingLaughing &ulls breedsolelyintheNewWorld.MostbreedingpopulationsarefoundalongtheAtlanticcoastoftheUnitedStates,alongthe95

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LaughingGullTable12.PeakconcentrationsofmigrantandwinteringLaughingGullsinthecoastalsoutheasternUnitedStates.NumberDateseenseenLocalitySourceNORTHCAROLINA1971 24 Nov.10,000vicinityofMoreheadCityTeulings1972aFLORIDA197118Dec.30,000SebastianInletStevenson1972awinter80,000StPetersburg-TampaAreaKale1979msaALABAMA1956 6Oct.11,000+MobileBay Imhof 1976b 195928Dec.1,631MobileImhof 1976b MISSISSIPPI 1960 8Sep.1,400PassChristianGandyandTurcotte1970 196022Sep.2,000GulfportGandyandTurcotte1970 196013Oct.1,000DeerIslandGandyandTurcotte1970 1969 7Dec.210 BaySt.LouistoGulfportJames 1970 197816Aug.3,500aroundsunkenfishingboat,Purrington1979GulfportGulfofMexico,andintheCaribbean,whileanunknown numberalsobreedonislandsalongthenortherncoastofSouthAmerica.TheLaughingGullisthemostabundantbreedingmarinebirdinthesoutheasternUnitedStates.However,theyarealsodifficulttocensusadequatelyandcensusesofthesoutheasternUnitedStateshavebeenfewandincomplete.Consequently,thetotalbreedingpopulationinthesoutheastcouldbeconsiderablylargerthanissuggestedbyTable11.LaughingGullsarealsoabundantbreedingbirdsalongthenorthernAtlanticcoastoftheUnitedStates.Surveysofthiscoastin1977revealedabout138,400 Lauglrlng GullsnestingfromMainetoVirginia(ca.460inMaine[Korschgen1979],400inMassachusetts,61,880inNewJersey,190inDelaware,4,460inMaryland,and62,390inVirginia[Erwin1979a]).Somepopulationsinthisareaarefarsmallerthantheywereinthelate1800'sandearly1900's.Breedingpopulationsattheirpeakon MuskegetIsland,Massachusetts,numberedover40,000birdsinthelate1930'sandearly1940's,andabouttwiceasmany96

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LaughingGullbredinMaineduringthe1960's(Nisbet1971a)thanatpresent.BreedingpopulationsontheNorthAtlanticcoast(andinthesoutheasternUnitedStates)mayvaryconsiderablyfromyeartoyear.Thecensustotalsforthe1977censusgivenbyErwin(1979a)andKorschgen(1979)maynotbeanadequateestimationofthenumbersusuallynestingintheNorthAtlanticstates.Erwinreportedthatabout61,880LaughingGullsnestedin25coloniesinNewJerseyduring1977,butBuckleyandMcCaffrey(1978)reportedabout70,488in31coloniesduringthesameyear.Anotherunspecifiedrecentcensus(J.GalliinBurger1981b)indicatedthatsome108,000birdsbredin75coloniesinNewJersey.AnundeterminednumberofLaughingGullsalsobreedinthesouthernGulfofMexicoandintheCaribbean.Breedingpopulationstheremaybesubstantial.WinterLaughingGullswinterprimarilyinthesoutheasternUnitedStates,theCaribbean,andalongthecoastsofMexicosouthtoPanamaandthenortherncoastofSouthAmerica.TheymaywinterasfarsouthalongtheAtlanticcoastofSouthAmericaasthemouthoftheAmazon,andalongthePacificcoastsouthtoPeru.Southern's(1980a)analysisof1,521bandingrecoveriesledhimtoconcludethatasignificantproportionwinterinCentralAmericaandnorthwesternSouthAmerica.Wetmore(1965)reportedthatlargeconcentrationswinterintheGulfofPanamaandLaughingGullsevidentlyarecommoninwinterinnorthernSouthAmerica(Murphy1936).Thelargestproportionofbandedbirdsrecoverednorthof19()N camefromFloridaandCubaandneighboringareas,withsmallernumbersreportedfromtheCarolinasandtheGulfcoast(Southern1980a).ThesizesofwinteringpopulationsarepoorlyknownbutreachtheirpeakintheUnitedStatesalongthecoastsofFloridaandTexas(Map2).YoungdispersingfromcoloniesonthewestcoastofFloridaoccurwidelyinwinterthroughoutthepeninsularwestcoastofFloridaandalsomigratetotheCaribbean.Fewfromthewestcoastwinterontheeastcoast(SchreiberandSchreiber1977).MigrationLittledetailedinformationisavailableonthemigrationofLaughingGulls.NorthwardmigrationbeginsinMarch(Southern1980a)andbyAprilandMaylargeflocksmaybeseenstreamingnorthclosetotheAtlanticcoast.MostfallmigrationtakesplacefromlateAugusttoNovember. NumberspeakinPanamainOctoberand November,andagaininApril(Wetmore1965).HABITATNestingHabitatschosenfornestingvaryfromareatoarea.LaughingGullsinthenorthernportionofthebreedingrangetendtonestintidalsaltmarshes;thosefurthersouthnestbothinsaltmarshesandindriersituationsonisolatedman-madeandnaturalislands,aswellasalongbarrierbeaches.Thosenestinginareasundisturbedbyfloodingconsistentlyreturnedtositesusedduringpreviousnestingseasons(BurgerandShisler1980a).MuchhasbeenpublishedonnestingareaschosenbyLaughingGullsintheeasternUnitedStates.Wesummarizesomeofthisinformationbystatebelow.97

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LaughingGullMassachusettsLaughingGullsatMonomoyIslandnestmostlyindensebeachgrassinhollowsbetweendunes,butsometimesunderoraroundbayberry(Myricapensylvanica)bushes.Mostnestsare1-5m(3-16ft)abovehighwater(Nisbet1976b).NewJerseyAtBarnegatBay,NewJersey,about90%oftheLaughingGullsnestinlowmarshdominatedbysaltmarshcordgrass(Spartinaalterniflora)(BurgerandShisler1980a).Manymovedtohighersitescontainingless arterniflora andmoreSpartinapatensduringayearwithunusuallyhightides.SomeuseJuncus,Phragmites,andbushesaswell,butnonenestonlowmarshislandsvegetatedonlywithalterniflora.Montevecchi(1978a)reportedthatnestsitesatBrigantineNWRweregenerallyonornearmatsofSpartinaintallgrassonlowgroundnearwater.Nestsnearpoolsinthemarshusuallywereclosertowater(2.7+0.3m[8.9+1.0ft])thanwerethosenestingneartidalcreeks(4.8+0.2 m [15.7+0.6-ft]),butgullsnestednearcreeksnearlytwiceasoften as nearpools.-Theheightofthegrass(Spartinaalterniflora)aroundnewlybegunnestsaveraged33.8+0.8cm[13.3 0.3in].LaughingGullson RingIsland,atidalmarshnearCape May,NewJersey,prefertonestinthetallestalterniflora(Bongiorno1970).Of156nests,86(55.1%)werebuiltinalterniflora0.8m[2.6ft]highorhigher.Thishabitatrepresentedonly17.9%ofvegetatedarea(1,540sqm[16,576sqft])availablefornesting.Inthistallgrassthenestdensitywas0.3nests/sqm(2.8/100sqft).Thesmallestproportionofnestswerefoundinalternifloralessthan0.4m(1.3ft)inheightandinSpartinapatens0.2-0.4m(0.61.3ft)tall.Intheseareasthenestdensitywas0.02nests/sqm(0.2/100sqft).AlthoughbothBongiorno(1970)andMontevecchi(1978a)foundnestsonwrack(pilesofdebriswashedup onthemarshbyhighorstormtides),noneoccuronsuchsitesatBarnegatBay,presumablybecausewrackisfoundthereonlyatthehightideline(BurgerandShisler1980a).Also,BurgerandShisler(1980a)didnotfindvegetationheightasignificantfactorinnestsiteselectionatBarnegatBay.NorthCarolinaLaughingGullsinNorthCarolinausedredge-spoilislandsasnestsitesalmostasfrequentlyasnaturalsites.In1973,53.3%ofthebirdsnestedonspoiland 46.7%nestedonnaturalislands(SootsandParnell1975a).Comparablefiguresfor1977were65.7% and34.3%,respectively(ParnellandSoots1979ms).Nestssiteswereusuallyonlow,grassyflats(primarilyS.patens),inswalesbetweenmoreelevatedground,andonthelowerslopesofdredge-spoilislands(ParnellandSoots1975).LaughingGullsinNorthCarolinatendtonestinmorevegetativecoverthananyothermarinebirdsstudiedexceptHerringGulls(ParnellandSoots1975,SootsandParnell1975a).Mostnestingcolonies(73%)containedSpartinapatens,andtypicalnestinghabitatwasmoderatelytodenselyvegetated,usuallywithmorethan50%groundcover.Plantsmostfrequentlyassociatedwiththenestsiteswereseasidegoldenrod(Solidagosempervirens),Mexican98

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LaughingGulltea(Chenopodiumambrosioides),andseaox-eye(Borrichiafrutescens)(SootsandParnell1975a).Florida-GulfCoastLaughingGullsnestingondredge-spoilislandsavoidopenareasofsandandgravelandnestamonglowgrassesandbushes(SchreiberandSchreiber1978).Neststendtobeclumpedaroundbusheswhenthesearepresent.Wherebushesareabsent,nestsarewidelyspaced,some2-3m(7-10ft)apart.Nestdensitiesrangefrom0.5/sqm(4.6/100sqft)inseasidepaspalum(Paspalumvaginatum)to0.33-0.5/sqm(3.1-4.6/100sqft)amonggroundsel(Baccharishalimifolia)anddogfennel(Eupatoriumcapillifolium).Wherethereislesscover,nestdensitybecomesaslittleas0.125/sqm(1.2/100sqft)(SchreiberandSchreiber1978).NestdensitiesatacolonyinBocaCiegaBay,southernPinellasCounty,forthethreeyears1975-1977were0.146, 0.165,and0.321/sqm(1.36,1.53,and2.98/100sqft),respectively(datarecalculatedfromSchreiberetal.[1979).LouisianacoastalbeachesandorS.alterniflora;noy-1977).LaughingGullnestsinLouisianaarefoundprimarilyalongsecondarilyinsaltmarshes.Nestsinmarshesareonshellthosealongbarrierbeachesaremostlyin patens(Port-TexasLaughingGullsnestinginTexasarefoundaboutequallyoftenonnaturalanddredgedsites(Chaneyetal.1978).Theyprefertonestnearbushesingrassyareas,buttheyhavealsobeenfoundonwind-blownmatsofSpartinaspp.FeedingLaughingGullsfeedinawidevarietyofhabitatsthatreflecttheirvariedfoodhabits.Theytypicallyforageinmarshes,alongbeachesandmudflats,andinbaysandinlets.Theyfrequentlyfeedinsmallflocksoffshore(Burleigh1958,Kale1979msa).IncoastalMainethesegullspreferredtoforageonmudflats(HuntandHunt1973).ManyoftheLaughingGullsbreedingatMonomoy,Massachusettsfedneartheirnestingareaintide-rips,overmudflatsatfallingtide,andalongtheshores(Nisbet1976b).Smallernumbersforagedfurtheraway,pickingfoodfromthesurfaceoftheseaorfeedingoversandbarsonschoolsofsmallfishoronsmallfishdriventothesurfacebypredatoryfish(Nisbet1976b).LaughingGullsfeedinginmarshesatBrigantineNWR,NewJersey,fedprimarilyduringlowtideand moreoftenduringarisingtidethanduringafallingtide(Burger1976).AlthoughLaughingGullsforagelesson dumpsinMainethanotherspeciesofLarus(HuntandHunt1973),dumpsinotherareashavebecomemajorsourcesoffood(SchreiberandSchreiber1980,Burger1981b).Burger(1981b)pointedoutthatrecentuseofdumpsbyLaughingGullsinNewJerseyhasincreasedconsiderablysince1975.Increasingdailyuseofland-movingequipmenthasmade moregarbageandrefuseavailabletoLaughingGullsforlongerperiods.ItalsoaffordsaerialLaughingGullsmoreopportunitiestoseizefoodbetweenmovingbulldozers,amanueveratwhichtheyaremoreadeptthanareHerringGulls.99

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LaughingGullOntheeasternshoreofMarylandandVirginia(Wolk1959;Clapp,pers.observ.)andinSouthCarolina(SpruntandChamberlain1949),LaughingGullsfrequentlyforageoninsectsandotherinvertebratesturnedupbytractorsandplows.Oberholser(1974)notedthattheyoccasionallyfeedashortdistanceinlandinfields,pastures,and onprairies.LaughingGullsreadilyforagebehindboatsandferriesalongtheAtlanticandTexascoasts,scavengingdiscardedfishremainsfromtheformerandseizingbreadthrownbytouristsfromthelatter.Theyalsoscavengediscardedgarbagealongwharfs(Burleigh1958).NonbreedingandOffshoreHabitatsusedinwinteraresimilartothoseusedduringthebreedingseason.Mostbirdsarefoundalongthecoasts(Murphy1936,Wetmore 1965)butsomearefoundinlandinpastures(Murphy 1936)oronlakesandwaterimpoundments (Wetmore1965).Blake(1977)indicatedthattheprimaryhabitatsinSouthAmericaareseacoasts,islands,andestuaries.LaughingGullsfrequentlyfeedoffshore.Kale(1979mSb)notedthathundredswereseeninNovember 1974feedingoveraschooloffishsome 80mi(130km)westofNaplesontheFloridaGulfcoast.InthesouthernChesapeakeBight,LaughingGullsseenoffshorefromearlyspringtolatefallarefoundmostlyinshoreofthe10-fathomcontour(19km[12miloffshore)(Rowlett1980).Transientsoccasionallyoccurfartheroffshoreandnonbreedingbirdsmayforageasmuchas95km(60mi)offshore.FOODANDFEEDINGBEHAVIORLaughingGullsengageinmuchaerialfeeding.GullsinMaine doabout22%oftheirforagingfromtheair(Hunt and Hunt1973).They hawkinsectsintheair(Forbush1924,Mayr1948),andfromthesurfaceofthewater(Lauro1977),andseizethemfromleaves,twigs,andbranches(Forbush1924).LaughingGullsdivingforfoodalongtheTexascoastusuallyhoverataheightof Less than10ft(3m)and swooptopickfoodfromthesurfaceofthewaterorslightlybelowit(Zusi1962,pers.comm.). Theyoftenlowertheirfeetwhilereachingforfoodandpaddletheminthewater.LaughingGullsseldomsubmergedeeplywhendivingandusuallyleaveatleastthewingsandtailexposed(Zusi1962).Tolonen(1970)notedsmallgroupsofLaughingGulls"ploughing"(i.e.,runningthroughthewaterwiththelowermandibleimmersedandseizinganysmallfishencountered).Healsosawlow-flyingbirdspaddlewiththeirfeetinthewaterwhilekeepingthelowermandibleconstantlyimmersed.LaughingGullsalso"tread"whenforaginginshallowwater(Wood1949).Theystamptheirfeetrapidlyand.alternately,apparentlytoscaresmall food itemStothesurfacewherethesemaybeseized-moreeasily(Wood1949).LaughingGullsalsofeedondeadfishandotheritemsonmudflatsandbeaches,andalightonthewatertoreachdown andseizefood(Zusi1962).Theyoftenfeedonscrapsbehindfishingvessels(Zusi1962)orferries,andreadilyseizebitsoffoodthrowntothem.Adultsandyoung-of-the-yearthatwerethrownpiecesofbread100

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LaughingGullinNorthCarolinaandTexaspreferredtoseizethepiecesinflight(Clapp,pers.observ.).Adultsareconsiderablymoreadeptatseizingfoodthrownintheairthanarejuveniles(Schreiberand Young1974;Clapp,pers.observ.).LaughingGullsalsokleptoparasitizeotherspeciesofbirds,chasingthemandforcingorfrighteningthemintodisgorgingordroppingtheirprey,whichisthenseizedbythegull.Chasingmaybedoneeitherbyindividualsorsmallgroupsofbirds.Asmanyas11gullschasedasingletern(Sternaspp.)atPetitMananIsland,Maine,butmostchasingwas donebygroupsof2-4gulls(Hatch1975).ThesegullsalsokleptoparasitizeBlackSkimmers(Zusi1958,Leck1968),and BrownPelicans(Pelecanusoccidentalis)(Murphy1936,Baldwin1946).Burger(1981b)describedfeedingbehaviorinthefallatadumpinEastBrunswick,NewJersey.LaughingGullshoveredoverthedump,dippeddown, andpickedupfooditemsexposedbybulldozers.Theagedistributionofbirdsusingthedumpwas53%adults,8%subadults(birds15to16monthsold),and39%young-of-the-year.TherateatwhichLaughingGullsfoundfoodvariedinverselywiththenumberof Herring Gullspresentandtherewereapparentlynodifferencesinfeedingsuccessrelatedtoage.Burger(1981b)suggestedthatthelackofdifferenceinfeedingsuccessbetweenagegroupsresultedfromcompetitionwithHerringGulls,i.e.,allagesofLaughingGullswerepreventedfromfeedingwhenconcentrationsofHerringGullskeptthemfromreachingfood.ThedietoftheLaughingGullishighlyvariedbutpoorlyknown.BirdstakeninAlabamainsummer hadfedlargelyoncrustaceansandinsects,whilemostfromtheAtlanticcoasthadeatenfish(Howell1932).ThosetakeninAlabama hadeatencrabsandshrimpand avarietyofinsectsbuthadalsoeatena numberofsmallcatfish(Howell1928).Anexaminationof32stomachsfrombothareasshowedthatabout47%ofthebulkwascrustaceansandabout43%wasfish(Howell1932).LaughingGullsalsooccasionallyeatyoungClapperRails(Ralluslongirostris)(Segreetal.1968),terneggs(Oberholser1938),andsmallpasserines.Daniels(1973)notedadultandimmatureLaughingGullschasingandcapturingsmallpasserines,evidentlyCatharusthrushes,overDelawareBayinearlyOctober.Inanotherinstance,anescapedGalapagosfinch(Camarhynchusparvulus)wasseizedandeatenasitfledashipatthemouthoftheRioGuayas,Ecuador(Wiggins1965).IMPORTANTBIOLOGICALPARAMETERSEggLayingLaughingGullsarriveandnestatnorthernlocalitiesabitlaterthanatsouthernones(Montevecchietal.1979).Peaksoflayingoccurduringthefirsttwo weeksofJuneinMassachusetts,inlateMayinNewJersey,andinthefirsttwo weeksofMayontheGulfcoastofFlorida(authorscitedbyMontevecchietal.1979).Thepeakofegglayingmayvarya weekormorefromyeartoyearinsomeareasbutitusuallyoccurswithina two weekperiodinMassachusetts,NewJerseyandFlorida(Schreiberetal.1979).Montevecchi101

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LaughingGulletale(1979)studiedpatternsofegglayingatBrigantineNWR,NewJersey,andfoundamoderatedegreeofsynchrony.Femalesinthecenterofthecolonyconsistentlylaidearlierthanthoseattheperiphery.Schreiberetale(1979)reportedthatmosteggsinFloridaarelaidbetween0700and1900hrs.MeanClutchSizeLaughingGullslaytwotofiveeggs,butmostnestscontaintwoorthreeeggs.Montevecchi(1978a)reportedthatbirdsinthecentralareasofthecolonyhavea modalclutchsizeofthreeeggs,comparedtotwoeggsforbirdslayingattheperiphery(Table13).At aFloridacolonyclutchsizewas althoughnotsignificantlyso,inareasofthecolonywhereeggswerefirstlaid(2.63vs.2.51foranareainwhichbirdslaidlater)(Schreiberetale1979).Schreiberetalefoundnosignificantdecreaseintheclutchsizeasthebreedingseasonprogressedinthiscolony.IncubationPeriodNisbet(1976b)reportedanaverageincubationperiodofabout29daysforLaughingGulls'nestingatMonomoyIsland,Massachusetts.LaughingGullsatBrigantineNWR,NewJersey,hadanaverageincubationperiodof21-23days(Segreetale1968).MostoftheeggsincubatedontheFloridaGulfcoasthatchedin23-25dayswith56%oftheincubationperiodsover23.5andunder25days(Schreiberetal.1979).Schreiberetalefoundnosignificantdifferenceinincubationperiodinrelationtowheneggswerelaid.Theythoughtthatlargersamplesmightshowsignificantdifferencesinincubationperiodsbetweendifferenteggsoftheclutch,butpointedoutthatthedegreeofvariabilitywassoslightthatitwasprobablyofnobiologicalimportance.HatchingSuccessReportedhatchingsuccessratesvary(Table14).Insomeareas,hatchingsuccessmaydependonhowcloseLaughingGullnestsaretothoseofthepredatoryHerringGull.Burger(1979)reportedthat86%oftheLaughingGulleggshatchedatClamIsland,NewJersey,wheretheLaughingGullswerewellseparatedfromHerringGulls.Wherethetwospeciesnestedincloseproximity,only60%oftheLaughingGulleggshatched.In1976inFlorida,astheseasonprogresBedhatchingsuccessofthree-eggclutchesdeclined,buthatchingsuccessoftwo-eggclutchesincreased(Schreiberetal.1979).AgeatFledgingSchreiberandSchreiber(1980)obtainedrecordsoffledgingperiodsaccuratetowithin1-3daysforyoungatacolonyontheFloridaGulfcoast.Youngfledgedata meanageof42.5days,witharangeof35-50days.Nisbet(1976b)suggestedthatLaughingGullsatMonomoyIsland,Massachusetts,fledgeatabout34days.FledgingSuccessSchreiberandSchreiber(1980)reportedfledgingsuccess(i.e.,percentofeggslaidthatresultedinfledgedyoung)as44%and36%fornestswiththreeand twoeggs,respectively,atonecolonyinFloridain1976.Thesenestsproduced1.32and0.71youngperpair,respectively.Hahn(1981)reportedaproductionof2.13youngperpairfornestswiththreeeggsinNewJersey.MortalityofEggs and YoungLaughingGullnestsandeggsareoftenlosttohighwaterorstorms(Hailman1960b,Bongiorno1968,SchreiberandSchreiber1980),sometimesresultinginthereestablishmentofthecoloniesonhigher102

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LaughingGullTable13.MeanclutchsizesreportedfortheLaughingGull(a).MeanNumberclutchofsizeclutchesLocalityandyearofobservationSource2.512.212.852.762.842.52 2.192.412.152.29941795138 1937016017NewJersey,BrigantineNWR,1972-74NewJersey,BrigantineNWR,1972-74Florida,lowerBocaCiegaBay, 1972Florida,lower Boca CiegaBay,1973Florida,lowerBocaCiegaBay, 1975Florida,lowerBocaCiegaBay,1976Florida,lowerBocaCiegaBay, 1977Texas,islandsinLagunaMadre,1977Texas,GalvestonBay,LittlePelicanIsland,1977Texas,GalvestonIsland,JigsawIslands,1977Montevecchi1978aMontevecchi1978a DinsmoreandSchreiber1974 Dinsmore andSchreiber1974Schreiberetal.1979Schreiberetal.1979Schreiberetal.1979 Chaneyetal.1978 Chaneyetal.1978 Chaneyetal.1978(a)Someofthesefiguresarederivedfromcountsofcontentsofnestsduringshort-termvisits.Theymaynotadequatelyrepresentclutchsizeforthepopulation,becauseaproportionofthepopulationmayhavestillbeenlayingeggs.ThefirstfigurelistedforNewJerseyisforbirdslayinginthecenterofthestudyarea;thesecondisforbirdslayingattheperiphery.ThefigurelistedforislandsintheLaguna Madreisacompositederivedfrom10nestsateachofsevendifferentlocalities.Meanclutchsizeattheselocalitiesrangedfrom1.80to2.80.ground.Montevecchi(1978a)regardedtidalfloodingasthegreatestthreattoLaughingGullnestsatBrigantineNWR,NewJersey.Hereportedthat70-100%oftheneststhereweredestroyedbyfloodsonaverageonceeverytwoyearsoveraten-yearperiod.Lossofeggsoryoungtopredatorsiscommon,butisnotamajorsourceofnestfailure.Predationwasbelievedtohavecausedonly3-10%ofthelossofeggslaidatoneFloridacolony(Schreiberetal.1979),andMontevecchi(1977)reportedthatonly4.4%oftheeggsinaNewJerseycolonywerelosttopredation..HerringGullsareoftenreportedaspredators(Montvecchi1977,Burger1981a),andtakebotheggsandyoung.Crows(CorvusossifragusandC.brachyrhynchos)alsofrequentlytakeeggs.Montevecchi(1977)stated thatlboth adultsandyoungarepreyeduponbyBarnOwls(Tytoalba)andGreatHorned Owls (Bubovirginianus),andsuggested,butdidnotdocument,predationbyShort-eare-d--Owls(Asioflammeus),TurkeyVultures(Cathartesaura),andNorthernHarriers (CircuS-CYaneus). ClapperRails(Rallus 10ngirostriS) alsooccasionallypreyoneggs(Segreetal.1968).103

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LaughingGullTable14.RatesofhatchingsuccessreportedfortheLaughingGull(a).PercentofeggslaidthathatchNumberofeggsLocalityandyearofobservationAllclutchesSource11.151.392.979.281.377.686.892.970.72739 126 265 344 237 22828116Texas,islandinLagunaMadre,1977Texas,GalvestonBay,1977Florida,lowerBocaCiegaBay,1972Florida,lowerBocaCiegaBay,1975Florida,lowerBocaCiegaBay,1976Three-eggclutchesFlorida,lowerBocaCiegaBay,1975Florida,lowerBocaCiegaBay,1976Two-eggclutchesFlorida,lowerBocaCiegaBay,1975Florida,lowerBocaCiegaBay,1976Chaneyetal.1978 Chaneyetal.1978 DinsmoreandSchreiber1974Schreiberetal.1979Schreiberetal.1979Schreiberetal.1979Schreiberetal.1979Schreiberetal.1979Schreiberetal.1979(a)ThedataforFloridain1972arerecalculatedfromthecitedsource.Eggsthatdisappearedataboutthetimehatchingwasexpectedarenotincluded,buttheeggs_lastnotedaspippedorcrackedareconsideredtohavehatchedsuccessfully.Extensivemortalityofyoung(andadult)LaughingGullsinCorpusChristi,Texas,in1978wasattributedtofeedingoninsectsincottonfieldsthathadbeenpoisonedwithparathion,anorganophosphateinsecticide(Whiteetal.1979b).Starvationwas amajorcauseofnestlingfatalityinoneinstance(SchreiberandSchreiber1980).RenestingBirdswhosenestsarefloodedoutearlyinthenestingcyclemayrenestatotherlocalities(BurgerandShisler1980a).Schreiberetal.(1979)observedthat11of134neststhatlostaclutchreceivedreplacementsatacolonyinBocaCiegaBay,Floridain1976.Althoughbirdsatthese11nestswereunmarked,Schreiberetal.believedthatbehavioralcharacteristicsofeightpairsidentifiedthemastheoriginalpairs.Eggsinsevenoftheseeightnestswerereplaced5to30daysafterthefirstsetdisappeared.Replacementclutchesin7of8instancescontainedthesame numberofeggsorlessthandidtheoriginalclutch.104

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LaughingGullAgeatFirstBreedingUnknown.MaximumNaturalLongevityTheoldestknownLaughingGullwas abirdbandedasanimmatureatMuskegetIsland,Massachusetts,thatwasfounddeadthereatanestimatedminimumageof15years,1 month(Clappetal.1982a).WeightDataonweightsofLaughingGullsforthesoutheasternUnitedStatesandelsewherearegiveninTable15.SUSCEPTIBILITYTOOILPOLLUTIONFewreportsexistofLaughingGullsdyingfromthedirecteffectsofoiling.RecordsinthebandingofficeatPatuxentWRC,Laurel,MD,revealedonlyfourLaughingGullswhosedeathwasattributedtooilpollution.ThefourbirdswerefromNewJersey,Panama,Texas,andLouisiana.Presumably,theLaughingGulls'aerialandterrestrialfeedingbehaviormake themlessvulnerabletooilspillsthanotherdivingandmoremarinespecies.BecausetheyareoneofthemostabundantbreedingseabirdsoftheAtlanticandsoutheasterncoasts,occasionaloilspillsarenotlikelytohavemucheffectonthepopulationsofthisspecies.Secondaryeffectsofoffshoreoildevelopmentandoilpollution(e.g.disturbance,lossoffeedingandbreedingareas)posea muchgreaterhazard.Long-term,low-leveloilpollutionintheenvironmentmighthaveadverseaffectsonthisspecies.Severalpapershaveexploredtheeffectofexperimentalcontamination.Whiteetal.(1979a)examinedsomeeffectsofoilonLaughingGulleggs.TheytreatedeggsatanislandinMatagordaBay,Texas,with20microlitersofNo. 2fueloilandrecordedembryonicmortalityafterfivedaysofnaturalincubation.Eighty-threepercentof58treatedeggsweredead,asopposedto 2% of56controleggs,asignificantdifference.Laboratorystudiesofartificiallyincubatedeggsrevealedalowerhatchabilityintreatedeggs(2of51ata20microliterdosagehatched,asopposedto26of51controleggs).King andLeFever(1979)examinedtheeffectofoiltransferredfromincubatingLaughingGullstotheireggs.Forty-twogullson SundownIsland,inMatagordaBay,werecapturedand2.5mlofNo.2fueloilwereappliedtotheirbreastfeathers.Significantembryonicmortalityoccurredintheeggsthesegullswereincubating,with41%mortalityinthetreatedgroupversus2%forthe20controls.Thesestudiesshowthatlow-leveloilingcancausesignificantreproductivelossesinLaughingGulls.Studiesareneededontheprevalenceandimpactofoilinginbreedingbirds.105

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LaughingGullTable15.Weights(ingrams)ofLaughingGulls(a).MeanNumberweightRangeweighedSample andseasonAreaSourceFledgedbirds182-36016adultsFloridaDinsmore andSchreiber1974 249150-345125adultmalesFloridaSchreiberandSchreiber1979 348282-39213adultmalesTexasZusi1962,pers.corom.29419malesFloridaHartman 1955 204137-30026males,1yearFloridaSchreiberandSchreiber1979 224150-300224adultfemalesFloridaSchreiberandSchreiber1979 306 8femalesFloridaHartman 1955 247female,MayMexicoLSU256female,Dec. MexicoLSU220 1female,Sep.Ohio Trautman1956178140-27222females,1yearFloridaSchreiberandSchreiber1979 3182ndwinter Hexico LSUfemale,Dec.Chicks30936-day-oldsFloridaSchreiberandSchreiber198028.823.1-34.117newlyhatchedNewJerseyRicklefsetal.197830-334newlyhatchedFloridaDinsmoreandSchreiber197425-35newlyhatchedFloridaSchreiberandSchreiber1980Alleggsofclutch(a)42.735.3-50.127NewJerseyRicklefsetal.197838.130.4-43.39FloridaSchreiberand Lawrence 1976 106

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Table15.Concluded.LaughingGullMeanweightRangeNumberweighed Sample andseasonAreaSourceAlleggsofclutch(continued)42.834.2-60.6716 1975FloridaSchreiberelal.197940.028.5-57.61061 1976FloridaSchreiberetal.1979Firsteggofclutch44.839.4-50.29NewJerseyRicklefseta1.197843.436.1-48.260 1975FloridaSchreiberetal.197942.536.0-47.5771976FloridaSchreiberetal.197943.338.6-47.0111975FloridaSchreiberetal.197941.534.5-47.5601976FloridaSchreiberetal.1979 Second eggofclutch42.836.0-49.69NewJerseyRicklefseta1.197843.234.6-50.4591975FloridaSchreiberetal.197940.933.0-48.0781976FloridaSchreiberetal.197942.935.7-48.7121975FloridaSchreiberetal.197938.728.5-47.5601976FloridaSchreibereta1.1979Thirdeggofclutch40.632.6-48.69NewJerseyRicklefseta1.197839.234.2-48.360 1975FloridaSchreiberetal.197936.330.5-41.8781976FloridaSchreiberetal.1979(a)Alleggweightsareforfreshornearfresheggs.Figuresforrangefrom Rickelefs etal.(1978)arethemean2 S.D.ThesecondsetoffiguresforthefirsteggandsecondeggsofaclutchfromSchreiberetal.(1979)areforsecondeggsoftwo-eggclutches.107

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LaughingGull BIBLIOGRAPHY 1982Reid,M.andC.S.Hacker.1982.SpatialandtemporalvariationinleadandcadmiumintheLaughingGull,Larusatricilla.Mar.Pollut.Bull.13:387-389.vandenBerg,A.B.1982.LaughingGullwithflame-scarletbillandlegs.DutchBirding4:55.1981Andrews,R.1981.ThegullpopulationonMonomoyIsland,Massachusetts,USA.(Abstractonly.)ColonialWaterbirds4:194.Burger,J.1981b.FeedingcompetitionbetweenLaughingGullsandHerringGullsatasanitarylandfill.Condor83:328-335.Burger,J.andM.Gochfeld.1981a.Age-relateddifferencesinpiracybehaviouroffourspeciesofgulls,Larus.Behaviour77:242-267.Erwin,R.M.,J.GalliandJ.Burger.useinAtlanticcoastseabirds.1981.ColonysitedynamicsandhabitatAuk98:550-561.Gochfeld,M.andJ.Burger.1981.Age-relateddifferencesinpiracYoffrigatebirdsfromLaughingGulls.Condor83:79-82.Hahn,D.C.1981.AsynchronoushatchingintheLaughingGull:cuttinglossesandreducingrivalry.Anim.Behav.29:421-427.Muse, C.J.andS.Muse. 1981. First gullrecordforSamoa.Elepaio41:130131.Telfer,T. C. andJ.K.Shisler.1981.RecordmovementofaLaughingGulltoHawaiifromNewJersey.J. Field Ornithol.52:340-341.Tessen,D.1981.ALaughingGullinManitowocCounty.PassengerPigeon43:63.1980Brock,K.J.1980.LaughingGullatMichiganCityHarbor.IndianaAudubonQ.58:19.Burger,J.,M.Fitch,G.ShugartandW.Werther.1980.PiracyinLarusgullsatadumpinNewJersey.Proc.1979Conf.ColonialWaterbirdGroup3:87-98.108

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LaughingGullNunnally,S.,D.NunnallY,R. Needham andR.Lennon.1979.Nocturnalfeedingofgullsatalightedpier.Chat43:63.Schreiber,E.A.,R.W.SchreiberandJ.J.Dinsmore.1979.BreedingbiologyofLaughingGullsinFlorida.PartI:Nesting,egg,andincubationparameters.Bird-Banding50:304-321.Schreiber,R.W.and E.A.Schreiber.1979.Notesonmeasurements,mortality,molt,andgonadconditioninFloridawestcoastLaughingGulls.Fla.FieldNat.7:19-23.White,D.H.,K.A. King andN.C. Coon.1979a.EffectsofNo.2fueloilonhatchabilityofmarineandestuarinebirdeggs.Bull.Environ.Contam.Toxicol.21:7-10.White,D.H.,K.A.King,C.A.Mitchell,E.F.Hilland T.G.Lamont.1979b.ParathioncausessecondarypoisoninginaLaughingGullbreedingcolony.Bull.Environ.Contam.Toxicol.23:281-284.1978Buckley,F.G.,M.GochfeldandP.A.Buckley.1978.BreedingLaughingGullsreturntoLongIsland.Kingbird28:203-207.Burger,J.1978.DeterminantsofnestrepairinLaughingGulls.Anim.Behav.26:856-861.Burger,J.andJ.Shisler.1978a.NestsiteselectionandcompetitiveinteractionsofHerringandLaughingGullsinNewJersey.Auk95:252-266.Condamin,M.1978.Nidificationsd'oiseauxdemerenGuyane.OiseauRev.Fr.Ornithol.48:115-121.[InFrenchwithEnglishsummary.]Green,J.C.1978.FirstadultLaughingGullforMinnesota.Loon50:167-168.Montevecchi,W.A.1978a.NestsiteselectionanditssurvivalvalueamongLaughingGulls.Behav.Ecol.Sociobiol.4:143-161.1978b.Corvidsusingobjectstodisplacegullsfromnests.Condor80: ------349. Ricklefs,R.E.,D.C.Hahn andW.A.Montevecchi.1978.TherelationshipbetweeneggsizeandchicksizeintheLaughingGullandJapaneseQuail.Auk95:135-144.Snodderly,D. M.,Jr.1978.EggshellremovalbytheLaughingGull(Larusatricilla):normativedataandvisualpreferencebehaviour.Anim.Behav.26:487506.110

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LaughingGullSchreiber,R.W.andS.N.Young.1974.EvidenceforlearningtofeedinLaughingGulls.Fla.FieldNat.2:16-17.1973Bohlen,H.D.1973.FirstspecimenofLaughingGullfromIllinois.WilsonBull.85:233.Daniels,G.1973.LaughingGullspreyonmigrants.Linn.Newsl.27:1.Hunt,G.S.,Jr.andM.W.Hunt.1973.HabitatpartitioningbyforaginggullsinMaine andnorthwesternEurope.Auk90:827-839.Impekoven,M.1973.TheresponseofincubatingLaughingGulls(LarusatricilIaL.)tocallsofhatchingchicks.Behaviour46:94-113-.--------Lincer,J.L.andD.Salkind.1973.Apreliminarynoteonorganochlorineresiduesintheeggsof fisr-eating birdsofthewestcoastofFlorida.Fla.FieldNat.1:3-6.1972Bierly,M.L.1972.LaughingGullinMauryCounty.Migrant43:93-94.Dawson,W. R., J.W.Hudson andR.W.Hill.1972.TemperatureregulationinnewlyhatchedLaughingGulls(Larusatricilla).Condor74:177-184.Forsythe,D.M.1972.LaughingGullbandrecoveriesfromSouthCarolina. Bird-Bandlng 43:264-266.Tucker,V.A.1972.MetabolismduringflightintheLaughingGull(Larusatricilla).Am.J.Physiol.222:237-245.Wallace,D.I.M.1972.LaughingGullintheIslesofScilly.Brit.Birds65:79-81.1971Hailman,J.P.1971.Theroleofstimulus-orientationinelicitingthebeggingresponsefromnewly-hatchedchicksoftheLaughingGull(LarusatricilIa).Anim.Behav.19:328-335.Impekoven,M.1971.PrenatalexperienceofparentalcallsandpeckingintheLaughingGull(LarusatricillaL.).Anim.Behav.19:475-480.Morton,A.and C.Nicholson.1971.LaughingGullinLondonCounty.Migrant42:89.Nisbet,I.C.T.1971a.TheLaughingGullinthenortheast.Am.Birds25:677-683.113

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LaughingGullNisbet,I.C.T.1971b.LaughingGullcoloniesinthenortheast.Mass.Audubon55:2-10.Stewart,P.A.1971.RejectionofTobacco Hornworm moths byLaughingGulls.Chat35:53.1970Beer,C. G.1970a.OntheresponsesofLaughingGullchicks(Larusatricilla)tothecallsofadults.I.Recognitionofthevoicesoftheparents.Anim.Behav.18:652-660.1970b.OntheresponsesofLaughingGullchicks(Larusatricilla)to-----thecallsofadults.II.Agechangesandresponsestodifferenttypesofcall.Anim.Behav.18:661-677.Bongiorno,S.F.atricilla) 1970.Nest-siteselectionbyadultLaughingGulls(LarusAnim.Behav.18:434-444.Hatch,J.J.1970.PredationandpiracybygullsataterneryinMaine.Auk87:244-254.Tolonen,K.E.1970. GullandLaughingGullcatchfishby"ploughing"and"skimming".WilsonBull.82:222-223.1969Beer,C. G.1969.LaughingGullchicks:recognitionoftheirparents'voices.Science166:1030-1032.Brown,N.R.andP.A.Pearce.SparrowandLaughingGull.1969.NewBrunswickspecimensoftheFieldCan.Field-Nat.83:403.1968Bongiorno,S.F.1968.EggpuncturingbehaviorinLaughingGulls.Auk85:697-699.desForges,G.1968.AnoldrecordofLaughingGullinSussex.Brit.Birds61:213-214.Hailman,J.P.1968a.Spectralreflectanceofgull'sbill:physiologicalandevolutionaryimplicationsforanimalcommunication.Science162:139-140.1968b.Visual-cliffresponsesofnewly-hatchedchicksoftheLaughing-----GullLarusatricilla.Ibis110:197-200.Leek,C.F.1968.AnadditionalobservationofLaughingGullsrobbingaBlackSkimmer.Cassinia50:27.114

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LaughingGullMarshall, R. V.A.1968.LaughingGullinEssex.Brit.Birds61:415-416.Segre,A.,J.p.Hailman and C.G.Beer.1968.ComplexinteractionsbetweenClapper Rails andLaughingGulls.WilsonBull.80:213-219.Tuck,L.M.1968.LaughingGull(Larusatricilla)andBlackSkimmers(Rynchopsnigra)broughttoNewfoundlandbyhurricane.Bird-Banding39-:-200-20-8-. -Weston,F.M.1968.LaughingGullnestingnearPensacola[Florida].Fla.Nat.41:35.1967Albrektsson,T.andP.Lindberg.1967.Field-charactersoftheimmatureLaughingGull.Brit.Birds60:159-160.Alsop,F.J.1967.LaughingGullinKnoxCounty.Migrant38:61-62.Bongiorno,S.F.1967.SubstrateandthedistributionofLaughingGull,Larusatricilla,nests.Ph.D.thesis,RutgersUniv./NewBrunswick,NJ.130pp.Buck,W.F.A. andD.W.Taylor.1967.LaughingGullinKent:aspeciesnewtoBritainandIreland.Brit.Birds60:157-159.Frohling, R. C.1967.ApartialalbinoLaughingGull.Bird-Banding38:235-236.Grant,P.J.1967.TheEuropeanrecordsofLaughingGullsin1964-66.Brit.Birds60:489-490.Hailman,J.P.1967.Theontogenyofaninstinct.ThepeckingresponseinchicksoftheLaughingGull(LarusatricillaL.)andrelatedspecies.BehaviourSuppl.15.vii Harriman,A.E.1967.LaughingGullsofferedsalineinpreferenceandsurvivaltests.Physiol.Zool.40:273-279.vonSchmidt,K.1967.(Larusatricilla).Note onthebreedingplumageoftheLaughingGullFla.Nat.40:103.1966Frohling,R.C.1966.AsocialflightoftheLaughingGull.Bird-Banding37:206-207.Hailman,J.P.1966a.FourcolorpreferencesoftheLaughingGull(Larusatricilla).(Abstractonly.)Am.Zool.6:288.115

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LaughingGullHailman,J.P.1966b.Mirror-imagecolor-preferencesforbackgroundandstimulus-objectinthegUllchick(Larusatricilla).Experientia22:257-258.1965Albrektsson,T.andR.Berndtsson.1965.Amerikanskskrattmas (Laru atricilIa)observeradiGoteborgsfiskhamn.[AmericanLaughingGull(Larus atrtCilla) observedatGothenburg.]VarFagelvarld24:289-293. SwedishwithEnglishsummary.]Schmerler,S.andJ.P.Hailman.1965.LaughingGull,LarusatricillaL.DiscriminationoforientationintheAm.Zool.5:655.Wiggins,I.L.1965.GalapagosFinchcapturedinflightbyLaughingGull.Condor67:82.1964Hailman,J.P.1964a.Theontogenyofaninstinct:thepeckingresponseinchicksoftheLaughingGull(LarusatricillaL.)andrelatedspecies.Ph.D.thesis,DukeUniv./Durham,NC.1964b.Codingofthecolourpreferenceofthegullchick.Nature204:------710.Medley,M.1964.ObservationsonLaughingGullsandFranklin'sGullsatSt.Joseph,Michigan.Jack-PineWarbler42:231.1963Nicholson,D.J.1963.ALaughingGull(Larusatricilla)seeninOrangeCounty.Fla.Nat.36:126-127.Penner,L.R.and B.Fried.trematodefrombirds.1963.Philopthalmusregenerisp.n.,anocularJ.Parasitol.49:974-977.Voous,K.H.1963.CarribeanSea.Terncolonies-inAruba,Curacao,andBonaire,SouthProc.Internatl.Ornithol.Congr.13:1214-1216.1962Hailman,J.P.1962a.LaughingGullcolonyonGullIslandinPamlicoSound.Chat26:19.1962b.PeckingofLaughingGullchicksatmodelsoftheparentalhead.------Auk79:89-98.116

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LaughingGull1961Hailman,J.P.1961.AgeofLaughingGullchicksindicatedbytarsallength.Bird-Banding32:223-226.1960Hailman,J.P.1960b.DestructionofPeaIslandLaughingGullcolonyin1960.Chat24:98.1960c.BandedPeaIslandLaughingGullrecoveredinCuba.Chat ------24: 24.1959 Walk,R.G.1959a.LaughingGullsfollowingtheplow.WilsonBull.71:387388.1958Moynihan,M.1958.NotesonthebehaviorofsomeNorthAmericangulls.II.Non-aerialhostilebehaviorofadults.Behaviour12:95-182.Zusi,R.L.1958.LaughingGulltakesfishfromBlackSkimmer. Condor 60:67-68.1957Preston,F.W.1957.Pigmentationofeggs:variationintheclutchsequence.Auk74:28-41.1956Gunter,G.1956.Onthereluctanceofgullstoflyunderobjects.Auk73:131-132.1953Nicholson,D.J.1953.FirstrecordoftheLaughingGullforOrangeCounty,Florida.Fla.Nat.26:136.Preston,F.W.and E.J.Preston.1953.Variationoftheshapesofbirds'eggswithintheclutch.Ann.CarnegieMus. 33:129-139.1952Parkes,K.C.1952.TaxonomicnotesontheLaughingGull.Proc.BioI.Soc.Wash. 65:193-196.117

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LaughingGull1951Gross,A.o.1951.LaughingGullsnestingatStrattonIsland.Bull.Maine AudubonSoc.7:52-54.Small,A.1951.LaughingGullonthecoastofsouthernCalifornia.Condor 53:101.1949Hawksley,O.andJ.Hawksley.1949.LaughingGullsnestingoffthecoastofMaine.Bull.Maine AudubonSoc.5:77.Wood,H.B.1949.LaughingGullstreadouttheirfood.Bird-Banding20:103.1948 Mayr,E.1948.Gullsfeedingonflyingants.Auk65: 600. 1946Baldwin,W.P.1946.LaughingGullrobsBrownPelican.Auk63:96-97.1945Gross,A.O.1945.TheLaughingGullonthecoastofMaine.Bird-Banding16:53-58.1943Noble,G.K.andM.Wurm.1943. ThesocialbehavioroftheLaughingGull.Ann. N.Y. Acad.Sci.45:179-220.1940Hickey,J.J.1940.CourtshipnoteontheLaughingGull.Auk57:111-112.Noble,G.K.and D.S.Lehrman. 1940.EggrecognitionbytheLaughingGull.Auk57:22-43.Noble,G.K.andM.Wurm.1940.TheeffectofhormonesonthebreedingoftheLaughingGull.Anat.Rec.78(Suppl.):50-51.Snyder,L.L.1940LaughingGullbreedinginNovaScotia.Auk57:568-569.1938Buckalow,H.[sic].1938. FoodofyoungLaughingGulls.Auk55:672.118

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LaughingGull1936Murray,J.J.1936.LaughingGullintheNorthCarolinamountains.Auk53:81-82.1935Sprunt,A.,Jr.1935.SecondrecordednestingoftheLaughingGullontheSouthCarolinacoast.Auk52:308-309.1934Lincoln,F. 1934. BandedLaughingGullrecoveredinElSalvador.Condor36:36-37.Weston,F.M.1934.AmelanisticLaughingGullatPensacola,Florida.Auk51:82-83.1933Burton,E.M.and E. B.Chamberlain.1933.LaughingGullbreedingontheSouthCarolinacoast.Auk50:360.1932Allen,R.P.1932.Eggsof TerninLaughing;Gull'snest.Auk49:219-220.1929Gudger, E. w.1929.HowtheLaughingGull,thefeatheredBuccaneerofFloridawaters,getsitsdinner.Sci.Mon.29:435-439.Miller,L.andA.J.vanRossem.1929.NestingoftheLaughingGullinsouthernCalifornia.Condor31:141-142.1928Phillips,C.L.1928.EarlyarrivaloftheLaughingGullinMassachusetts.Auk45:366-367.1925Hix,G.E.1925.AbundanceoftheLaughingGull(Larusatricilla)aboutNewYorkCity.Auk42:125.1924Forbush,E.H.1924.GullsandternsfeedingontheSeventeen-yearCicada.Auk41:468-470.119

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LaughingGullStone,W.1924.Franklin'sGullatPhiladelphia--acorrection.Auk41:468.1921Alexander,E.G.1921.LaughingGull(Larusatricilla)capturedbysnappingturtle.Auk38:596.Pearson,T.G.1921.Notes onthebird-lifeofsoutheasternTexas.Auk38:513-523.1916Williams,J.1916.LaughingGull(Larusatricilla)nestingnearSt.Marks,Fla.WilsonBull.28:36.1907 Brownson,W.H.1907.VisittoacolonyofLaughingGulls.J.MaineOrnithol.Soc.9:57-66.1898Mackay,G.H.1898.TheternsofMuskegetIsland,Massachusetts.PartIV.Auk15:168-172.1897 Mackay,G.H.1897.TheternsofMuskegetIsland,Massachusetts.PartIII.Auk14:383-390.1896Mackay,G.H.1896.TheternsofMuskegetIsland,Massachusetts.PartII.Auk13:47-55.1895MacKay,G.H.1895.TheternsofMuskegetIsland,Massachusetts.Auk12:32-48,178.1893 Mackay,G.H.1893.ObservationsonthebreedinghabitsofLarusatricillainMassachusetts.Auk10:333-336.120

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FRANKLIN'SGULL(Laruspipixcan)[DA:Praeriemage;FR:MouettedeFranklin,SP:ApipizcadeFranklin,GaviotadeFranklin;SW:Prariemas,Franklinmas]GENERALDISTRIBUTIONNorthAmericaFranklin'sGullsbreedonlyininteriorNorthAmerica.InCanada,theybreedintheeasternhalfofAlberta,centralandsouthernSaskatchewan,andsouthwesternManitoba(Godfrey1966).IntheUnitedStates,theybreedfromtheCanadianbordersouthtocentral-easternOregon,northwesternUtah,easternNorthDakota,northeasternSouthDakota,southwesternMinnesota,andnorthwesternIowa(AOU1957).Franklin'sGullswinterinsmallnumbersonthecoastsofLouisianaandTexas,andfromGuatemalasouthtotheGulfofPanama(AOU1957).ThemainwinteringgroundsareonthePacificcoastofSouthAmericasouthtoArauca,Chile(Blake1977).MostmigrantsfollowanarrowroutesouththroughtheGreatPlains,Texas,andeasternMexico(DuMont1941,Blake1977),andthencesouthward.Franklin'sGullsarepronouncedwanderers.Totheeastandsoutheast,theyhavestraggledtoNewBrunswick(Godfrey1966);VirginiaandMaryland(Burford1963,Wierenga1976);sixtimestoEngland(BilletandGrant1971,Rogers1972,Blick1979,Brown1979,Harrison1979,Rogersetal.1981,1982);twicetoFrance(Beaudoin1979,Kerautret1981);twicetoSweden(JonssonandWennberg1981);toNorway(Rogersetal.1981)andofftheFaroeIslands(Grandjean1981);toSt.BartholomewandPuertoRicointheWestIndies(Bond1971);andtotheAtlanticcoastofSouthAfrica(CooperandWilliams1975).Tothesouthandwest,Franklin'sGullshavestrayedtotheGalapagosIslands(AOU1957);theMarquesasIslands(Dupont1976);Hawaii(Berger1972);JohnstonAtollandtheLineIslandsintheCentralPacific(Clapp1968,AmersonandShelton1976);theMarshallIslands(Anderson1978);andsouthwesternAustralia(ServentyandWhittell1976).One wasalsoseenrecentlyatMarionIslandintheIndianOcean(Sinclair1981).IntheNewWorldFranklin'sGullshavestrayedasfarnorthasCookInlet,Alaska(GibsonandMacDonald1971),andasfarsouthastheStraitsofMagellan(PetersonandWatson1971).WorldDistributionSeeabove.DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESFranklin'sGullshavebeenrecordedfromallthesoutheasternstates,buttheirstatustherevariesfrombeingaveryrarevagrantinthenortheasttoanabundantmigrantinthesouthwest.121

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Franklin'sGullNorthCarolinaFranklin'sGullsareaccidentalinNorthCarolina(Wray andDavis1959).TheonlyrecordforthestateisofafemalecollectedinlandontheCatawbaRivernearCharlotteon 12October1952(Chamberlain1953).SouthCarolinaFranklin'sGullisacasualvagrantinSouthCarolinathatisknownonlyfromfoursightrecords.Tworecordsarefrominland.Onewasseenabout1 miSEofTownville8May1975 (LeGrand1975)andanotherwasseenatLakeGreenwood 2April1978(Teulings1978a,Lewis1979).AnotherwasseenalongthecoastatHuntingtonBeachStateParkon 26September1976(LewisandLewis1977,Teulings1977a)and oneinsummer plumage wasseenatBarnwellIslandnearthemouthoftheSavannahRiveron 18 March 1978(Smith1978).GeorgiaTwosightrecordsofFranklin'sGullshavebeenmadeinlandatColumbus,butadequatedocumentationoftheiroccurrenceinthestateisstilllacking.Thefirst,anadult,wasseen24April1965(WellsandWells1965);thesecond,afirst-yearbirdwasseen20May1966(Parnell1966b).ThebirdseennearBarnwellIslandontheSouthCarolina-GeorgiaborderalsooccurredinGeorgia(Smith1978).FloridaFranklin'sGullsarerarevisitorstoFlorida,wheretheyhavebeenrecordedtwiceasfrequentlyontheGulfcoastasontheAtlantic.MostrecordsfromstatesnorthofFloridaareapparentlyofstragglingmigrants,whereasmostFloridarecordsareapparentlyofwinteringbirds.AllbutoneoftheAtlanticcoastrecordsarebetween6 November and 8December,andallbuttwooftheGulfrecordsarebetween24Octoberand26February.Florida-AtlanticCoast1938 4 Dec.foundinlandatLakeport,LakeOkee-McClanahan 1941chobee(bandedinManitoba)195617Nov.remainsfoundnearPrinceton,MiamiStevenson1957aMiami 1956 18 Nov. ,3,1seennearPrinceton,MiamiStevenson1957a,8Dec.1957b 1961 15 Mar.seeninlandatLakeFlorencenearStevenson1961 Cocoa 197511Nov.imm.seen,photogr.atNewSmyrnaBeachEdscorn1976 1976Dec.seen,photogr.atCocoaStevenson1977 1977 6 Nov.(1?)seenoffCanaveralEdscorn1978 1977 19 Nov.(1?)seenatMayportEdscorn1978 122

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Franklin'sGullFlorida GulfCoast1918 26Feb.1955 12June1July195710-11Dec.1957-28Dec.,58 28Jan.19589-10Nov. 1958 12 Nov. 1958 30 Nov. 1960 4Dec.1961 24Feb.1963Dec.197727Nov.1977-10Dec.78 6Jan.1978 20Apr.5(l?)2seenatSt.Petersburgad.colI.inlandatLakeJackson,Leon Co.seennearFranjoseennearFlamingoimm.seenatFairpoint,3miSPensacolaseen(4imm., 1ad.)1 miSWPace,SantaRosa Co. imm.seen3 mi WPace,SantaRosa Co.seennearFranjoseenatNaplesseenatSt.MarksseenatAlligatorPointseenatSt.petersburg,Toy townDumpad.seenatSt.Petersburg,Toy townDumpPangburn1919Stevenson1959aStevenson1958bStevenson1958b Monroe1959,Newman1959 Monroe 1959 Monroe 1959Stevenson1961Stevenson1961Stevenson1964aEdscorn1978Stevenson1978Kale1978a197824OcL17Nov.,etseq.1,2seenatToy townDump,PinellasCo.Edscorn1979 1978 28Oct.seenatFt.DesotoPark,PinellasCo. 1979to12Jan.3(?)seeninPinellasCo.Edscorn1979Stevenson1979aAlabama Imhof(1976b)consideredFranklin'sGullararemigrantinAlabama.Hepointedoutthattheymayoccurregularlyinfall,buthavebeenoverlookedbecauseofdifficultiesindistinguishingthisspeciesfromtheLaughingGull.AtpresentthereareonlyninerecordsfromAlabama.Mostoftheseareofwinteringbirds,andonlytwoarefromcoastalareas.ThefrequencywithwhichFranklin'sGullshavebeenreportedrecentlyfromtheGulfcoastofFlorida,inareasconsiderablyfartherfromthemainmigratorypathwaysthanAlabama,123

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Franklin'sGullsuggeststhatImhofiscorrectandthatFranklin'sGullsprobablyaremore commoninAlabamathanthefewsightingsindicate. 1958 Junead.seeninlandatDecaturImhof 1976b19604Jan.ad.seeninlandatDecaturImhof 1976b197025Oct.2ad.seenatCochraneCauseway,MobileImhof 1976b 1972 4Julyad.seeninlandatHorseshoreBendDam,Imhof1976b,FranklinCo.Stewart1972 1972 14Dec.ad.seeninlandatBirminghamImhof1976b 1972 26Dec.3ad.seeninlandatDecaturImhof1976b 197724-25Apr.ad.seeninlandatDecaturImhof19771977-winterad.seen[nolocalitygiven)Hamilton1978 78 1979 28Jan.ad.seenatDauphinIslandHamilton1979MississippiThestatusofFranklin'sGullinMississippiisinadequatelyknown.Wehavefoundonlyninerecords,includingabirdreportedbyBurleigh(1944)asaBonaparte'sGullthatisapparentlythefirstspecimenofFranklin'sGulltakeninMississippi.ItseemsclearthatFranklin'sGullsarelargelyuncommonwinterresidentsinthestate.1938 25 Nov.1960earlyNov. 1961 24Dec.1968 6Jan.1972 27Dec.87femalecolI.(USNM#432219)atGulfportthispaperjuv.seenatGulfportWilliamsandClawson1963juv.femalecolI.atGulfport, GandyandTurcotte1970seeninlandatLegionLake,BolivarCo.James1968ad.seeninlandatSardisLakePurrington1973b1976 31Dec.1979 23Oct.198010Nov.198111May1221seeninlandonHattiesburgCBCimm.seeninlandatHattiesburgad.seenonmudflatatBaySt.Louisad.seenatPascagoulaRiverMarsh 124Gates1977JacksonandSchardien1980Hodges andToups,1981Jackson1981

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Franklin'sGullLouisianaOberholser(1938)statedthatFranklin'sGullwereararewinterresidentonthecoastandlistedonlyeightrecordsforthestate.Lowery(1974)indicatedthatthisgulloccursregularlyinsmallnumbersinsouthwesternLouisiana.Theyaremuchlesscommoninthesoutheasternpartofthestate,eventhoughthelargestcountrecordedinthiscentury(175birds)wasseenoverLakeponchartrainon 25October1959(Lowery1974).AsinMississippi,Franklin'sGullsareprobablymorecommoninLouisianathanispresentlythoughtbecauseofthedifficultyindistinguishingthisspeciesfromotherblack-headedgulls.TexasFranklin'sGullsarecommontoabundantmigrantsincoastalTexas.TheyoccurinfallmigrationfromlateSeptembertoearlyDecember,andinspringmigrationfromearlyApriltoearlyJune(Oberholser1974,Blacklock1978ms).PeaknumbersarepresentinOctoberandNovemberandinApril(Blacklock 1975 ms,Table16).Smallnumberswintercasuallyalongthecoast(Oberholser1974).SYNOPSISOFPRESENTDISTRIBUTIONANDABUNDANCEBreedingFranklin'sGullsbreedsonlyintheNearctic,wheretheyareconfinedtofreshwatermarshesofcentralNorthAmericainanareabetweenabout43and54Nlatitudeand90and110Wlongitude(Southern1980a).MigrationInthesoutheasternUnitedStates,Franklin'sGulloccursmainlyasamigrantbutoccasionallyasawintervisitor.TheyareraretoaccidentalalongtheAtlanticcoastandraretouncommonintheeasternGulf(Table16),buttheirstatusinthenorthernGulf(Alabama,Mississippi,Louisiana)isill-definedandthespeciesmay bemorecommontherethaniscurrentlythought.Migrationoccurswithinabriefspanoftime,primarilyinlateOctoberandNovemberandfrommid-AprilthroughearlyMay(Tables17,18).ThespeciesfollowsanarrowmigrationpathfromitsnorthernbreedingareasthattakesitalongtheTexascoast,whereenormousconcentrationshavebeenseen.WinterSmallnumberswinterintheGulfofMexicoandonthePacificcoastofCentralAmerica.MostofthepopulationwintersalongthePacificcoastofSouthAmerica.HABITATNestingFranklin'sGullsnestcoloniallyinmarshesorsloughs,ofteninshallowwaterwithsemi-openemergentcover(R.Stewart1975).Bent(1921),Guay(1968),andBurger(1974)agreedthatFranklin'sGullsprefertonestnearopenwaterandthattheearliestnestsoftheseasonarealongtheouteredgeofthereeds.Thenestsusuallyareplacedonaclumpofemergentvegetationthatwaseitherbentoverorbrokenfromthepreviousyear(Guay1968).125

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Franklin'sGullTable16.ApproximatenumberofFranklin'sGullsrecordedbymonthfortheareafromNorthCarolinatoMississippi(a).MonthsState/regionJANFEBMARAPRMAYJUNJULSEPOCTNOVDECNorthCarolinaSouthCarolina11Georgia11Florida-AtlanticCoast7 3Subtotal-ATLANTICCOAST2 2 2 1 7 3Florida-GulfCoast62 1 1 2 10 6Alabama2 1 1 2 1 4Mississippi81 123 9Subtotal-GULFCOAST16 2 2122 5 34 19TOTAL-ALLAREAS16 2 2 4 3 2 2 64122(a)Birdsfounddeadinthefirst10daysofamontharearbitarilyassignedtotheprecedingmonth.Ifthesourcedidnotmakeitclearwhetheroneormore wasseen,weassumedonlyonewasseen.Ifthesourceindicatedmorethanonewasseenbutdidnotspecifiyhow many(e.g."several","afew"),weassumedtwowereseen.Birdsseeninmorethanonemonthwerecountedseparatelyineachmonth.Birdsrecordedintwostatesarelistedunderthestateinwhichfirstseen.Recordswithunspecifieddates(e.g.,"winter","spring")wereomitted.Guay(1968)consideredthemostimportantfactorinthechoiceofasuccessfulnestsitetobethepresenceofappropriateemergentvegetation.IntheAlbertamarshwherehestudiedthisgull,theimportantplantsarebulrush(Scirpusacutus)orcattail(Typhalatifolia).Inotherareasotherspeciesofplantscomprisethe importanc-emergent vegetation.R.Stewart(1975)notedthattheimportantspeciesinNorthDakotaarebulrush,alkalaibulrush(Scir paludosus[=maritimus]),whitetop(Scolochloafestucacea),andphragmites(Phragmitescommunis[=australis]).At acolonyatDavisLake,Montana,thepredominantemergentvegetationisalkalaibulrush,withsomesoftstembulrush(Scirpusvalidus)andcattail(Typhalatifolia)(Rothwiler1960).Waterdepthinthemarshrangedfromato6in(0-15cm).Burger(1974)concludedthatbothdensityanddispersalofemergentplantsareimportantinthechoiceofanestsite.IncentralAlberta,Guay(1968)foundlittlevariationinnestdensitybetweenstudyplots.Overalldensitywas 1nestper80.7sqft(1/7.5sqm),butmoredesirablenestingareashadgreaternestdensity(1/68.6sqft[=1/6.4sq126

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Franklin'sGullTable17.DatesofoccurrenceforFranklin'sGullsinthecoastalsoutheasternUnitedStates(a).StateNumberofoccurrencesDatesofoccurrence1 3 2 8ca.159512Oct.26Sep. 2Apr.24Apr.20May6 Nov. -15Mar. 24Oct. 20Apr.(12June-1July)25Oct.-25Apr.(1June-4July)23Oct 11May25Sep.-17MaylateSep. earlyJune(29June mid-July)??-scatteredreportsmanyNorthCarolinaSouthCarolinaGeorgiaFlorida-AtlanticCoastFlorida-GulfCoastAlabamaMississippiLouisianaTexas(a)Particularlyunusualdatesarelistedinparentheses.Table18.PeakconcentrationsofmigrantFranklin'sGullsincoastalTexas(a).NumberDateseenseenLocalitySource195813Apr.500+AustinWebster1958b 197214Apr.1,000BartlettWebster1972c 1979 17Apr.1,800moving NoverAustinWebster1979c195921Apr.200 SanAntonioWebster1959 1962 25Apr.4,550AustinWebster1962b 195826Apr.827+AustinWebster1958b 196029Apr.1,640+AustinWebster1960b 1961 30Apr.6,000+overLakeTravis,15miWebster1961b WofAustin1960 1May2,170+AustinWebster1960b 195211Oct. 15,000.:!:. overCove Goldman and Watson1953a1952 14Oct.1,000feedingnearHockleyGoldman and Watson1953a1955 29Oct.20,000+CoveWebster1956a1951 30Oct. 20,000+" overElCampoGoldman and Watson 1952 1951 1 Nov. 50,000+" overRockportGoldman and Watson 1952 1959 2 Nov. 1O,000I roostingon LakeTravisWebster1960a 1967 10 Nov.1,200+NeucesareaWebster1968a196411Nov.20,000+betweenFreeportandAngletonWebster1965 1967 12 Nov. 7,500.:!:.LosFresnos,NofBrownsvilleWebster1968a(a)Recordsarearrangedchronologicallybytimeofyear.127

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Franklin'sGullm]).Nestdensitywaslower(1/100.5sqft[=1/9.3sqm])inlessdesirablenestingareas.Burger(1974)obtainedsimilarresultsinMinnesota.Shefoundanaverageof12.6nestsper6sqmsampleplotin areas nexttoopenwater,anda meanof4.0forplotsentirelywithinstandsofcattail.Eveninopenareasthedensityofnestswasinverselyproportionaltothedensityofcattailstems(butthepresenceofsomecattailswasnecessary).Coloniesvarygreatlyinsize.Theyusuallyconsistofafewhundredpairsbutmaycontainasmanyas50,000pairs(McNicholl1971).Franklin'sGullsevidentlyarelessfaithfultotheirnestsitesthanmanyotherspeciesofgulls.From 1969through1971,onlyoneofsevencolonieswasusedcontinuouslyatAgassizNationalWildlifeRefugeinMinnesota(Burger1974).FeedingFranklin'sGullsforageinmarshesandfields.BreedingbirdsinAlbertafedbothoverthenestingareaandoveradjacentlandundercultivation.MostbirdsbreedingatacolonyinnorthwesternMinnesotafedwithin16km(10mi)ofthecolony,butonewasseenasfarawayas39km(24mi)(Burger1974).BirdsbreedingatSandLakeRefuge,SouthDakota,mainlyfedinfieldstoatleast36 mi(58km)fromthecolony(DuMont1941).NonbreedingandOffshoreWinteringbirdsinSouthAmericaareabundantalongthecoast,onsandybeachesandshores,inharbors,andincultivatedfields(Murphy1936).FOODANDFEEDINGBEHAVIORFranklin'sGullsobtainmostoftheirfoodbyaerialpursuitofflyinginsects(Guay1968).Whenoverwater,theyfeedonfishbysurface-dipping(Murphy1936)andfrequentlysurface-seize(Burger1974),i.e.,swimonthesurfaceofthewaterpickingpreyatorjustbelowthesurface.Franklin'sGullsarealsonotedforthepropensitywithwhichtheyfollowploughs.Franklin'sGullsfeedonawidevarietyoffooditemsbutaremainlyinsectivorous,particularlyduringthebreedingseason.Thelittleinformationavailableformigrantandwinteringbirdssuggeststhatthisgullfeedsmoreopportunisticallyduringthoseperiods.McAteeandBeal(1912)examined93stomachs,almostallfrombirdsonthebreedinggrounds.Animalfoodmade up 94.5%ofthedietbyvolume;vegetablefoodmadeup5.5%.Themostimportantfoodwasgrasshoppers(43.4%).Mostoftherestofthefoodwereinsects(beetles,crickets,dragonflynymphs,ants,Maybeetles).Theonlyplantfoodofanysignificancewaswheat,whichmorethanhalf-filledtwostomachs.Cottam(1944)reportedthreeFranklin'sGullsfromManitobaandNorthDakotawithstomachscontainingsignificantamountsofoatorwheatkernels.MostoftheremaininginformationonthedietofFranklin'sGullisfrombreedingareasandissummarizedbelow.128

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Franklin'sGullAlbertaGuay(1968)examined27stomachscollectedinJuneatHaysLake.Byvolume,insectspredominated(88%).Overall,Coleoptera(46%),Diptera(12%),andHemiptera(7%)werethemostimportant,butwhatwasmostimportantatanyonetimevariedseasonally.DuringMay,dytiscidbeetlesweremostimportant,followedbyCorixidaeandmidges(Chironomidae).DuringmostofJune,Corixidaeweremostimportant,withCarabidaeandScarabeidaeeatenmorelaterinthemonth.InJuly,ScarabeidaewerefollowedinimportancebyCarabidaeandthenLepidoptera.MontanaRothweiler(1960)reportedthefrequencyofoccurrenceoffoodbyfamilyfrom 108stomachscollectedonthebreedinggroundsatFreezeoutLake.Plants(mainlywheatandbarley)werefoundinenoughstomachssothathebelievedthesefoodswerenottakenincidentally.Hisdatashow aseasonalchangeindiet.InAprilandMay, 88.9%ofthestomachscontainedplants,27.8%containedamphibians,andinsectswerecommon(Scarabeidae72.2%;Carabidae-44.4%;Chironomidae38.9%;Dytiscidae-27.8%;Muscidae-27.8%).InJuneandJuly,plantswerefoundin92.6%ofthestomachs,andamphibiansin14.8%.AstrongshiftindietoccurredinAugustandSeptember.plantswerestillfoundin75.0%ofthestomachs,butamphibians(Ambystomatigrinum)inonly1.6%.Theinsectseatenchangedgreatly,withgrasshoppers(Acrididae)occurringin87.5%ofthestomachs;beetleswerenextinimportance(Carabidae-64.1%;Scarabeidae-37.5%).NorthDakotaAughey(1878)examined10stomachscollectedinMayfrom1868-1877.Thesecontainedlocusts,frogs,fish,snails,crayfish,andalizard.Saugstad(1940)reportedonanother10stomachs,ninefromadultsandjuvenilesinJuly,andonefromanOctoberadult.Theformercontainedmostlygrasshoppers(Melanoplusmexicanus,M.bivittatus,M.differentialis),aswellassomebeetlesandplantbugs;thelatterlargelycontainedbeetles(Carabidae).MinnesotaRoberts(1900)madegeneralcommentsbasedonobservationsin1892-1893andin1899.InJune,adultsandyoungfedlargelyondragonflynymphs,butlaterfedextensivelyonearthwormsandcockchafergrubs;grasshopperswerefavoredattheendofthebreedingseason.Burger(1974)foundevidenceofconsiderableseasonalchangeinfoodhabits.InearlyMay,birdsatemanyearthwormsandtookbarleyandoatsaswell.InlateMayandlateJuneadultsfedmainlyonmidges(Chironomidae).InearlyJune,samplesfromyoungbirdsmostlycontainedearthworms.Migrantsapparentlyarealsolargelyinsectivorous.StomachsfrombirdscollectedinLouisianainfallcontainedgrasshoppers,truebugs,andbeetles(McAteeandBeal1912).AFranklin'sGullcollectedinfallinKansashadeatenstinkbugs(Pentatomidae),anadultprairievole(Microtusochrogaster),andanadultwesternharvestmouse(Reithrodontomysmegalotis)(EasterlaandDamman 1977). BirdsseeninSouthCarolinainearlyMayandinFloridainmid-Novemberwerehawkinginsects(Monroe1959,LeGrand1975).Winteringbirdsevidentlyfeedlargelyonfishandcrustaceans.ThestomachsofsixofeightbirdscollectedinJanuaryatPacasmayo,Peru,allshowedtracesoffish(Murphy1936).Oneheldabout300smallcrustaceans,andthe129

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Franklin'sGullremainsofamolluscwerefoundinanother.IMPORTANTBIOLOGICALPARAMETERSEggLayingMostnestingtakesplaceinthelatterhalfofMay,althougha fewnestsmaybestartedinearlyMayandinJune(Rothweiler1960,Guay1968,Burger1974).Theperiodofegglayinginanygivenyearisshortcomparedtoothergulls(Burger1974),andmaylastno morethan21days(Guay1968,Burger1974).MeanClutchSizeBurger(1974)statedthatFranklin'sGullsnormallylay3-eggclutches,althougha fewnestscontain4eggs.At Hays LakeincentralAlberta,therangewas1-3eggs;themodal numberofeggswas two(in56%of208nests)(Guay1968).Guayreportedmeanclutchsizesof2.16,2.09,and2.39for58,117,and 33nestsin1964,1965,and1966,respectively.The meanclutchsizefor42nestsatSanFranciscoLakeinsouthernAlbertain1964 was2.45;threeeggswerefoundinhalfthenests(WolfordinGuay1968).The mean numberofeggslaidatHays Lake was morethanthatfoundbyinspectingnestsbecauseexaminationof68reproductivetractsshowedthat90%ofthefemaleshadlaidthreeeggs(Guay1968).Iftherestofbirdsexaminedlaidtwoeggs,thiswouldresultina meanclutch size of2.90.IncubationPeriodGuay(1968)gavearangeof22-28daysforincubationofthefirsteggofaclutch(mean=24.6,n=11);21-26daysforthesecondegg(mean=24.4,n=38);and24-27daysforthethirdegg(mean=25.0,n=9)incentralAlberta.The meanincubationperiodforall80eggsmonitoredwas24.6days.InanotherstudyatSanFranciscoLakeinsouthernAlberta,Wolford(inGuay 1968)reporteda meanincubationperiodof23.5daysfor11clutcheswitharangeof21-25days.HatchingSuccessIncentralAlbertain1964,75.7%ofalleggsoranaverageof1.69eggspernesthatched;in1965,60.4%hatchedforanaverageof1.27pernest(Guay1968).AgeatFledgingInAlbertain1964,ageatfirstflightrangedfrom23to33days,witha meanof30days.In1965,itrangedfrom24to31days,witha meanof28days(Guay1968).FledgingSuccessTheaveragenumberofbirdsfledgedpernestwas0.34in1964incentralAlberta.In1965,theaveragewasonly0.16(Guay1968).MortalityofEggs and YoungOf418eggslostin1964and1965atcoloniesinAlberta,66%weredestroyed,2%disappeared,26%wereabandoned,1%droppedintothewater,and5%diedduringhatching(Guay1968).DestructionwascausedprimarilywhenneighboringFranklin'sGullspeckedatorateeggsinunattendednests.Consideringthelargeproportionofabandonednests,onesuspectsthatthemajorfactorinegglossduringthisstudywasdisturbancebytheinvestigator,ahazardalsonotedbyBurger(1974).Guay(1968)stated,130

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Franklin'sGullhowever,thatareaswhichhedidnotdisturbalso"showedreducedclutchsizesandevidencesofpeckedandeateneggs.Competitiveinteractionwithotherspeciesmayalsocauseeggloss.Burger(1974)observedanAmericanCoot(Fulicaamericana)aggressivelydisplaceanestinggullfromitseggsandtheneattheeggs.Inanotherpartofthecolonyinanareaaroundacoot'snest,shefound19gullnestswithpeckedeggs.Othersourcesofnestlossincludefloodingandpredation.InoneMinnesotacolony,highwaterpullednestsfromtheirsites(Roberts1900);inanothercolony,asnappingturtle(Chelydraserpentina)killedanadultandatetheeggs(Preston1886).InAlbertain1964-65lossofyoung wasattributedprimarilytoattacksbyneighboringadults,andtoexposuretoheavyrains(Guay1968).Guaysuggestedthatthislossresultedinlargepartbydisturbancecausedbytheobserver.Avianpredatorsalsoreducefledgingsuccess.InMinnesota,Burger(1974)sawNorthernHarriers(Circuscyaneus)attackingadultsandchicks,andsawonecarryingadeadadult.ShealsoreportedthataGreatHornedOwl(Bubovirginianus)killed12adultsandfournearly-grownjuvenilesina days.MammalianpredatorsmaynotbeassignificantafactorinnestlossinFranklin'sGullsasinothercolonialwaterbirds,possiblybecausethisspeciestendstonestoutneartheedgeofthewater(Burger1974).However,aquaticmammalscanbeasourceofpredation.Burger(1974)wastoldthatmink(Mustelavison)hadcausedsomedestructionofaFranklin'sGullcolonyinNorthDakota,butfoundvisibleevidenceofpredationbythisspeciesinonlyoneyearofher3-yearstudyinMinnesota.InearlyJuly1971,shefoundover45deadanddyingfledglingsonnestplatformsandinthewaterthatsheattributedtopredationbymink.RenestingTheextenttowhichFranklin'sGullsreplacelostnestsisalmostunknown.OvarianinspectionofgullsatacolonyinAlbertasuggestedthatatleastthreefemaleshadreplacedclutcheslostearlier(Guay1968).AgeatFirstBreedingMostFranklin'sGullsprobablydonotbreeduntiltheyareatleasttwoyearsold.Examinationoftheovariesofbirdsintheimmatureplumagecharacteristicofone-year-oldsshowedthattheyoccasionallylayeggs(Guay1968),buttheproportionthatdosoissmall.MaximumNaturalLongevityAFranklin'sGullbandedasajuvenileinMontanahadreacheda minimumageof9yearsand5monthswhenrecovered(Clappetal.1982a).WeightsWefoundfewpublishedweightsofFranklin'sGulls(Table19).Theweightofadults,excludingbirdsthatareevidentlyunder-oroverweightvariesbetween210-340g(7.4-12.0oz).Guay(1968)foundnosignificantdifferenceintheweightsof100adultmalesand69adultfemalesfromcentralAlberta.131

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Franklin'sGullTable19.Weights(ingrams)ofFranklin'sGulls(a).MeanWeightRange NumberofbirdsSample andSeasonAreaSource157ad.male,MayPacificOcean,USNMJohnstonAtoll240280.8220-335268.7325 4521294male,Junebreedingad.malesmale,Augustad,Oct.ad.males,Oct.IndianaMinnesotaMinnesotaMissouriMinnesotaMumford 1962Burger1974MarshallandErickson1945EasterlaandDamman1977Burger1974201.2191-212173.5171-176278.6250-3252211ad.females,MayMexicoLSUad.females,MayPacificOcean,USNMPalmyraAtollbreedingad.femalesMinnesotaBurger1974208.4243 318209.5375 263 26313-323240-28620-352479ad.female,Oct.imm.male,Oct.imm.males,Nov.irom.female,Dec.juveniles,Oct.fledgingyounghatchingchicksHondurasMexicoMichiganMexicoMinnesotaAlbertaAlbertaLSULSUFisheretal.1966LSUBurger1974 Guay 1968 Guay 1968(a)RangefiguresforfledgingyounginAlbertaarethemean+2S.D.132

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Franklin'sGullSUSCEPTIBILITYTOOILPOLLUTIONFranklin'sGullsseldomdiveintowaterbutreadilysitonthesurface.Foodobtainedinflightmaybeeatenwhilethebirdrestsonthewater(Wierenga1976).Thesegullsaremainlyinsectivorous,however,andareaslikelytobeseeninterrestrialsituationsasinaquaticones.ConsequentlytheyarelesssusceptibletooilpollutionthanothermarinebirdsoccurringinthesoutheasternUnitedStates.Throughoutmuchofthesoutheast,theFranklin'sGullisrareoraccidental,buttheyareseasonallyabundantinTexas.Thosemonthswhenitpassesthroughthestateingreatnumbers(April.lateOctober,andNovember)aretheonlytimeswhen asignificantnumbermightbelikelytobeadverselyaffectedbyoilspills.BIBLIOGRAPHY1981Grandjean,P.1981.PraeriemageLaruspipixcanvedFaeroerne1976.Dan.Ornithol.Foren.Tidsskr.75:146-147.[InDanishwithEnglishsummary.]Grant,P.J.1981.Mysteryphotographs.55.Brit.Birds74:298-299.Hodges,M.F.andJ.A.Toups.1981.AflockofFranklin'sGullsinBaySt.Louis,Mississippi.MississippiKite11:17.Jonsson,P.E. andO.Wennberg.1981.PrariemasenLaruspipixcanpatraffadiSverige.[RecordsofFranklin'sGullLaruspipixcaninSweden.]VarFagelvarld40:263-269.[InSwedishwithEnglishsummary.]Kerautret,L.Calais.1981.UneMouettedeFranklin(Laruspipixcan)dansIePas-deOiseauRev.Fr.Ornithol.51:337338.Littlefield,C.D.andS.P.Thompson.1981.HistoryandstatusoftheFranklin'sGullonMalheurNationalWildlifeRefuge,Oregon.GreatBasinNat.41:440-444.Weseloh,D.V.[C.]inginAlberta.1981.AprobableFranklin'sXRing-billedGullpairnestCan.Field-Nat.95:474-476.Weseloh,D.V.C.andM.T.Myres.1981.bersatCalgary,Alberta,CanadaandColonialWaterbirds4:132-137.1980Seasonalfluctuationsingullnum a modelforestimatingtheirnumbers.Allaire,P.N.1980.Noteworthyspecies(includingFranklin'sGull)inBellCounty.KentuckyWarbler56:18-20.133

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Franklin'sGullSchulenberg,T.S.1980.AFranklin'sGull(Laruspipixcan)insoutheasternPeru.Gerfaut70:403-404.Southern,W.E.1980a.ComparativedistributionandorientationofNorthAmericangulls.Pp.449-498inJ.Burger,B.C.OllaandH.E.Winn(eds.).BehaviorofMarine AnimalS7 Vol.4:MarineBirds.PlenumPress,NY.xviiand 515pp.1979Beaudoin,J.-C.1979.UneMouettedeFranklinMaine-et-Loire:premieredonneefrancaise.45-49.[InFrenchwithEnglishsummary.](Laruspipixcan)aAngers,OiseauRev.Fr.Ornithol.49:Blick,M.A. 1979.Franklin'sGullinCleveland.Brit.Birds72:478-479.Brown, B.J.1979.Franklin'sGullinSuffolk.Brit.Birds72:479-482.Grant,P.J.1979.FieldidentificationofwestPalearcticgulls.PartII.Common,Mediterranean,Ring-billed,Laughing,andFranklin'sGulls.Brit.Birds72:142-182.Harrison,G.R. 1979.Franklin'sGullinNorfolk.Brit.Birds72:476-478.Lewis,B.1979.Franklin'sGullinGreenwoodCounty,S.C.Chat43:63-64.Wennberg,o.1979.Franklinmas(Laruspipixcan)forforstagangenantraffadiSverige.Bubo 1978:25-26.1978Abbott,J.M.andD.F.Abbott.1978.Franklin'sGullinAlexandria,Virginia.Raven49:67-68.Anderson,D.A.1978.AFranklinsGullfromtheMarshallIslands.Micronesica14:361-362.Hochbaum,G.andG.Ball.1978.AnaggressiveencounterbetweenaPintailwithabroodand aFranklinGull.WilsonBull.90:455.Nickerson,A.W.achusetts.1978.SightingofaFranklin'sGullinVineyardSound,MassBirdObs.East.Mass.6:222-223.Ryder,R.A.ments.1978.GullsinColorado:theirdistribution,status,and moveProc.1977Conf.ColonialWaterbirdGroup1:3-9.Smith,T.C.1978.Franklin'sGullnearSavannahRivermouth.Oriole43:54.134

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Franklin'sGull1977Bahl,A.K.and B.S.Pomeroy.1977.ExperimentalexposureofFranklin'sGulls(Laruspipixcan)andMallards(Anasplatyrhynchos)toaturkeyinfluenzaAvirusA/Turkey/Minn/BF/72(Hav 6Neq2).J. WildL Dis.13:420-426.Burger,J.1977b.RoleofvisibilityinnestingbehaviourofLarusgulls.J.CompoPhysiol.Psychol.91:1347-1348.Carrolan,T.1977a.Franklin'sGullontheSt.LawrenceRiver.Kingbird27:86-87.1977b.Franklin'sGullontheSt.LawrenceRiver.Kingbird27:142.Easterla,D.A.andD.L.Damman.1977.UnusualfooditemofFranklin'sGull.Auk94:163.Giezentanner,K.I.and W. H.Clark.1977.FirstrecordofFranklin'sGull(Laruspipixcan)inNevada.Southwest.Nat.22:141-142.Hughes, R.A.1977.Franklin'sGulls(Laruspipixcan)atLakeTiticaca,Peru.Biotropica9:52.Lewis,B. andA.Lewis.1977.Franklin'sGullatHuntingtonBeachStatePark,S.C.Chat 41:96-97.1976McColl,J.G.andJ.Burger.1976.Chemicalinputsby acolonyofFranklinGullsnestingincattails.Am.MidI.Nat.96:270-281.Wierenga,H.L. 1976.Maryland'sfirstFranklin'sGulls.Md.Birdlife32:113-114.1975Cooper,J.and A.J.Williams.1975.Franklin'sGullinSouthAfrica.Ostrich46:117.LeGrand,H.E.,Jr.1975.Franklin'sGullinSouthCarolina.Chat 39:92.1974Burger,J.1974.BreedingadaptationsofFranklin'sGull(Laruspipixcan)toamarshhabitat.Anim. Behav. 22:521-567.Houston,C.S.1974.SouthAmericanrecoveriesofFranklin'sGullsandSwainson'sHawksbandedinSaskatchewan.BlueJay32:156-157.135

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Franklin'sGullTaboada,P.,D.A.,S.,E.Zarate.,R.,M.T.Valderrama.1974.DeterminaciondealgunoshelmintosparasitosdoLaruspipixcanWagler,'GavotadeFranklin'.Rev.Peru.BioI.1:194-196.1973Bry,E. 1973. TheFranklin'sGull.N.Dak.Outdoors36:2-3.Burger,J.1973a.CompetitionbetweenAmericanCootsandFranklin'sGullsfornestsitesandeggpredationbythecoots.WilsonBull.85:449-451.1973b.EggsizeandshellthicknessintheFranklin'sGull.Auk90:423-426.Campbell,R.W.andR.G.Foottit.1973. TheFranklin'sGullinBritishColumbia.Syesis5:137-167.1972Burger,J.1972a.BreedingadaptationsofFranklin'sGulls(Laruspipixcan)toamarshhabitat.Ph.D.thesis,Univ.Minnesota/Minneapolis,MN.1972b.Dispersalandpost-fledgingsurvivalofFranklin'sGulls.Bird-Banding43:267-275.1972c.Theuseofafish-eyelenstostudynestplacementinFranklin'sGulls.Ecology53:362-364.Cowan,P.J.1972.Thecontrastandcolorationofsea-birds:anexperimentalapproach.Ibis114:390-393.Moos,L.M.1972.GullbandinginMontana.Proc.Mont. Acad.Sci.32:20-23.Rogers,M.J.1972.Franklin'sGullinSussex.Brit.Birds65:81-82.1971Billett,D.F.andP.J.Grant.newtoBritainandIreland.1971.Franklin'sGullinHampshire:aspeciesBrit.Birds64:310-313.Burger,J.1971. A methodformarsh-trappingbreedingFranklin'sGulls.Bird-Banding42:123-124.Peterson,R.T.andG.E. Watson.1971.Franklin'sGullandBridledTerninsouthernChile.Auk88:670-671.1968 Guay,J.W.1968. ThebreedingbiologyofFranklin'sGull(Laruspipixcan).Ph.D.thesis,Univ.Alberta/Edmonton,AB.119pp.136

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Franklin'sGullSipe,J.P.1968.RarebirdscollectedatLakeSaintMarys,Ohio.OhioJ.Sci.68:334.1967Schultz,Z.M.1967.SightrecordsofFranklin'sandSabine'sGullinSeattle,Washington.Murrelet48:19.1966Fisher,D.,J.Hubbard andP.DeBenedictis.1966.FirstMichiganspecimensoftheLittleGullandFranklin'sGull.Jack-PineWarbler44:50.1965 McCaskie,G.and E.A.Cardiff.1965.NotesonthedistributionoftheParasiticJaegerand some membersoftheLaridaeinCalifornia.Condor67:542-544.Swales,M.K. and R.C.Murphy.1965.AspecimenofLaruspipixcanfromTristandaCunha.Ibis107:394.Wells,L. A. and R.Wells.1965.Franklin'sGullatColumbus.Oriole30:9394.1964 Medley,M.1964.ObservationsonLaughingGullsand GullsatSt.Joseph,Michigan.Jack-PineWarbler42:231.1963Burford,F.C.1963.Franklin'sGullatNorfolk,Virginia.Raven34:52.1962Fisher,H.I.1962.ThehatchingmuscleinFranklin'sGull.WilsonBull.74:166-172.Williams,L.E.,Jr.1962.White-frontedGoose andFranklinGullinMississippi.Miss.Ornithol.Soc.Newsl.7:8.1960Rothweiler,R.A.1960.Foodhabits,movements andnestingofgullson awaterfowlarea,FreezoutLake,TetonCounty,Montana. M.S.thesis,MontanaSt.Coll./Bozeman,MT.29pp.1959King,J.E.1959.FranklinGullinthecentralPacific.Condor61:226.137

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Franklin'sGullMonroe,B.,Jr.1959.Franklin'sGull.Fla.Nat.32:95-96.Moynihan,M.1959.NotesonthebehaviorofsomeNorthAmericangulls.IV. Theontogenyofhostilebehavioranddisplaypatterns.Behaviour14:214-239.Stevenson,H.M.1959d.Franklin'sGull(Laruspipixcan).Fla.Nat.32:146147.1957Collias,E.C. andN.E.Collias.Gulltoparentalbill-color.1957.TheresponseofchicksoftheFranklin'sAuk74:371-375.1956 Moynihan,M.1956.NotesonthebehaviorofsomeNorthAmericangulls.I.Aerialhostilebehavior.Behaviour10:126-178.1954Sooter,C.A.1954.Franklin'sGullsridingwhirlwindandfeeding.Condor56:313.1953Pearse,T.1953.Franklin'sGullonPacificcoastof-BritishColumbia. Condor55:219.1949Jewett,S.G.1949.TheFranklinGullinOregon.Condor51:189-190.1944Cottam,C.1944.Gullsasvegetarians.Condor46:127-128.1943Slipp,J.W.1943.FurthernotesontheFranklin'sGullinthePacificNorthwest.Condor45:38-39.1942Slipp,J.W.1942a.FranklinGullinthestateofWashington:afirstrecord.Murrelet23:18.1942b.FranklinGullinIdaho.Condor44:226-227.138

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Franklin'sGull1941 DuMont,P.A.Plains.1941.RelationofFranklin'sGulltoagricultureontheGreatTrans.N.Am.Wildl.Conf.5:183-189.Low,J.B.1941.Gadwall andFranklin'sGullnestinginIowa. IowaBirdLife11:31-32.McClanahan,R.C.1941.Franklin'sGullanadditiontotheFloridalist.Auk58:405-406.1940 Meade,G.M.1940.Franklin'sGullinNewYorkState.Auk57:251.Saugstad,S.1940.FeedinghabitsoftheFranklin'sGullinNorthDakota.N.Dak.Agric.Exp.Stn.Bimon.Bull.2:14-16.1939 McLean,D.D.1939.EuropeanJackSnipeandFranklinGullinCalifornia.Condor 41:164.Olsen,O.1939.TheoccurrenceoftheflukePlagiorchispotaniniSkrjabin1928inFranklin'sGull(LaruspipixcanWagl.)inNorthAmerica.Proc.HelmintholeSoc.Wash. 6:20.1938 DuMont,P.A. 1938.Franklin'sGullsnestingonSand LakeRefuge,SouthDakota.Oologist55:3-5.1936 Lowe,W.P.1936.Franklin'sGullLaruspipixcan(olimfranklini)inDevon.Ibis(13thSer.)6:378-379.1934 Bradshaw,F.1934.Grasshoppersroutedbygulls.Can.Field-Nat.48:68-69.1932Leopold,A.1932.AflightofFranklin'sGullsinnorthwesternIowa.WilsonBull.44:116.1928Brodkorb,P.1928.Franklin'sGullinIllinois.Auk45:91.139

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Franklin'sGull1927Gardner,L.L.1927. Therelationofbirdstoanoutbreakoflocusts.BirdLore 29:180-182.Lewis,W.E.1927.Franklin'sGullsinnorthwesternOklahoma.WilsonBull.39:109-110.Rowan,W.1927.BandingFranklin'sGullsinAlberta.WilsonBull.39:44-49.1926Bailey,A.H.and R.J.Niedrach.1926. TheFranklinGullinColorado.Condor 28:44-45.Hall,J.W.1926.Franklin'sGullsasinsectdestroyersintheNorthPlatteValleyofNebraska.WilsonBull.38:36-37.1924Stone,W.1924.Franklin'sGullatPhiladelphia--acorrection.Auk41: 468. 1921Morse,A.P.1921.Franklin'sGullinNewEngland.Bull.EssexCo.Ornithol.Club 1921:68-69.1915'Law,J.E.1915.FranklinGull:a newrecordforCalifornia.Condor 17: 96. 1912Conover,F.L.1912.Franklin'sGullinWisconsin.Auk29: 388.Stone,W.1912.Franklin'sGull(Larusfranklini)atPhildelphia.Auk29:99-100.1910Job,H.K.1910. TheFranklin'sGull.Bird-Lore12:124.1902 Smyth, E. A. 1902.Franklin'sGullintheVirginiaMountains.Auk19:74-75.1900Roberts,T.S.1900.AnaccountofthenestinghabitsofFranklin'sRosyGull(Larusfranklinii),asobservedatHeron LakeinsoutheasternMinnesota.Auk17: 272 283. 140

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Franklin'sGull1886Preston,J.W.Oolog. 11: 1886.BreedingofFranklin'sGullinMinneaota.54-55.1885Orn!thol.Willard,S.W.1885.Theoccurrence of Chroicocephalusfranklin!1nWisconsin.Auk2:222.1878 Aughey, S. 1878.NotesonthenatureofthefoodofthebirdsofNebraska. Appendix II.Pp.13-621n U.S. Entomol.Comm.Annu. Rep. 1877.GPO, D.C.AdultBonaparte'sGullinwinterplumage.PhotographbyClaytonTaylor.141

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BONAPARTE'SGULL(Larusphiladelphia)[DU:KleineKokmeeuw,NW:KanadiskHettemake]GENERALDISTRIBUTIONNorthAmericaBonaparte'sGullsbreedinlandinwesternandcentralAlaskaandinCanadafromtheMacKenzieDelta,NorthwestTerritories,andcentralYukonsouthtotheFraserRiverValley,BritishColumbia,andeastacrossforestedareasofAlberta,Saskatchewan,Manitoba,andOntariotoJames Bay (Godfrey1966).ThesegullsmigratesouthalongthePacificcoastandthroughtheGreatLakes(AOU1957)southalongtheAtlanticcoast.Inthewesttheywinterfrom WashingtontosouthernBajaCalifornia.IntheeasttheywintercommonlyalongthesoutheasternAtlanticcoastfromsoutheasternVirgina(VirginiaSocietyofOrnithology1979)southtoFlorida(AOU1957) andtheGreaterAntilles(Bond1974),aswellasalongtheGulfofMexico fromFloridathroughTexas(Map3)totheYucatan(AOU1957).TheywinterinsmallnumberscasuallyasfarnorthassouthernOntarioandNewEnglandintheeast,andsouthernBritishColumbiainthewest(AOU1957).WorldDistributionBonaparte'sGullsbreedonly in theNearcticbuthavestraggledbotheastandwest.Tothewest,thisgullhasbeenreportedfrom Hawaii(Berger1972).Totheeast,ithasbeenrecordedfromGreenland,France,Heligoland(AOU1957),theNetherlands(Kist1961),and Norway (Ree1974).ThespeciesusuallyisreportedonceortwiceayearinBritainandIreland(Grant1978).TheyhavealsobeenreportedfromthenorthernLesserAntilles(Bond 1971)andPanama(Pujals1973).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESBonaparte'sGullisaspeciesthatisseldommentionedintheseasonalreportsappearinginAmericanBirds.Thus,mostofourregionalinformationcame fromstatebirdbooks andmaybeoutofdate.NorthCarolinaBonaparte'sGullsarecommonmigrantsandwinteringbirdsontheNorthCarolinacoast.Potteretal.(1980)regardedthisspeciesaserraticalongthecoastsoftheCarolinas:usuallyfairlycommontocommon,butsometimesabundant.BirdshavebeenrecordedinNorthCarolinafrom asearlyas8AugusttoaslateasMay(Wray andDavis1959).PeakconcentrationsofwinteringbirdsareshowninTable20.SouthCarolinaBonaparte'sGullarelocallycommonwinterresidentsinSouthCarolina(SpruntandChamberlain1949),usuallypresentfromOctober142

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linterDistri ..MapfirSOItlieastn UnitedStatesBIRDSPER'0PARTY-HOURS c:=::J lessthanone _'-5 5-20 1"&1 More than20 (Adapted "0"' 1,ltrca, 1974)INDIVIDUALSOBSERVEDDURINGCHRISTMASBIRDCOUNTS,1973-1977(ARITHMETICMEAN) @ Number of individuals8 less than oneindividual None observed TEGULFOFMEXICOMap3BIRONAME' 30 28

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Bonaparte'sGullthroughMay.Occurrenceson 20 August and 9Junewereconsideredparticularlyearlyandlate(SpruntandChamberlain1949).AccordingtoBurton(1970),thesebirdsaremuchmorerestrictedtocoastalareasthanareHerringor Ring billedGulls.GeorgiaBonaparte'sGullsareerraticallycommonwintervisitorsalongtheGeorgiacoast.Mostoccurinthestatebetween21Septemberand25May(Burleigh1958, Dentonetal.1977),butone wasseenaslateas20JuneintheSavannahRiverDelta(Tomkins1958).TheseasonaldistributionofthisspeciesinGeorgiaisprobablysimilartothatinFlorida,withthebulkofthewinteringpopulationarrivingaftermid-December.FloridaSprunt(1954)calledBonaparte'sGullscommoninwinter,notingthatfewarriveinfallandthatthemaininfluxoccursinearlyormid-winter.Kale (1979msa)consideredthemcommonwinterresidentsfromOctobertoMayalongtheAtlanticcoast,andregular(1979msb)alongtheGulfcoastinvaryingnumbers fromOctobertoMarch.RelativelylownumbersarefoundalongtheAtlanticcoast,andevenfewerarefoundontheGulfcoastofthePeninsula.Thesegullsaremostcommon along theFloridapanhandle(Map3),wheretheyarriveearlierthanelsewhereinthestate.TheTampa-St.PetersburgareahasmanymoreBonaparte'sGullsinJanuary-MarchthaninDecember(W.Hoffman,pers.comm.). Alabama Imhof(1976b)regardedthisspeciesasuncommontoabundantontheAlabamaGulfcoast,whereextremedatesofoccurrenceare1Octoberand14May.WinterconcentrationsinJanuaryandFebruaryoften tothreethousandbirds(Imhof 1976b,Table20).MississippiBonaparte'sGullsarecommoninsmallflocksinwinteralongthecoast.Extremedatesofoccurrenceare25August(Jacksonand Cooley 1978a) and21May(Burleigh1944).Gullsseen27Juneand23July(JacksonandCooley 1978a)mayhavebeenbirdsthatsummeredsouthofthebreedingrange.ThespeciesevidentlydoesnotoccurinaslargeconcentrationsinMississippiasinneighboringAlabama(Table20,Map3).LouisianaBonaparte'sGullswinterregularlyinsmallnumbersincoastalLouisiana(Lowery1974).Lowerylistedextremedatesofoccurrencetobefrom29Augustto14June,butindicatedthatpeaknumberswerepresentfrommid-Octobertomid-April.TexasBlacklock(1978 ms)consideredthisgulluncommoninwinteronthecoast,wheretheyreachpeaknumbersinMarch.Oberholser(1974)reportedthattheprimaryperiodofoccurrenceisfromearlyOctobertolateApril,butBlacklock(1978ms)believedthatmostarriveinNovember. 144

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Bonaparte'sGullTable20. PeakconcentrationsofwinteringBonaparte'sGullsinthecoastalsoutheasternUnitedStates.NumberDateseenseenLocalitySourceNORTHCAROLINA194719Dec.3,500AlbemarleSoundChamberlainandChamberlain1948 1979 8Apr.5,000BirdShoalnearBeaufortLeGrand 1979d 195925Apr.21WilmingtonChamberlain1959bGEORGIA1965 2Jan.5,000offSap,eloIslandParnell1965bFLORIDA196128Dec. 200Panacea(Gulf)Stevenson1962b 1958 1Jan.450PortCanaveral(Atlantic)Stevenson1958b 196718Jan.41KeyWest(Keys)Robertson1967 1950 5Jan.2,500Pensacola(Gulf)Lowery1950 1950 11-12Jan.2,000Pensacola(Gulf)Lowery1950 192011Feb.1,500SaintAndrews Bay(Gulf) 'Howell 1932 1979Feb.500Eastpoint(Gulf)Stevenson1979a 196410Mar.3,000Pensacola(Gulf)James 1964 195725Mar.6,000nearPensacola(Gulf)Newman1957a 194828Mar.1,000Pensacola(Gulf)Weston 1948ALABAMA197222-23Jan.1,400FortMorgantoGulfShoresImhof 1976b 195831Jan.1,200+AlabamaPointNewman1958b 197419Feb.2,750+GulfShoresImhof 1976b 197818Mar.20,000AlabamaPointImhof 1978 MISSISSIPPI 198020Nov. 200+ Clermont HarborJackson1981 1977 3 Dec. 200 RossBarnettReservoirJacksonand Cooley 1978a 198027Dec. 214SardisLakeJackson1981 1978Jan-Feb.to160 HornIslandJacksonand Cooley 1978a 1978 5 Mar. 250PascagoulaRivermarshJacksonand Cooley 1978a 145

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Bonaparte'sGullTable20.Concluded.Number DateseenseenLocalitySourceLOUISIANA193231Dec. 32CalcasieuLakeOberholser1938 1937 2Apr.50GrandIsleOberholser1938TEXAS196911Jan.100FreeportjettiesWebster1969b 196913Jan.300+Gilchrist,BolivarPeninsulaWebster1969b 196711Feb.80feedingalongPortAransasWebster1967bjetty196814Apr.102PortAransasjettyWebster1968bSYNOPSISOFPRESENT DISTRIBUfiON ANDABUNDANCEBreedingBonaparte'sGullsbreedintheinteriorof Alaska andnorthernCanadaintheborealforestzone.WinterMostBonaparte'sGullswinteralongthePacificcoastfrom WashingtonsouthintoMexico,and ontheAtlanticcoastfromNewEnglandtotheYucatanPeninsula.Becausetheyarriveonthewinteringgroundwellafterthecompletionofthebreedingseason,ChristmasBirdCountsmaynotreflectthetruesizeofwinteringpopulations.RecentChristmasCounts(Map3)suggestthatthelargestwinteringpopulationsinthesoutheasternUnitedStatesarefoundalongthenorthcoastoftheGulfofMexico.SubstantialnumbersarealsopresentontheAtlanticcoastofFloridaandNorthCarolina.Insomeyears,largeconcentrationsoccurinthesoutheasternstates(Table20).MigrationCooke(1915)remarkedthatmostBonaparte'sGullswintertotheeastofthebreedingrange.MostofthebirdsthatwinteralongtheAtlanticcoastfollowthenorthernlimitoftreegrowththroughtheGulfofSt.Lawrence,butothersmovesouthby Lake WinnipegtotheGreatLakes andthencetothecoast.Cookebelievedthatonly small numbersfollowtheMississippiRiversouthtotheGulfcoast.Bonaparte'sGullswinteringalongthePacificcoastwerebelievedtomigratesouthacrosstheRockyMountainstothecoastofsouthernAlaskaandthencesouth.Cooke(1915)suggestedthatthespringroutebasicallyretracedthattakeninfall.Beardslee(1944)laterpointedoutthatverylargenumbersfollowtheNiagaraRiverwherepeaksoffallmigrationoccurfromearlyAugustthroughSeptemberandinNovember and December. ApeakofspringmigrationtakesplacetherefromearlyAprilthroughearlyMay.The146

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Bonaparte'sGullfallmigrantsarelargelyadults,whichledBeardsleetosuggestthatjuvenilestravelsouthwardoveranotherroute.Cooke(1915)remarkedthatthemostnotablecharacteristicofthefallmigrationofBonaparte'sGullisitsearlybeginning;birdsaremigratinginthenorthernportionoftherangebylateJuly.ArrivaldatesforlocalitiesalongtheAtlanticcoastarefromearlyAugusttomid-September,withprogressivelylater dates atmoresouthernlocalities.He noted.othat peakmovementsoccura monthtosixweekslaterandthatthespeciesismostabundantofftheNewEnglandcoastinOctober.Thelatestdatesofoccurrenceforbirdswinteringinthesoutheastlistedby Cooke(1915)rangefromlateMarchatNewOrleans,Louisiana,tothefirstweekofAprilinFlorida,tothefirsttwo weeksofMayinNorthandSouthCarolina.HABITATNestingBonaparte'sGullsusuallynestinspruceandotherconiferoustreesnearlakesand muskegsofCanadianandAlaskanborealforests(Bent1921, Henderson 1926,Twomey1934,JehlandSmith1970).Nestsareusuallyfromabout4-20ft(1-6m)abovetheground(authorscitedbyBent 1921, Henderson1926).Symons(1968)recordedanatypicalnestina clumpofbulrushesinaswampinSaskatchewan.OthersnestsinSaskatchewanhavebeenfoundinreedsinamarsharoundalakeand oneevenwasfoundonthemudflatofatemporarypothole(Lamont1980).FeedingBonaparte'sGullsfeedovermarshy ponds intheinterior(Bent1921).Transientandwinteringbirdsfeedinawidevarietyofhabitatsincludingfreshwatermarshes,rivers,lakes,estuaries,saltmarshes,beaches,bays,and openocean.Tomkins(1958)reportedthatsewermouthswerefavoredasfeedingareasintheSavannahRiverDeltaareainGeorgia.Bonaparte'sGullswinteringon LongIsland,NewYork,prefertofeedoverinlets,particularlyathightide(Lauro1980).NonbreedingandOffshoreInthesoutheasternUnitedStates,Bonaparte'sGullsusefreshwaterlakes,estuaries,andinlets,aswellasthenearshorewatersoftheGulfofMexico andtheAtlanticOcean. Theyregularlyroostonsandbarsandbeaches,butsomebirdsstayoffshoreforextendedperiods.Those onMontereyBay,California,primarilywinterinshoreandsecondarilyuseoffshorewaters.Theyalsooccurinlandalongsloughsandrivers(Baltzand Morejohn1977).InthenorthernChesapeakeBight,winteringandmigrantbirdsoccurlargelyovershallowcoastalwaters(Rowlett1980).MostoftheBonaparte'sGullsobservedintheBightwerefoundwithin20km(12mi)ofshore,butonefeedingflockof300birdswasseen44km(27mi)eastofOceanCity,Maryland,on 4 December 1976.Bonaparte'sGullshavealsobeenseenintheGulf upto40 mi(64km)westoftheFloridacoast(Buhrman and Hopkins1978).147

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Bonaparte'sGullFOODANDFEEDINGBEHAVIORBonaparte'sGullsfeedextensivelybyplungingforsmallfish(Wolf andGill1961).Theyareadeptathawkinginsectsandoccasionallyforageonfootalongbeaches.Lauro(1980)suggestedthatmostBonaparte'sGullswinteringon LongIslandprefertofeedeitherbysittingonthewaterandpickingitemsfromthesurfaceorbyseizingitemsatthesurfacewhileinflight.GullsfeedingatsettlingtanksinBaltimore,Maryland,prefertoseizefoodwhileinflightbutalsopickupitemswhilesittingonthesurface(R.Blum,pers.comm.). Atthislocalitytheyseldomorneverplungeforfood.MigrantsatOlsenCreek,PrinceWilliamSound,Alaska,feedondrifteggsofsalmon(Oncorhynchusspp.)bybrieflylandingonthewateranddivingforthem (Moyle1966).InthewesternLakeErieregion,migratingBonaparte'sGullscongregateinthethousandsduringthefalltofeedonemeraldshiners(Notropisatherinoides)intheharboratPut-In-Bay(Ligas1952).There,theydiveforfish,submergingcompletely,andoftenfeedoninsectseitherbyseizingthemwhileflyingorbypickingthemfromthesurfaceofthewater.King(1973)reported"foot-paddling"todisturbsmallmarineorganismsintidalpools.Bonaparte'sGullshavebeenobservedrobbingDunlins(Calidrisalpina)ofearthwormsinMichigan(PayneandHowe1976),andabirdcollectedinNewJerseyhadeatenalargequantityofwalnutmeat(Frohling1967).InFlorida,theyoftenfeedonschoolingbaitfish.AttimestheyfeedontheschoolsinflocksWith Horned Grebes(Podicepsauritus)(Dusi1968; W. Hoffman,pers.comm.).OthersseenwithRed-breastedMergansers(Mergusserrator)atBlackburnBay,Florida,fedonitemsstirredupbyfeeding lucks and Stedman1979).Insectsarethemajoritemsofdietonthenestinggrounds,asaremarineorganismsinwinter(Bent1921).Informationonthedietbasedontheexaminationofstomachcontentsisscanty.Seventy-eightstomachstakenbytheBiologicalSurveycontainedaboutone-thirdinsects."aboutone-halffishes,andtheremaindersmallsnailsandcrustaceans"(Howell1932).FourstomachsofsevenBonaparte'sGullscollectedontheBassIslandsinWesternLakeEriecontainedonlyinsects,therestonlyemeraldshiners(Ligas1952).Stinkbugsweretheinsectfoundmostfrequently,butsmallbeetles,flies,andspiderswerealsoeaten.LigasreportedthatanadultBonaparte'sGullheldcaptivefor40dayssubsistedentirelyonshinersandatefrom63.0-147.7g/day(2.2-5.2.oz/day)(mean=95.6g[3.4oz]).Thecaptivebirdweighed176 g(6.2oz)attheendoftheexperiment.Thestomachcontentsof23migrantBonaparte'sGullsthatwerecollectedinAugust and SeptembernearDeerIsland,NewBrunswick,wereexamined by Braune andGaskin(1982a)whoreportedthatthesebirdshadfedmostlyoneuphausiidcrustaceans(Meganyctiphanesnorvegica,Thysanoessainermis)andinsectsfromavarietyoforders.Representativesofbothgroupsoccurredinequalfrequency,buteuphausiidsconstitutedamajorityofthedietbyweight.Smallherring(Clupeaharengus),pollock(Pollachiusvirens),andrainbowsmelt(Osmerus mordax)werealsosignificantinthediet.148

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Bonaparte'sGullStomachsof12Bonaparte'sGullswinteringonMontereyBay,California,containedinvertebrates(93.1% byfrequency)andsmallfish(6.9%)(Baltzand Morejohn1977).Bonaparte'sGullhadthemostvarieddietofanyofthesixspeciesofgullsexamined.Twohundredinsectpupae(47.6%ofallfooditemsfound),62crabmegalops(16.9%),40unidentifiednereids(9.5%),and 30euphausiidcrustaceans('fhysanoessaspinifera7.1%) werethefooditemsmostfrequentlyfound.FoodsidentifiedtoeitherspeciesorgenuswereThysanoessa,rockfish(Sebastesspp.-3.8%),smallcrustaceans(Euphausiasp.-3.1%;Pasiphaeapacifica1.4%; Hyalesp.1.2%;Idotheaspp.-0.9%),andthenorthernanchovy(Engraulismordax -1.7%).Oneeachofthreebenthicspeciesoffish(plainfinmidshipman[Porichthysnotatus],redbrotula[Brosmophycismarginata],andspottedcusk-eel[Chilarataylori])wasalsoidentifiedinthestomachcontents.IMPORTANTBIOLOGICALPARAMETERSThebreedingbiologyofBonaparte'sGullisvirtuallyunstudied.AsHenderson(1926).indicated, the habitofbreedingnoncolonially,orinverydispersedcoloniesinblackfly-infestedmuskeg swamps,hasdeterredmostornithologistsfromobservingtheirhabits.EggLaying1968),lateMay1934),andmid-NestinghasbeenreportedinlateMayinSaskatchewan(SymonstoearlyJuneinAlberta(Henderson1926,Farley1931,TwomeytolateJuneinManitoba(Jehland Smith1970).MeanClutchSizeFewdataareavailableonmeanclutchsize(Table21).Mostnestscontainthreeeggs,althoughsomecontainonlytwo.Clutchesoffourarerarelyfound(Bent1921).Table21.MeanclutchsizesreportedfortheBonaparte'sGull.Meanclutchsize2.833.002.95Numberofclutches6420LocalityandperiodofobservationAlberta,nearAthabascaLandingSaskatchewan,nearMeotaManitoba,Churchill,1964-1967SourceHenderson1926 Symons 1968Jehland Smith 1970IncubationPeriodJehland Smith(1970)reportedanincubationperiodof23-24days.HatchingSuccessAtChurchill,Manitoba,16of38eggshatched,arateof42%(Jehl1971).149

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Bonaparte'sGullAgeatFledgingUnknown.FledgingSuccessUnknown.MortalityofEggsandYoung Most (96%)oftheeggslostfromnestsoftheBonaparte'sGullatChurchill,Manitoba,werelosttoinclementweather;eggswere blownoutofthenestsbyhighwinds(Jehl1971).RenestingHenderson(1926)describedoneclutchasanapparentreplacementclutch.AgeatFirstBreedingAgeatfirstbreedinghasnotbeendetermined.Bonaparte'sGullsapparentlygainfullbreedingplumageattwoyearsofage(Dwight1925)andtheymaybreedthen.MaximumNaturalLongevityNodataareavailable.Weight Gross(1937)reportedtheweightofanunsexedlatemigranttakenintheMagdalenIslandsinJuneas201g(7.1oz).WeightsoffourbreedingmalestakennearNapaskiak,Alaska,inJunerangedfrom182to202 g(6.4-7.1oz),andtheweightofasinglebreedingfemalewas 169 g(6.0oz)(Williamson1957).The meanweightoffiveBonaparte'sGulls,allevidentlywinteringbirdstakenatMontereyBay,California,was176.8g(6.2oz)(BaltzandMorejohn1977).SUSCEPTIBILITYTOOILPOLLUTIONTherearefewrecordsofoiledBonaparte'sGulls.Kramer and Kramer(1945)foundthreedead,oiledbirdsofthisspeciesamong92birdsof11speciesfounddead,mostoiled,inalO-milestretchofbeachinNewJersey.DuMont(1977)sawanoil-stainedbirdamong agroupof150gullsatOceanCity,Maryland.Phillips(1974)saw abirdinCornwall,England,thatwasoiledonthelowerbellyandvent.Fivedayslaterthisbirdwascleanandapparentlysufferingnoill Phillipsbelievedthatthiscleaningresultedfrompreening,butMather(1974)expressedskepticismaboutgulls'abilitytocleantheirplumagesolelythroughpreeningandsuggestedthatthecleanappearancemay have beenduetomolt.Bonaparte'sGullsoccurinlargenumbersinthesoutheasternUnitedStates,sometimesinlargeconcentrations,bothneartheshoreandwelloutfromit.Theyfrequentlyalightonthewaterandfeedfromthesurface.However,theytendtooccurinscatteredflocks,and numberspresentinanyareavaryconsiderablyfromyeartoyear.Consequently,amajoroilspillprobablywouldhavelittleeffectonthetotalpopulation.Long-termchronicoilpollutionofthecoastmighthavemoreseriousconsequences,butmoreinformationonthenumbers andstatusofthisgull,bothwithinthesoutheastandelsewhere,isneededtoevaluateadequatelythepotentialeffectsofoiling.150

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Bonaparte'sGullBIBLIOGRAPHY1982Braune,B.M.andD.E.Gaskin.1982a.FeedingecologyofnonbreedingpopulationsoflaridsoffDeerIsland,NewBrunswick.Auk99:67-76.1982b.Feedingmethods anddivingratesofmigratinglaridsoffDeerIsland,NewBrunswick.Can.J.Zool.60:2190-2197.1980 Lamont,S.M.1980.AtypicalnestofBonaparte'sGull.BlueJay38:44-45.Lauro,A.J.1980.ofLongIsland.ThewinterecologyofBonaparte'sGullonthesouthshoreLinn.Newsl.34(1):1-3.1979Braune,B.M.1979.FeedingecologyofmigratingpopulationsofBonaparte'sGulls,CommonTerns,andArcticTerns.M.S.thesis,Univ.Guelph/Guelph,ON.Britton,D.1979.IdentificationofBonaparte'sGull.Brit.Birds72:339340.Stedman,S.andA. 1979.FeedingassociationbetweenBonaparte'sGullandRed-breastedMergansers.Fla.FieldNat.7:27.1978Grant,P.J.1978.FieldidentificationofwestPalearcticgulls.Brit.Birds71:145-176.Grow,R.1978.AkleptoparasiticactionbyBonaparte'sGullsonYellowlegs.IndianaAudubonQ.56:178-179.Ryder,R.A.ments.1978.GullsinColorado:theirdistribution,status,and moveProc.1977Conf.ColonialWaterbirdGroup1:3-9.1977Baltz,D.M.and G.V.Morejohn.1977. Foodhabitsandnicheoverlapofseabirdswinteringon Monterey Bay,California.Auk94:526-543.Vermeer,K.1977.SomeobservationsonArcticLoons,Brandt'sCormorantsandBonaparte'sGullsatActivePass,BritishColumbia.Murrelet58:45-47.151

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Bonaparte'sGull1976Payne,R. B. andH.F.Howe.1976.Cleptoparasitismbygullsofmigratingshorebirds.WilsonBull.88:349-351.1974Mather.J.R.plumage.1974.[Commenton]OiledseabirdssuccessfullycleaningtheirBrit.Birds67: 483-484.Phillips,J.1974. seabirdssuccessfullycleaningtheirplumage.Brit.Birds67: 483.Ree,V.1974.KanadiskhettemakeobservertiNorge.[Bonaparte'sGullobservedinNorway.]Sterna13: 109-115.[InNorwegianwithEnglishsummary.]1973King,B.1973.organisms.Bonaparte'sGullinCornwall"foot-paddling"todisturbBrit.Birds66: 447.Pujals,J.J.1973.FirstphotographofBonaparte'sGullinPanama.Am.Birds27: 840.Raines,R.J.1973.Bonaparte'sGulloffBirkenhead.CheshireBirdRep.1973:26-27.1972Alsop,F.J.,III.1972.Bonaparte'sGullinGreatSmokyMountainsNationalPark.Migrant43:72-73.Seymour,N.R.1972.Successofthreegullspeciesfeedingon swarmingantsinAntigonishCounty,NovaScotia.Can.Field-Nat.86: 391-392. 1971Jehl,J.R.1971.Patternsofhatchingsuccessinsubarcticbirds.Ecology52:169-173.1970Jehl,J.R.andB.A.Smith.1970.BirdsoftheChurchillregion,Manitoba.Manit.Mus.ManNat.Spec.Publ.1.86pp.1968Burger,J.and R.Brownstein.1968.ThestatusofBonaparte'sGullinNewYorkState.Kingbird18:9-20.152

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Dusi,J.L.Grebes.Bonaparte'sGull1968.FeedinginteractionbetweenBonaparte'sGullsand HornedAuk85: 129.Parkin,D.T. andP.Parkin.1968.Bonaparte'sGullinSutherland.Scott.Birds5:175.Rusk,M.S.andF.G.Scheider.1968.AdditionstoBonaparte'sGullrecords,Region5.Kingbird18: 84.Symons,R.D.1968.AtypicalnestingofBonaparte'sGullinSaskatchewan.BlueJay26:70-74.1967Burger,J.1967.NotesontheroostingbehaviorofBonaparte'sGull.Kingbird17: 213.Frohling,R.C.1967.Bonaparte'sGullfeedingonwalnutmeat.WilsonBull.79: 341. 1966Moyle,P.1966.FeedingbehavioroftheGlaucous-wingedGullonanAlaskansalmonstream.WilsonBull.78:175-190.1965Schmidt,G.D.1965.Corynosomabipapillumsp.n.fromBonaparte'sGullLarusphiladelphiainAlaska,withanoteon C.constrictumVanCleave,1918.J.Parasitol.51: 814-816. 1964Post,P.W.1964.OnthereluctanceofBonaparte'sGullstoflyunderobjects.Kingbird,14:91.1961Kist,J.bij34: 1961.WaarnemingenvandeKleineKokmeeuw,Larusphiladelphia(Ord),Scheveningen.[Larusphiladelphia,newtotheDutchlist.)Limosa220-224.[InDutchwithEnglishsummary.) Wolf, L.L.andF.B.Gill.1961.FlockfeedingbehaviorinmigrantBonaparte'sGulls.WilsonBull.73: 389-390. 1956Bard,F.G.1956.Bonaparte'sGull.BlueJay14: 80.Little,B.1956.Bonaparte'sGullinNorthumberland.Brit.Birds49: 324-325. 153

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Bonaparte'sGullMoynihan,M.1956.Notes onthebehaviorofsomeNorthAmericangulls.I.Aerialhostilebehavior.Behaviour10:126-178.1954 Moule,G.W.H.1954.Bonaparte'sGullinDorset.Brit.Birds47:272-273.1953Taber,W.1953.WinterstatusoftheBonaparte'sGullinMaine.Bull.Maine AudubonSoc.9:35-36.1952Harber,D.D.1952.Bonaparte'sGullinSussex.Brit.Birds45:333.1948Brackbill,H. 1948.Bonaparte'sGulland AmericanPipitinwinter.Md.Nat.18:14-15.1944Beardslee,C.S.1944.Bonaparte'sGullontheNiagaraRiverandeasternLakeErie.WilsonBull.56:9-14.1940Cruickshank,A.D.1940.Albinismingulls.Proc.Linn.Soc.N.Y.50-51:31-32.Nichols,D.G.1940.FeedinghabitofBonaparteGull.Gull22:41.1936 Munro,J.A.1936.NestingoftheBonaparteGullinBritishColumbia.Condor38:60-61.1934Twomey,A.C. 1934.BreedinghabitsofBonaparte'sGull.Auk51:291-296.1931Bunyard,P.F.1931.[OntheeggsofTringaamericanaandLarusphiladelphiafromAlberta.]Bull.Br.Ornithol.Club51:9-12.Farley,F.L.1931.centralAlberta.NestingofBonaparte'sGull(Larusphiladelphia)inCan.Field-Nat.45:138-139.154

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Bonaparte'sGull1930Murray,J.J.1930.Bonaparte'sGullatLexington,Virginia.Auk47:72.1929Saunders,W.E.1929.Bonaparte'sGullsinunusualnumbers.Oologist46:2.1928Stubbs,A.P.1928. SummeringBonaparte'sGullatLynn, Mass.Auk45:91-92.1927Nichols,J.T. 1927. BrownPelicanand summeringBonaparte'sGullsatNewport, R.I. Auk44:555.1926Bunyard,P.F.1926. [Note onthenestandeggsofLarusphiladelphiaandEreneutespusillus.]Bull.Br.Ornithol.Club 46:108-112.Henderson,A. D. 1926.Bonaparte'sGullnestinginnorthernAlberta.Auk43:288-294. Rowan, W.1926.OnthebreedingoftheBonaparteGull.Bull.Br.Oologic.Assoc.1:112-116.1924Urner,C.A.1924.Bonaparte'sGullsummeringnearPerthAmboy,N.J.Auk41:149.1910Seguin-Jard,E. 1910.Premierecaptured'unLarusphiladelphiasurlescotesdelaVendee.NantesBull.Soc.Sci.Nat.10:97-100.1902Eifrig,G.1902. Remarkable flightofgullsatCumberland,Md.Auk19:75.1901 Loomis,L.M.1901.OnthesouthernlimitofthewinterrangeofBonaparte'sGull(Larusphiladelphia).Auk18:104-105.155

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RING-BILLEDGULL(Larusdelawarensis)IGE:Ringschnabelmowe,SP:Apipizcapinta]GENERALDISTRIBUTIONNorthAmericaRing-billedGullsbreedalmostentirelyintheinteriorofNorthAmericafromnortheasternCalifornia,centralOregon,andcentralWashingtoneastthroughthenorthernGreatBasin(southtoColorado)andtheRockyMountainStates,northinthewesternGreatPlainstonorthernAlberta,Saskatchewan,southcentralManitoba,andtheGreatLakes,andlocallyalongtheSt.LawrenceRiver,tonortheasternNewfoundland andNewBrunswick,southernandsoutheasternLabrador,andsouthtoLakeChamplaininNewYork(AOU1957,Godfrey1966,Southern1980a).TheyhavebeenreportedbreedingalongJames Bay,butthisneedsconfirmation(Godfrey1966).Recently,theywerefoundbreedingontheWashingtoncoast(PenlandandJeffries1977) and on Lake CalumetinIllinois(Kleen1975).Ring-billedGullsthatbreedwestof960WwinteronthePacificcoast(Southern1974a)southfromsouthernBritishColumbia(AOU1957,Godfrey1966)commonlytonorthernBajaCalifornia,butarerarefarthersouth(AOU1957,Southern1980a).PopulationsfromtheGreatLakesand CanadawinterprimarilyontheAtlanticandGulfcoasts,although some winterintheGreatLakeswhereveropenwaterremains(SouthernandMoore1974).ThemajormigrationrouteiseastthroughtheGreatLakestostagingareasonLakesErieandOntario,eastthroughPennsylvaniaandNewYorktotheAtlanticcoast,thensouthtoFloridaandtheGulfofMexico(Southern1967a,1974a).MostofthebirdswinteringontheAtlanticcoastandeasternGulfofMexicoapparentlyfollowthisroute.BirdswinteringinthewesternGulfofMexico,however,probablymovesouththroughtheplainsandtheMississippiValleyfromtheCanadianPrairieProvincesandthenorthernPlainsStates.BirdsfromtheGreatLakesoccasionallywanderoutsidetheirnormalrangewestandnorthwesttotheYukon,BritishColumbia,Saskatchewan,James Bay, Quebec,Oregon,California,andColorado(Southern1974b,Southernand Moore1974).WorldDistributionRing-billedGullsoccuralmostsolelyinNorthAmerica.SomewinterregularlyintheGreaterAntilles,andothersoccurcasuallysouthtoTrinidad(ffrench1973),Aruba (Voous1977),Honduras(Southern1974b, BrownandMonroe1974,SouthernandMoore1974),Panama (Wetmore1965,Southern1974b,Southernand Moore1974),and Colombia(Southern1974b,Southernand Moore1974).TheywandertoHawaii(Berger1972) andtheLineIslands(King1955).IntheOldWorld,Ring-billedGullshavebeenreportedinBritain(Hume1973,P.J.Grant1979),Ireland(Mullarney1980),Spain(Southern1974b,Southernand Moore1974),andcentralGermany(Berndtand Rahne1968).Atleast64werere-156

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Ring-billedGullcordedinBritainandIrelandthrough1980(Rogersetal.1982).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESAlthoughtheoccurrenceofraremarinebirdsinthesoutheastisusuallywelldocumented,thepresenceofcommontransientorwinteringspeciesisusuallypoorlyreported.TheRing-billedGullisnoexception.AmericanBirds,whichreportsseasonalobservationsofbirds,madeonlya fewscatteredreferencestoRing-billedGullsinthesouthduringaperiodof10years,althoughthisgulloccursthereinthethousandsduringthewinter.Consequently,wehavelittledetailedinformationonitsstatusinthesewaters.NorthCarolinaRing-billedGullsareabundantinwinteralongthecoastandareregularasnonbreedersinsummer(Pearsonetal.1942).AccordingtoSouthern(1974a),mostoftheGreatLakesRing-billedGullspassthroughNorthCarolinaonfallmigration.However,ChristmasCountdata(Map4)indicatethatlargenumbers,asmanyas16,000percoastalcount,mayremaintowinter.SouthCarolinaSpruntandChamberlain(1949)regardedRing-billedGullsasabundantinwinterinSouthCarolinabutmuchlesscommoninsummer.Forsythe(1973)foundthatthenumberofRing-billedGullsintheCharlestonmetropolitanarearangedfromunder100birdsinsummertoover3,500duringtheperiodfromlateNovemberthroughmid-April.InDecember andFebruary,somewhatlowernumbers(butstillseveralthousand)werepresent.Forsythe(1972)examinedSouthCarolinaband recoveries ofRing-billedGullsandfoundmostofthemtobefromtheGreatLakesregion;a fewwerefromnortheasternAtlanticareas.Mostrecoverieswereoffirst-year(53%) andsecond-yearbirds(17%).Forsythe's(1973)analysisofChristmasCountsshowedthatpopulationsofthisspecieshadincreasedintheCharlestonareafrom1957through1972.Our summaryofrecentChristmasCountdata(Map4)suggeststhatfarfewerRing-billedGullswinterinSouthCarolinathaninNorthCarolina;theaveragesforfourcoastalcountswereallbelow1,500.GeorgiaDentonetal.(1977)statedthatRing-billedGullsareabundantontheGeorgiacoastinfall,winter,andspring,andthata fewnonbreedingbirdsarepresentinsummer.Norris(1939)foundthisspeciesmuch moreabundantinJuneatSt.SimonsIslandthantheresidentLaughingGull.Most(ca.80%)oftheRing-billedGullsseenby himwereimmatures.Atthenorthernborderofthestate,Ring-billedGullsarriveinOctober,butbegintoreturnnorthinlateApril(Tomkins1958);mostaregonebyearlyMay(Burleigh1958).FloridaThestatusoftheRing-billedGullinFloridahaschangedmarkedlyinthiscentury.Howell(1932)calledthespeciesacommonwinterresidentinonlythenorthernpartofthestate,andindicatedthatitwaslesscommonthantheHerringGull.Hehad norecordsfortheFloridaKeys.HealsonotedthatthebulkofthepopulationarrivedinlateOctoberand November.Sprunt(1954)notedthattheRing-billedGullhad become morecommonandwidespread,andoccurredallthewaytoKeyWest.Bandrecoveries(KadlecandDrury1968a,157

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linter listrillltillapferSdemn litedStatesBIRDSPER10PARTY-HOURS c::::::J Lessthan10 _10-50_ 50-200_ More than200 (Aclapted 'rolll Iyltrak. 19741INDIVIDUALSOBSERVEDDURINGCHRISTMASBIRDCOUNTS,1973-1977(ARITHMETIC MEAN)@Number of individuals o Lessthan one individual None observed 92-GULFOFMEXICOMap4 BIRO NAME'

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Ring-billedGullSouthern1974a)indicatethatFloridaisnowthemainwinteringareaforGreatLakesRing-billedGulls.Southern'sdatashowthat68%ofallFebruaryrecoveriesand54%ofallJanuaryrecoveriesofGreatLakesRing-billedGullswere fromFlorida.Inaddition,73%ofthesoutheasternrecoveriesofjuvenilesbandedatRogersCity,Michigan,wereinFlorida.Southernalsonotedthatamajority(upto62%)oftheFloridaencountersoccurredontheAtlanticcoast.Florida-AtlanticCoastTheRing-billedGullisanabundantwinterresidentalongtheFloridaAtlanticcoast,whereKale(1979msa)considereditthemost numerouswinteringgull.Flocksofseveralthousandareseenandmanycongregateneargarbagedumps,fishingvessels,andinlets(Kale1979msa).Averagesfor eight-coastalAtlanticChi!stntaSiCounts ranged from about450toover11,000.Fiveofthesecountsaveragedover1,000gulls(Map4).FloridaKeys Hundley andHames(1960-62)describedtheextensionofthewinteringrangeintotheFloridaKeys. ThefirstFloridaKeysrecordmentionedwasin1941,butbythelate1950'sRing-billedGullsweremorecommonat Key WestthanwereHerringGulls.The Key Largo/Plantation Key and Lower KeysChristmasCountsaverageabout100Ring-billedGullsayear.TheexpansionoftheRing-billedGull'swinterrangeinSouthFloridaandtheKeysparallelsboththeremarkablegrowthoftheGreatLakesbreedingpopulationsandthecommercialdevelopmentofSouthFlorida,whichhasincreasedthehabitatavailabletocommensalgulls.Florida-GulfCoastRing-billedGullsareabundantinwinterontheFloridaGulfcoast,particularlyneargarbagedumps (Kale msb),andaremuchmorecommonthanontheAtlanticcoast(SchreiberandSchreiber1977).TheaveragesfortenGulfCoastChristmasCountsrangefroma fewhundredtoover12,000.ThehighestcountsarefromtheTampaBayandSarasotaareas(Map4),whicharethemosturbanareasonthiscoast.About43,000Ring-billedGullswerepresentatSt.Petersburgduringthewinterof1976-1977(Kale1979msb).AlabamaRing-billedGullsareabundantontheAlabamacoast,especiallyinwinter.Mastof61bandedRing-billedGullsrecoveredinAlabama hadcomefromtheGreatLakes,butoneeachhad comefromNorthDakota andSaskatchewan(Imhof1976b).RecentChristmasCountsgiveaveragecountsinthelowhundredsforfourcoastalAlabamaareacounts(Map4).MississippiRing-billedGullsarecommonwinterresidentsontheMississippicoast,andsmallflocksarefound'therefromearlyOctobertolateApril.FromJuneuntilSeptemberonlysmallnumbersarepresent(Burleigh1944).Fewquantitativedataareavailableonwinterpopulations.RecentJacksonCountyChristmasCountsaveraged261Ring-billedGulls(Map4),suggestingthatwinterpopulationsmaybeinthelowthousands.LouisianaOberholser(1938)and Lowery(1974)bothconsideredRing-billedGullsabundantinwinter,butrareinsummer. Lowery(1974)indicatedthatmigrantsarriveinnumbersinOctoberanddepartmostlyby 1 May.RecentNewOrleansandSabineChristmasCountsaverage3,420and1,044Ring-billedGulls,159

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Ring-billedGullrespectively.TexasRing-billedGullsarecommontoabundantinwinterontheTexascoast,anduncommoninsummer(Oberholser1963ms,1974).ApparentlyfewGreatLakesbirdsreachTexas.Southern(1967a)reportedno Texasrecoveriesamong376recoveriesofjuvenileandyearlinggullsfromaMichigancolony;hehad78recoverieselsewhereinthesoutheast.A morerecentanalysis(Southern1980a),basedonnearly21,500recoveriesforRing-billedGullsbandedthroughouttheirrange,showsthatthisspeciesisfairlycommonontheTexascoastinwinter.MostoftheRing-billedGullsinTexas andLouisianaprobablycomedirectlyfromthePlainscolonies.RecentChristmasCountsaverageabout200-2,000Ring-billedGullspercount;thehighestcountstendtocomefromurbanareas.SYNOPSISOFPRESENTDISTRIBUTIONANDABUNDANCEBreedingRing-billedGullsbreedinmaritimeCanada,throughouttheGreatLakes,acrossthePrairieProvincesofCanada,andinthenorthernGreatPlains,theRockyMountains,andtheGreatBasinofthewesternUnitedStates.Somefiguresareavailablefortheeasternpopulations.Ludwig(1974)estimatedtheGreatLakespopulationin1967as335,000breedingpairs.OntheU.S.portionoftheGreatLakestherewereabout90,000breedingpairsin1976, andabout103,000breedingpairsin1977(Scharf1978).In1977, LakesMichigan,Ontario,and Huronsupportedmostofthepopulation(33.3%,31.9%, and 25.1%,respectively);muchsmallerproportionsnestedon LakeErie(6.8%) and LakeSuperior(2.9%).TheCanadianportionofLakeOntarioandtheupperSt.LawrenceRiverheldabreedingpopulationofabout45,700 pairs in1976(Blokpoel1977).Most(ca.75%)bredattwosites:onGullIslandinPresqu'ileBay(ca.23,700pairs),andatLeslieSpit,theEasternHeadlandoftheTorontoOuterHarbor(ca.10,400pairs).Populationsinthisareahavebeenincreasingsince1967(Blokpoel1977).About62,000Ring-billedGullsnestedontheEasternHeadlandin1979(CourtneyandBlokpoel1980a).About36,600pairsofRing-billedGullswerefoundnestinginAlbertaduringapartialcensusoftheprovincein1976-1977 (Weselohetal.1978).Abouthalfwerefoundinsevencoloniesintheshort-grassprairieregion,andtherestinfourcoloniesinaspenparkland;figuresforcoloniesintheborealforestregionwerenotavailable.The numberbreedingintheshort-grassareawas nodifferentfromthatfoundduringacensusin1968 (Weselohetal.1978),butthenumberinaspenparklandwas morethandoublethatfoundduringtheearliersurvey.Mostofthisincreasewas duetoatenfoldincreaseinthenumberbreedingatMiquelonLake.Weselohetale(1978)suggestedthattheincreaseatthiscolonymayhavebeenduetothegulls'discoveryofthefoodresourcesprovidedbygarbagedumpsatEdmonton,approximately35 mi (56km)fromthecolony.The numberofRing-billedGullsnestinginthewesternUnitedStatespriorto1930 wassmall,perhapsontheorderof4,700-6,700birds,butithasincreasedsubstantiallysincethattime(Conover and Conover1981).About17,500Ring-billedGullsbredinWashingtonin1977(Conoveretale1979b).Recent160

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Ring-billedGullpopulationestimatesforbirdsbreedinginotherwesternstatesareunavailable.Insummary,incompleteinformationonbreedingpopulationsofRing-billedGullssuggeststhatthetotalworldbreedingpopulationisatleastahalfmillionbirds.WinterThewesternpopulationswintermainlyonthePacificcoastofNorthAmerica.YoungfromwesternAlbertawintermostlyinsouthernCaliforniaandalongthewestcoastofMexico (Vermeer1970).TheGreatLakespopulationwintersontheAtlanticandGulfcoasts,althougha fewremainintheinterior.Floridahasthelargestwinteringconcentrationsintheeast.SmallernumberswinterintheBahamas andtheGreaterAntilles.ManyofthebirdsintheGulfofMexicoevidentlycome fromthenorthernGreatPlains.MigrationRing-billedGullsnestinginwesternNorthAmericamigratesouthorsouthwestinfall,whilethosefromtheGreatLakesprimarilymovesouthorsoutheast.Young banded onthenorthshoreoftheGulfofSt.Lawrencedispersedwidelyafter withsome movingnortheasttoLabradorandotherswanderinguptheSt.LawrenceRiver(Lewis1941).Most moveddowntheAtlanticcoastinOctober,reachingNorthCarolinaby November, andFloridaby December.DeparturefromthebreedingcoloniesintheGreatLakesregionbeginsinmid-JulyandAugust;byJanuarymostofthebirdsareonthewinteringgrounds.SpringmigrationbeginsinlateFebruaryandpeaksinMarch andApril.RingbilledGullsoftenarriveatcoloniesasearlyaslate intheGreatLakes(Southern1980a)andasearlyasAprilinwesternAlberta(Vermeer1970).HABITATNestingRing-billedGullsnestprimarilyonislandsandoccasionallypeninsulas(Baird1976)inlargefreshwaterlakes(Southern1980a).A fewcoloniesareonoceanicislandsinmaritimeCanada(Southern1980a),orincoastalsituationsinWashington(PenlandandJeffries1977).Coloniesareusuallylocatedinlowvegetationratherthanon abaresubstrate.Thisspeciesfrequentlynestsinmixedcolonieswithotherlarids,includingHerringGullsandCaliforiaGulls(Laruscalifornicus).Ring-billedGullsatseverallakesinMontananestednearlow,sparsevegetation;meanheightsofthesurroundingvegetationrangedfrom17.0to135.5cm(6.7-53.3in),and meanvegetativecoverrangedfrom59.9%to86.4%(Baird1976).NestdensityatthreecoloniesontheFreezeoutPeninsulaand ArodIslandvariedfrom0.246nests/sqm(2.3nests/100sqft)to0.539nests/sqm(5.0nests/100sqft),butBairdfoundnosignificantcorrelationbetweennestdensityordistancetothenearestneighborandheightofvegetationoramountofcover.Ring-billedGullson twoislandsinMiquelonLake,Alberta,prefertonestinflat,elevated,sparselyvegetatedareasnearthecenterofcoloniesthey161

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Ring-billedGullsharewithCaliforniaGulls(Vermeer1970).Vermeer(1970)andBlokpoelandCourtney(1980)pointedoutthatnest-siteselectionisrelatedtonestlocationfromthepreviousyear.BlokpoelandCourtneynotedthatchangesinnestsitelocationfromyeartoyearwerelessthan6m(20ft)in13of24cases.Theyalsoreportedthatthetendencyofabreedingbirdtoreturntoitsformercolonyincreaseswithage.Southern(1977)reportedthat90%ofthebirdsbandedasadultsatonecolonyweresubsequentlyrecapturedthere,butindicatedthatyoungaremuchlesslikelytoreturntothenatalcolony.Thepresenceofotherseabirdsmayalsodeterminenest-siteselectionbyRing-billedGulls.Vermeer(1970)remarkedthatthepresenceofCaliforniaGullsinRing-billedGullcolonieslimitedthechoiceofnestsitesforthelatter.Whenthetwospeciesnesttogether,theCaliforniaGullsnestclosetothewaterattheperipheryofthecolonyandtheRing-billedGullsnestintheinterior.WhentheCaliforniaGullsnolongernestinanareatheyformerlyoccupied,Ring-billedGullsmoveintotheseareasandthennestclosertothewater.Lewis(1941)reportedthatRing-billedGullsonthenorthernshoreoftheGulfofSt.LawrenceinQuebecnestedmostlyonsmalltomoderatelysizedrockyislands20-60ft(6-18m)high.Mostoftheseislandsweresparselyvegetatedandtreeless,andgrassyareaswereoftenusedasnestsites.FeedingRing-billedGullsfeedinplowedfieldsandpastures(SpruntandChamberlain1949,Hailman1960a,Kirkham andMorris1979),ontidalflatsinsaltmarshes,alongtheshore,alongbeaches,andinshallow waters.RingbilledGullsbreedingatFreezeoutLake,Montana,foragedata meandistanceof10.8km(6.7mi)fromthecenterofthecolony;918of1,029(89.2%)birdsseenforagingwerewithin15km(9mi)ofthecolony,buta fewwereseenfeeding muchas31km(19mi)away(Baird1976).ThegullsatFreezeoutLakepreferredtofeedonirrigatedratherthandryfarmlands.InpreviousstudiesatFreezeoutLake,Rothweiler(1960)observed7 markedbirdsfeedingatdistancesrangingfrom 4to20 mi(6-32km)fromthecolony.NonbreedingandOffshoreMostRing-billedGullswinteringinthesoutheasternUnitedStatesoccuronornearthecoastbutsomewinterinlandonreservoirs,lakes,ponds,andstreams(Sprunt1954,Lowery1974,Imhof 1976b,Oberholser1974).NearCharleston,SouthCarolina,Ring-billedGullsoccurinlandmoreoftenthanotherspeciesofgulls;theyarefoundupto30 mi(48km)fromtheoceanorriversin"sanitaryfills,golfcourses,shoppingcenters,andshortgrassfields"(Forsythe1973).Onthecoasttheyfrequentbays,estuaries,beaches,mudflats,andthenearshorewatersoftheAtlanticOcean andGulfofMexico.Theyroostonexposedsandbarsandislands,and donotoftensleeponthewater.Ring-billedGullsareoftencommonaroundwharves,docks,andharbors(Burleigh1958),andtheyaremuchlesspelagicthanothergullswintering in thesoutheast.TheyareuncommoninoffshorewatersofthenorthernChesapeakeBightinfallmigrationandwinter(Rowlett1980),althoughthenabundantininshoreareas.MostofRowlett'soffshoreobservationsofRing-billedGullswerefrom NovemberthroughAprilandwerewithinthe10-fathomline(atabout19km[12mil).Hismaximumcounts,162

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Ring-billedGull21within15km(9mi)ofshoreinlateApril,and15within10km(62mi)ofshoreinearlyDecember,weremade whenthesegullswereundoubtedlypresentinthethousandsonshoreandthusemphasizethescarcityofthisgullinpelagicwaters.FOODANDFEEDINGBEHAVIORDuringthebreedingseason,Ring-billedGullsfeedextensivelyoninsects,grains,andsmallfish.Smallrodents,earthworms,andrefusemayalsobeimportantinthediet.WinteringRing-billedGullsfeedonfish(Tomkins ,1964, Leck1971),marineinvertebrates,insects,andenormousquantitiesofcarrionandgarbage(Forsythe1974b;W.Hoffman,pers.comm.). Theyalsoeatthefruitofthecabbagepalmetto(Sabalpalmetto)inFlorida(Nicholson1948c,Cruickshank1950,Harper1950).Aflockofabout65wasseeneatingtheripefruitofthedatepalm(Phoenixdactylifera)inCalifornia(G.Grant1979).Thegullsfedondatesbeneaththetrees,landedinthetreestopickfruit,orseizedfruitwhileonthewing.Ring-billedGullsfeedprimarilyduringthedaybutwillfeedatnightifthereissufficientlight.Leck(1971)sawabout250Ring-billedGullsfeeding pn fishundermercury-vaporlightsalongtheChesapeakeBayBridge,Virginia,inOctober;Nunnallyetal.(1979)saw themfeedingonbaitfishatalightedpieratWrightsvilleBeach,NorthCarolina,inDecember. Kirkham andMorris(1979)reportedtwopeaksoffeedingforchicksatGullIsland,Ontarioduringthesummerof1977;eachlastedabouttwohours.The first occurredshortlyaftersunrise,andthesecondjustbeforesunset.Ring-billedGullshawkforflyingantsandotherinsects(Alcorn1943,Sheppard1945,Trautman and Trautman1945,Pettingill1958a,Burton1960,MuellerandBerger1965,Tolonen1970,Seymour1972,JarvisandSouthern1976,KirkhamandMorris1979).Theyforageextensivelyinpasturesand plowedfields(Kirkham andMorris1979; R.Clapp,pers.observ.),wheretheypickuparthropods andearthworms.Someofthegroundsquirrelseatenareprobablyscavengedalongroads (Munro 1936,Vermeer1970),althoughthegullsareknowntohuntandkillvoles.Scavengingalongbeaches,inparks,andatgarbagedumpsmaybethemainwayofforagingbyRing-billedGullswinteringinthesoutheasternUnitedStates.Ring-billedGullsalsokleptoparastize(stealfood)fromotherspecies.GullsatadumpinNewJerseystole food fromStarlings(Sturnusvulgaris)(BurgerandGochfeld1981a).Adultsandsubadultsweremoresuccessfulatstealingfoodthanwereyoungbirds.Ring-billedGullshavealsobeenseenstealingfoodfromotherRing-billedGulls(PayneandHowe1976,Elstonetal.1978),Bonaparte'sGulls,Dunlins(Calidrisalpina)(PayneandHowe1976),CommonGoldeneyes(Bucephalaclangula),Buffleheads(Bucephalaalbeola),LesserScaup(Aythyaaffinis)(Siegfried1972,Grace1980),Canvasbacks(Aythyavalisineria)(Grace1980),CommonMergansers (Mergus merganser)(Lamore1953),RedbreastedMergansers(Mergusserrator)(Bunker1946),Pied-billedGrebes(Podilymbuspodiceps)(Scott1977),and AmericanCoots(Fulicaamericana) (Bartlett 163

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Ring-billedGull1957,Hardin1971,Grace1980).TheyhavebeenseenattemptingtostealfoodfromMallards(Anasplatyrhynchos)(Grace1980),andCommonEgrets(Casmerodiusalbus)(Leck Grace(1980)reportedthatRing-billedGullsobtainfoodfromwaterfowlby oneoftwomethods.Mostfrequently(ca.90%ofthetime),agullrestingonthewaterfliestowardafeedingduckandforcesittodive,subsequentlysettlingonthethewatertoseizethedroppedfood.Lessfrequently,RingbilledGullsinflightsweep down on afeedingduckandalightonthewatertoseizeadiscardedfish.Fishwereobtainedon13.3%ofallattempts.Otherfeedingtechniquesinclude:(a)foot-paddling---agullstandinginwaterstampsitsfeetup anddownonthesubstratetouncoverorflushsmallinvertebrates(Buckley1966);and(b)skimming---thebirdrunsthroughshallowwaterwithitsbeakopenandpartiallyimmersedinthewater,seizingpreyoncontact(Tomkins 1964,Tolonen1970)(1).Ring-billedGullseatavarietyofvertebrates,asidefromfish,includingtheeggs(Courtney1979b)andyoung(Hunter1976inCourtneyandBlokpoel1980a)ofCommonTerns,duckeggs(Vermeer1970),aswellasyoungFranklin'sGulls,Black-crownedNight-Herons(Nycticoraxnycticorax)(Wolford1966inVermeer1970),Double-crestedCormorants(Phalacrocoraxauritus),AmericanAvocets(Recurvirostraamericana)(Bent1921),SpottedSandpipers(Actitismacularia)(BlokpoelandHaymes1979a),and Canada Geese(Brantacanadensis)(Munro1936).Passerinesreportedasfooditemsincludedwarblers(Oporornissp.)(BlokpoelandHaymes1979a),andpossiblyyoung Red-wingedBlackbirds phoeniceus)(JarvisandSouthern1976),andseveralspeciesofsparrows:SavannahSpar r.;; (Passerculussandwichensis),White-throatedSparrow(Zonotrichiaalbicollis),SongSparrow(Melospizamelodia),andChippingSparrow(Spizellapasserina)(BlokpoelandHaymes1979a).Ring-billedGullshavealsobeenfoundscavengingtheremainsofWesternMeadowlarks(Sturnellaneglecta)(Vermeer1970).Mammalseatenincludevoles(Microtusspp.) (Vermeer1970,JarvisandSouthern1976,Blokpoeland Haymes1979a),especiallymeadowvoles(Microtuspennsylvanicus)(Vermeer1970,BlokpoelandHaymes1979a),deermice(Peromyscusmaniculatus)(HaymesandBlokpoel1978b),Peromyscussp.(Rothweiler1960, Vermeer1970),housemice(Musmusculus)(Anderson1965),short-tailedshrews(Blarinabrevicauda)(Blokpoeland Haymes1979a),Richardson'sgroundsquirrel(Spermophilusrichardsonii)(Komarek1929,Munro1936),northernpocketgophers(Thomomystalpoides)(Vermeer1970),andrabbits(Leporidae)(Anderson1965).FewdetailedaccountsoffoodhabitsofRing-billedGullsareavailable,andweknowofnonefromthesoutheasternUnitedStates.Wesummarize someoftheimportantstudiesbyareabelow.AlbertaMunro(1936)examined37pelletsregurgitatedbygullsatBitternLakeinMay1933 andJune1934.Whilesomemayhavebeenregurgitated(1)Tolonentermedthisbehavior"ploughing",reservingtheterm"skimming"forsimilarmanueversconductedwhileinflight.164

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Ring-billedGullbyCaliforniaGulls(Laruscalifornicus),thevastmajoritywereundoubtedlyfromRing-billedGulls.Itemsoccurringmostfrequentlyinthepelletswerewheat(Triticumaestivum-in57.6%ofthepellets),miscellanousvegetabledebris(51.5%),oats(Avenasativa-27.3%),flypupae(Diptera-24.2%),cowhair(30.3%),miscellaneousinsects(15.2%),wildoats(Avenasp.-12.1%),andvoles(12.1%).Munrosuggestedthatthecowhairwasingestedwhendipteridpupaewerescavengedfromcarrion,andindicatedthatthemostimportantfooditems,inorderofimportance,weregrain,Richardson'sgroundsquirrels,carrion,groundbeetles(Carabidae),andmice[=voles,probablyMicrotussp.].Vermeer(1970)reportedthatRing-billedGullsatMiquelonLakein1965fedmostlyonplantmaterialinMay,insectsinJune,andrefuseinJuly.Plantsmade up68%ofthetotalvolumeoffoodforadultgullsinMaybutwerenoteateninJuly.Arthropodsmade up54%oftheadultdietand50%ofthedietofchicksinJune,butwereofnegligibleimportanceinJuly.Refusemade up76%and66%ofthetotalvolumeoffoodforadultsandchicks,respectively,inJuly,butmade up no morethan16%ofthedietforeitherageclassduringthetwoprecedingmonths. mammals,particularlymeadowvoles,werealsoimportant;theymade up 10-29%ofthetotalbulkofthedietforadultgullsfromMaythroughJuly,andwereevidentlyimportantinAprilaswell.Arthropods mostfrequentlyeatenweregroundbeetles(Carabidae)foundin48%of25esophagiandregurgitationsexamined,anddamselflynaiads(Coenagrionidae),foundin32%.Plantfoodstakenwerechieflygrains.Vermeer(1970)alsoexaminedfoodeatenatothercoloniesinAlberta,anddiscoveredthatrodentswereofconsiderableimportanceinsoutheasternAlbertabutwerereplacedbyfishinmorenorthernpartsoftheprovince. Ontario Haymes andBlokpoel(1978b)analyzed147samplescollectedatLeslieSpitinTorontoOuterHarborin1977.EachsampleconsistedofthefoodregurgitatedbyfiveRing-billedGullchickslessthan10daysold.Asreportedinthestudyby KirkhamandMorris(1979),mostofthefoodconsistedoffish,arthropods,andearthworms.Fishmade upabout50%ofthediet(byvolume)throughouttheseason.Arthropodswereasimportantasfishearlyintheseason,butbecamelessimportantlateron.Earthwormsweremostimportantduringthemid-andlatebreedingseason,whentheymade up23%and28%ofthediet,comparedtoonly8%earlierintheseason.Birds,mammals, andrefusealsowereingestedbutformedaninsignificantportionofthediet.Mostofthefood(77%)takenwasfromaquatichabitatsearlyintheseason,butfoodsofterrestrialoriginbecameprogressivelymoreimportantlateron and made up42%and51%ofthedietinmid-andlateseason,respectively.Thefishmostfrequentlyidentifiedwerealewife(Alosapseudoharengus)andrainbowsmelt(Osmerusmordax).Smeltweremoreimportantearlyinthebreedingseasonandalewivesweretakenmoreofteninmid-andlateseason.Smallnumbersofshiners(Notropissp.)andyellowperch(Pereaflavescens)werealsoidentified(HaymesandBlokpoel1978b).HaymesandBlokpoel(1978b)reportedthatatleast95familiesand13ordersofarthropodswererepresentedwithmidges(Chironomidae)andleaf-hoppers(Cicadellidae)makingup morethan50%oftheindividualinsectseatendur-165

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Ring-billedGullingtheseason.Non-insectarthropodseatenincludedspiders(Arachnida),centipedes(Chilopoda),millipedes(Diplopoda),andisopods(Crustacea).Kirkham andMorris(1979)reportedthatfoodsfedtoyoungRing-billedGullsatGullIslandinPresqu'ileProvincialParkduringthesummerof1977wereprimarilyfish,insects,andearthworms.Chicksuptofivedaysoldwerefedinsectsmorefrequentlythanolderchicks,butallageclasseswerefedfishmorefrequentlythananyotherfood.Fooditemsfedtochicksalsovariedwiththeprogressofthebreedingseason.InsectswerethedominantfooditeminMayandfishwerethesolefoodinJuly.EarthwormswerefedtotheyoungsporadicallythroughMayandJune,butmorefrequentlyafterperiodsofhighhumidityorrain.CaliforniaThestomachcontentsof29adultRing-billedGullscollectedinJuneandJuly1963atHoney LakeWaterfowlManagementAreainLassenCounty,consisted,byvolume,mainlyofinsect fragments (33.3%),meadowmice(Microtussp.-28.2%),garbage(9.9%),leavesofgrasses(Graminaea-7.6%),andfishbonesandremains(6.9%)(Anderson1965).MichiganLudwig(1966)foundthatfish,especiallyalewivesandsmelt,andinsectswerethetwo mostimportanttypesoffoodon Lakes HuronandMichiganin1963-1965(1).Alewiveswerefoundin70%ofthe372foodsamplesobtained,andsmeltwerefoundin18%ofthesamples.Sixotherspeciesoffish(yellowperch,gizzardshad[Dorosomacepedianum],brooksticklebacks(Culaeainconstans),ninespinesticklebacks[Pungitiuspungitius],rockbass[Ambloplitesrupestris],andwhitesuckers[CatostomuscommeFsoni])weretheotherspeciesoffishidentified.JarvisandSouthern(1976)analyzedthecontentsofregurgitationsobtainedatamainlandcolonyatRogersCityandattwoislandcoloniesinLakeMichiganand Lake Huronduringthebreedingseasonsin1964and1971.Agreatvarietyoffooditemswaseatenin1964,butfish(76%)andinsects(22%)predominated.InsectsweremostimportantinMayandJune,butfishmade up agreaterproportionofthevolumeofthefoodastheseasonprogressed.EarthwormswereimportantinMay,forming26%ofthefoodatthattime,butwereoflittleornoimportancelater.InsectsmostimportantinthedietatRogersCityin1964werecicadas(Okanaganarimosa,Cicadidae-19%byvolume),Ephemoptera(15%),andleaf-hoppers(Cicadellidae-12%).The mostimportantfishwereyellowperch(11%),ninespinesticklebacks(4%),andalewives(4%).Samplestakenin1971attheRogersCitycolony,acolonyatlIeauxGaletsinLakeMichigan,and acolonyonBirdIslandinLake Huron,revealedthatfish(75.5%)andinsects(ca.23.5%)againformedthebulkofthediet.Cicadas,mayflies(Ephemeroptera),andchironomidmidgeswerethemostimportantinsectfoods.Smelt,alewives,andsticklebackswerethemostimportantfish.Earthworms(Annelida)werealsoamajorsourceoffoodinMayforbirdsnestingattheRogersCitycolony.(1)SomeofLudWig'smaterialmayhavebeenobtainedontheCanadianportionofthesetwolakes.166

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Ring-billedGullMontanaTwenty-fivestomachscollectedinAugust1958andfromMaytoSeptember1959atFreezeoutLakecontainedvegetationand mostcontainedinsectremains;twostomachscontainedtheremainsofsmallmammals(Rothweiler1960).Rothweilerindicatedthat"wheat andbarley[Hordeumvulgare]kernelsandstemswerethemostcommonplantitemsfound"inthestomachsofthethreespeciesofgullsexamined.InsectsmostfrequentlyfoundinthestomachsofRing-billedGullsweregrasshoppers(Acrididae--foundin32%ofthestomachs)andbeetles(Carabidaein 32%. andCurculionidaein28%).Someflies(Tabanidae.Muscidae).moths(Phalaenidae).crickets(Oryllidae).andotherbeetles(Elateridae.Tenebrionidae.Scarabeidae)werealsoeaten.Earthworms(Oligochaetain32%)werealsofrequentlyfoundinthestomachs.IMPORTANTBIOLOGICALPARAMETERSEggLayingMosteggsarelaidinMaybutthetimingvariesslightlyfromcolonytocolony;birdsinsoutherncoloniesinitiatelayingearlierthanthoseinmorenorthernareas.In1971Ring-billedGullsatKaginawLake.Manitoba.laideggsfromthelasttwo weeksofMaythroughthefirsttwo weeksofJune.withapeak22-28May(Vermeer1973c).In1964 and 1965inAlberta.thepeakoflayingwasinearlyMay(Vermeer1970).In1977layingintheTorontoOuterHarborbeganinmid-AprilandcontinuedintoJuly(Haymes andBlokpoel1980).In1973 on LakeSuperior.layingoccurredfromearlyMaytoearlyJune.andpeakedinmid-May(Ryder1975).In1959 and1974-75inMontanaeggswerefirstlaidinthelastweekofAprilorthefirstweekofMay(Rothweiler1960.Baird1976).A numberoffactorsinfluencetheinitiationoflaying.Vermeer(1970)reportedthatadelayinlayingatcoloniesinAlbertawasrelatedtodecreasedmeanairtemperatureandsuggestedthathumandisturbancecanalsodelaytheonsetoflaying.Egglayingmayalsovaryfromareatoareawithinacolonyandwiththeageoftheparent.Ryder and Ryder(1981a)reportedthatpeaklayingin1977inthecenterofthecolonyatGraniteIslandinnorthernLakeSuperioroccurred6-10May.abouta weekearlierthanlayingattheperipheryofthecolony..Three-year-oldandolderbirdsatLeslieSpitin hadmeandatesofclutchinitiationfromthelastfivedaysofAprilthroughthefirstfivedaysofMay.Two-year-oldbirds.however.hada meandateofclutchinitiationabout21May;nestsofthelatterageclasswerealsofartherfromthecenterofthecolonythanwerethoseofolderbirds(HaymesandBlokpoel1980).MeanClutchSizeRing-billedGullslayonetofoureggs.butthemodalclutchsizereportedbymoststudiesisthreeeggs(Vermeer1970.Dexheimer andSouthern1974.Ryder1975.Baird1976. Ryder and Ryder1981a).OnetoeighteggshavebeenfoundinnestsinnorthernLakeSuperior(Ryderand Ryder1981a.Kovacs and Ryder1981).NestsatMiquelonLakeinAlbertacontained2-4eggsin1964 and1-5eggsin1965 (Vermeer1970).butthemajorityoffemaleslaidthreeeggs(Table22).NestsatcoloniesontheFreezeoutPeninsulaand ArodIsland.Montana.held1-6eggs(Baird1976).asdidthoseatHoneyLake.California(JohnstonandFoster1954).Ryder and Somppi(1979)andConoveretal.1979a)believethatthelargerclutches(5-8eggs)arelaidbypairsoffemales.167

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Ring-billedGullTable22.MeanclutchsizesreportedfortheRing-billedGull(a).Meanclutchsize2.922.85Numberofclutches265 435LocalityandyearofobservationALBERTAMiquelonLake,1964(3-86%) MiquelonLake,1965 (3-79%)ONTARIOSourceVermeer 1970 Vermeer 19702.80(b)147GraniteIsland,1973(3-56%) Ryder 19752.86(c)107GraniteIsland,1973 (3-55%) Ryder 19752.79(d)33GraniteIsland,1973(3-67%) Ryder 19752.00(e).7GraniteIsland,1973 (2-43%) Ryder 19753.19(f)325GraniteIsland,1977 Ryder and Ryder 1981a3.08(g)80GraniteIsland,1977 Ryder and Ryder 1981a2.86(h)204TorontoOuterHarbor,1977 (3-85%) HaymesandB1okpoe1 1978a2.88(i)9TorontoOuterHarbor,1977 Haymes and B1okpoe1 19802.95(j)76TorontoOuterHarbor,1977 HaymesandB1okpoe1 19802.78(k)23TorontoOuterHarbor,1977Haymesand B1okpoe1 19803.00(1)13TorontoOuterHarbor,1977 B1okpoe1 1980CALIFORNIA2.85(m) 717 HoneyLake,1953(3-71%)JohnstonandFoster1954MICHIGAN2.920 GreenIsland,1953Em1en19562.966700' Lakes Huron andMichigan,1960-65Ludwig 19662.8380RogersCity,1972Dexheimer andSouthern19742.73102 Thunder Bay, 1972 Dexheimer andSouthern19742.711680SouthManitouIsland,1972Shugart1977b2.725230SouthManitouIsland,1973Shugart1977b2.965400SouthManitouIsland,1974Shugart1977b2.745175SouthManitouIsland,1975Southerneta1.19802.835317RogersCity,1975Patton19792.583935SouthManitouIsland,1976Southerneta1.19802.796452RogersCity,1976Patton19792.892686SouthManitouIsland,1977Southerneta1.19802.837246RogersCity,1977Patton19792.612919 South ManitouIsland,1978Southerneta1.19802.694928RogersCity,1978Patton19792.531931SouthManitouIsland,1979Southerneta1.19802.68(n)5625RogersCity,1979SouthernandSouthern1982a 168

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Ring-billedGullTable22.Concluded.MeanclutchsizeNumberofclutchesLocalityandyearofobservationSourceMICHIGAN(CONT.)2.64(0)1415RogersCity,1979SouthernandSouthern1982a2.61(n)6495RogersCity,1980SouthernandSouthern1982a2.64(0)1242RogersCity,1980SouthernandSouthern1982aMONTANA2.63(p)2.07(p)2.41(p)6474376FreezeoutPeninsula,1973(3-62%)FreezeoutPeninsula,1974 (3-38%) ArodIsland,i974(2-42%)Baird1976Baird1976Baird1976(a)Someofthesefiguresarederivedfromcountsofcontentsofnestsmadeduringshort-termvisits.They donotadequatelyrepresentclutchsizeforthepopulationsincesomeproportionofthebirdsmaystillhavebeenlayingeggsandothernestsmayhavelosteggsbeforethecolonywasvisitedbytheobservers.Inyetothernests,noeggsmayhaveeverbeenlaid(Ryder1976).Wherepossible,welistedin parentheses themostfrequentlyobservedclutchsizeandthepercentageofneststhatcontainedthisnumberofeggs.(b)ClutchsizeforallpairscalculatedfromdatainTable2ofcitedsource."Clutches"of5,6,and 7areexcludedfromthesefiguresandforthenextthreefigureslisted.(c)Dataforpairsinwhichbothbirdshadmatureplumage.(d)Dataforpairsinwhichonebirdhadimmatureplumage.(e)Dataforpairsinwhichbothbirdshadimmatureplumage.(f)Clutchsizeatnestsincenterofcolony.(g)Clutchsizeatnestsatperipheryofcolony.(h)Clutchsizeforallnestsstudied.(i)Earlynests(egglayingbegun onorbefore1May)withparents3yearsoldoryounger.(j)Earlynestswithatleastoneparent4yearsoldorolder.(k)Latenests(egglayingbegunafter1May)withatleastoneparent3yearsoldoryounger.(1)Latenestswithatleastoneparent4yearsoldorolder.(m)Figuresarederivedfromasinglecountofnestcontentsandincludeneststhatcontainedeggsandyoung,oryoung.(n)Clutchsizeatnestsinunalteredbreedinghabitats.(0)Clutchsizeatnestsinareathatwascoveredwithfillandgradedduringthefallof1979.(p)Figuresforpercentofmodalclutchsizeareapproximatedfromfigures5-1and5-2ofthecitedsource.169

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Ring-billedGullMeanclutchsizesreportedforRing-billedGullsvaryfromyeartoyearbothwithinandbetweencolonies(Table22).Ryder and Ryder(1981a)reportedthatthemeanclutchsizeattheperipheryofthecolonyisnotsignificantlydifferentfromthemeanclutchsizeinthecenter.SouthernandSouthern(1982a)reportedthatmeanclutchsizedidnotdifferbetweennestsfromareasthathadbeenbulldozedandareasinunalterednestinghabitat.Meanclutchsizeincreaseswiththeageoftheparentuptoaboutfouryears(Haymes andBlokpoel1980,Table22),suggestingthatnestsfartherfromthecolonycenterhavesmallerclutches.Ryder(1975)alsoreportedthatlargerclutchesarelaidbyolderbirds.IncubationPeriodVermeer(1970)reportedthatincubationofthethirdeggin64clutchestook23-28days(mean=25.0)inManitobain1964.Allbutthreeeggstook24to26daystohatch.Duringthefollowingyear,whenthecolonywasdisturbedmore by humansandnocturnalpredators,theonlytwoincubationperiodsrecordedwere28and29days.Baird(1976)reportedarangeofincubationperiodsof26-27daysforRing-billedGullsnestinginMontana.HatchingSuccessRatesofhatchingsuccessvarywidely(Table23) and depend on a numberoffactors.Hatchingsuccessdecreasedasthenestingseasonprogressedin1977atGraniteIsland(Ryderand Ryder1981a).Three-eggclutcheshadthehighesthatchingsuccessatGraniteIslandinboth1973 (Ryder 1975) and 1977(Ryderand Ryder1981a),andatArod Lake andtheFreezeoutPeninsula,Montana,in1974(Baird1976).Hatchingsuccessdecreasedinlatenestsin1977atLeslieSpitintheTorontoOuterHarbor,butincreasedwiththeageoftheparentstoamaximumof90.9%for5-year-oldbirds(Haymes andBlokpoel1980).ThisisconsistentwithRyder's(1975) report thathatchingsuccessishigherinpairsofmature-plumagedbirdsthaninpairsinwhichatleastonebirdhasimmatureplumage.Inanotherstudy(Ryder1976),moreeggshatchedinnestsbegunbyadultsearlyinthenestingseasonthaninnestsstartedlater,butRydernotedthatthisrelationshipdoesnotholdfornestsinitiatedby youngpairs.Dexheimer andSouthern(1974)reportedhigherhatchingsuccessfornestsinthecenterofthecolonythanfornestsattheperipheryfortwocoloniesinMichigan(Table23),butRyderand Ryder(1981a)found nodifferenceinhatchingsuccessforperipheralandcentralnestsatGraniteIsland.Hatchingsuccessinsupernormalclutches(thosewith5to8eggs)ismuchlowerthaninnestswithnormalclutches.HatchingsuccessatGraniteIslandin1978for37supernormalclutcheswasonly8.2%(Ryderand Somppi1979),comparedtoahatchingsuccessof80-90%fornormalclutches(Somppi 1978inRyder and Somppi1979).Conoveretal.(1979a)reportedthatfertilityinclutchesof5to6eggsatthreecoloniesinWashingtonwas muchlower(65.9% and 69.9%,respectively)thaninsmallerclutches(ca.90%) .AgeatFledgingVermeer(1970)reportedanaverageageatfirstflightas37.2days(range34-41).MeanageatfledgingatGullIsland,Ontario,wassimilar(36.4+2.36days;n=28) (KirkhamandMorris1979).170

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Ring-billedGullTable23.RatesofhatchingsuccessreportedfortheRing-billedGull(a).PercentofeggslaidNumberthatofhatchedeggsLocalityandyearofobservationSource86 MiquelonLake,1964 Vermeer 197043.4(b)412GraniteIsland,1973 Ryder 197551.6(b)306GraniteIsland,1973 Ryder 197522.8(b)92GraniteIsland,1973 Ryder 19750.0(b)14GraniteIsland,1973 Ryder 197559.5(c)1037GraniteIsland,1977 Ryder and Ryder 1981a58.5(c)246GraniteIsland,1977 Ryder and Ryder 1981a48.1-89.5(d)TorontoOuterHarbor,1977 HaymesandBlokpoel1978a96.2(e)26TorontoOuterHarbor,1977HaymesandBlokpoel198088.4(f)224TorontoOuterHarbor,1977 HaymesandBlokpoel198073.4(g)64TorontoOuterHarbor,1977HaymesandBlokpoel198084.6(h)39TorontoOuterHarbor,1977 HaymesandBlokpoel198091Lakes Huron andMichigan,1960-65Ludwig 196648.6(i)288 LakeHuron,1972 Dexheimer andSouthern197476.5(j)230 Lake Huron, 1972 Dexheimer andSouthern197464.4149SouthManitouIsland,1972 Shugart;, 1977b73.3375SouthManitouIsland,1973Shugart1977b70.1364SouthManitouIsland,1974Shugart1977b60.2(k)14174SouthManitouIsland,1975Southernetal.198015.0(k)10163SouthManitouIsland,1976Southernetal.198061.57756SouthManitouIsland,1977Southernetal.198037.0(k)7629SouthManitouIsland,1978Southernetal.198018.1(k)4591SouthManitouIsland,1979Southernetal.1980(a)Numberoeggsiscalculatedfromthenumberofnestsand meanclutchsizeoreggsperpairforallpapersexceptthoseby Dexheimer andSouthern(1974)andSouthernetal.(1980).MiquelonLakeisinAlberta,GraniteIslandandtheTorontoOuterHarborinOntario,andSouthManitouIslandinMichigan.(b)HatchingsuccessforallpairsascalculatedfromTable2ofcitedsource.Datafor"clutches"of5,6,and 7areexcludedfromthefigures.Firstisanoverallaverage;thenextthreefigureslistedare,respectively,forpairsinwhichbothbirdshadmatureplumage,onehadimmatureplumage,andbothhadimmatureplumage.(c)Thefirstfigureisfornestsatthecenterofthecolony;thesecondfornestsattheperiphery.(d)Rangeofvaluesforeightstudyplots.(e)Earlynestswithatleastoneparent3yearsoldoryounger.(f)Earlynestswithatleastoneparent4yearsoldorolder.(g)Latenestswithatleastoneparent3yearsoldoryounger.171

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Ring-billedGullTable23.Concluded.(h)Latenestswithatleastoneparent4years old orolder.(i)DataforfringenestsattwocoloniescombinedfromTable1ofcitedsource.(j)DataforcentralnestsattwocoloniescombinedfromTable1ofcitedsource.(k)Thesefiguresareforyearsinwhichthecolonywasaffectedbypredationbyresidentredfoxes(Vulpesvulpes);thecolonywasnotvisitedbyfoxesduringincubationin1977.FledgingSuccessFledgingsuccess,likehatchingsuccess,varieswidely.In1964, 34.3%oftheeggslaidresultedinfledgedyoungatcoloniesinAlberta,butno youngfledgedthefollowingyear(Vermeer1970).Vermeer(1970)alsoreportedthatlateclutchesfledgefeweryoungthanearlyones.HaymesandBlokpoel(1978a)reportedfledgingsuccessesof22.6-31.8%peregglaidin1977fortwostudyplotsattheTorontoOuterHarborwhereRing-billedGullsnestwithCommonTerns.AtfourstudyplotsoccupiedonlybyRing-billedGulls,fledgingsuccessperegglaidrangedfrom 11.1%to57.9%.SurvivalofyoungatacolonyatRogersCity,Michigan,wasnotsignificantlydifferentfornestsatthecenterandperipheryofthecolony(Dexheimer andSouthern(1974);atacolonyinThunder Bay,survivalofyoung wassignificantlygreater.atcentralnests.ThedifferenceinsurvivalattheThunder BaycolonymayhavebeenbecausethiscolonywassubjecttofloodingwhereasthecolonyatRogersCitywasnot(DexheimerandSouthern1974).Figuresforproductivity(Table24)revealsimilarvariationfromareatoareaandfromcolonytocolony.MortalityofEggsandYoung Vermeer(1970)reportedthat13.8% and 84.0%oftheeggsfailedtohatchatcoloniesinAlbertain1964 and 1965,respectively.Inbothyears,infertilityorembryonicdeathandthedisappearanceofeggsaccountedformostofthelosteggs.Vermeerattributedthemuchgreaterloss'in1965toacombinationofbadweatherandincreasednocturnalpredation.Predationhasbeenreportedasamajorcauseofegglosselsewhereaswell.Emlenetale(1966)reportedthatnocturnaldisturbancebyaraccoon(Procyonlotor)indirectlyledtothelossofmanyeggsandsmallyoungatRogersCity,Michigan.Redfox(Vulpesvulpes)predationonthegullcolonyonSouthManitouIsland,Michigan,reducedhatchingratesforRing-billedGullsto60%,15%,37%,and18%in1975,1976,1978,and1979,respectively,and noorveryfew youngfledged(Southernetale1980).Redfoxesalsoreducedchickproductionatthiscolonyin1974(Shugart1977b).Mostofthereductioninnestingsuccesswasnotattributedtopredationbyfoxesforfood,butmortalityapparentlyresultedeitherfromeggschillingwhengullsfledtheirnests,orasaresultofthefoxeskillingmorepreythantheycouldeat.NocturnaldisturbancebyGreatHorned Owls (Bubovirginianus)resultedinchickdeathsandineggsbeingknockedoutofnestsatRogersCityin1978(SouthernandSouthern1979).Anestimated18%oftheeggsin841nestsfailedtoproduceyoungasaresultofthisdisturbance.Aswiththefoxpredation,the172

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Ring-billedGullTable24.ProductivityinRing-billedGullcolonies.Numberofyoungproducedperpair(ornest)1.01.03(a)1.06(b)0.27-1.76(c)0.64-1.20(d)LocalityandyearofobservationALBERTAMiquelonLake,1964ONTARIOGraniteIsland,1977GraniteIsland,1977TorontoOuter'Harbor,1977TorontoOuterHarbor,1977MICHIGANSourceVermeer 1970 Ryder and Ryder 1981a Ryder and Ryder 1981aHaymesandBlokpoel1978a Haymes andBlokpoel1978a1.740.671.891.211.830.681.561.651.360.07 0.011.540.000.00(a)(b)(a)(b)(e)(e) (e) (e)LakesHuron,Michigan,1960-65GreenIsland,1952RogersCity,1972RogersCity,1972 Thunder Bay, 1972 Thunder Bay, 1972SouthManitouIsland,1972SouthManitouIsland,1973SouthManitouIsland,1974SouthManitouIsland,1975SouthManitouIsland,1976SouthManitouIsland,1977SouthManitouIsland,1978SouthManitouIsland,1979 Ludwig 1966 Emlen 1956 Dexheimer andSouthern1974 Dexheimer andSouthern1974 Dexheimer andSouthern1974 Dexheimer andSouthern1974Schugart1977bSchugart1977bSchugart1977bSouthernetal.1980Southernetal.1980Southernetal.1980Southernetal.1980Southernetal.1980(a)Nestsatthecenterofthecolony.(b)Nestsattheperipheryofthecolony.(c)RangeofvaluesforfourstudyplotsoccupiedonlybyRing-billedGulls.(d)RangeofvaluesfortwostudyplotsoccupiedbyRing-billedGullsandCommonTerns.(e)Thesefiguresareforyearswhenproductionwasaffectedbyaresidentpopulationofredfoxesand waslessthan0.1in1978 and 1979..directeffectofpredationbytheGreatHornedOwlwas muchlessthantheindirecteffect.173

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Ring-billedGullOthersuggestedorknown mammalianpredatorsofeggsoryoungincludecoyotes(Canislatrans)(Vermeer 1970,Baird1976),long-tailedweasels(Mustelafrenata)(Baird1976),mink(Mustelavison) (9onover andMiller1979),andstripedskunks(Mephitismephitis)(SouthernandSouthern1979).Commongartersnakes(Thamnophissirtalis)havebeenobservedseizingandeatingnewlyhatchedchicksatMugg'sIslandinTorontoHarbor(Fetterolf1979a),butpredationbysnakesisuncommon. Vermeer(1970)listedseveralsourcesofmortalityforRing-billedGullchicks.Disappearanceforunknownreasonsaccountedfor58%and63%ofthechickmortalityatMiquelonLake,Alberta,in1964 and 1965,respectively.TheprincipalknowncauseoflossofchickswasattacksbyneighboringRingbilledGullsandpredationbyCaliforniaGulls(Laruscalifornicus).OtherminorformsoflosswereknownorsuspectedpredationbyGreatHorned Owls (Bubovirginianus)andRed-tailedHawks(Buteojamaicensis),deathfromchillingfollowingstormsordisturbance,anddisease.Emlen(1956)reportedthatmostmortalityofchicksin1952atacolonyon GreenIsland,Michiganwastheresultofattacksbyneighboringadults;someyoungwerealsolosttopredationbyHerringGulls.FoxeskilledmorechicksthantheyateduringvisitstogullcoloniesatSouthManitouIsland,Michigan,butchillingwasthemajorcauseoffox-relatedmortality(Patton1979).Humandisturbanceofgullcoloniescan.beaprimarysourceofnestingfailureinRing-billedGulls(Burger1981a).Fetterolf(1981),inapreliminaryreportonreproductivesuccessinminimallydisturbedRing-billedGullcolonies,notedthatyoungfledgedatarateof2.10to2.53perpair,arateconsiderablyhigherthananylistedinTable24.RenestingSomeRing-billedGullsrelayafterlossofanest.Theproportionthatdoessoisnotknownbutisprobablysmall.Vermeer (1970) foundthatonlyoneof154pairsofRing-billedGullsrelaidwhentheirclutcheswereremovedduringthepeakoflaying.AgeatFirstBreedingLudWig(1966)reportedthatnearlyhalfofthetwoyear-oldbirdSatcoloniesintheGreatLakesbreedandthatallRing-billedGullsbreedbeforetheirfourthyear.Southern(1977)reportedthatmostRingbilledGullsatastablecolonynearRogersCity,Michiganbreedforthefirsttimeasthree-year-olds.Ludwig'sobservationsweremadeduringaperiodofrapidpopulationgrowth;mostRing-billedGullsprobablywaituntilagethreeinstableorspace-limitedcolonies.MaximumNaturalLongevityA youngbirdbandedinOntariowas found deadinthesamearea25yearslater(Clappetal.1982a).WeightWeightsofbreedingadultRing-billedGullsvaryfromabout375650 g(0.8-1.4lb);malesareheavierthanfemales(Table25).174

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Ring-billedGullTable25.Weights(ingrams)ofRing-billedGulls(a)Meanweight533 566511499 46446347143237735.5 38.5Range482-650 379-563 260-50334.3-37Number weighed Sample andseason19ad.males,summer48ad.males,May-June 1ad.male,May1ad.male,Feb.1imm.male,Feb.20ad.females,summer51ad.females,May-June 1imm.female,Feb.30fledgingyoung 3ca.day-oldyoung39newlyhatchedyoungAreaAlbertaOntarioCaliforniaAlabama AlabamaAlbertaOntarioAlabamaAlbertaCaliforniaAlbertaSourceVermeer1970Ryder1978Moffitt1942StewartandSkinner1967StewartandSkinner1967Vermeer1970Ryder1978StewartandSkinner1967Vermeer1970Moffitt1942Vermeer197034.626.6-42.6newlyhatchedyoungMichiganDawsonetal.197653.9 53.446.5-61.42445.3-61.528fresheggsfresheggsOntarioOntarioRyderetal.1977Ryderetal.1977(a)Rangefiguresfrom Dawsonetal.(1976),Ryderetal.(1977)andRyder(1978)arethemean+2SO.Thefirstsetoffiguresfrom Ryderetal.(1977)isforthree-egg-clutchesfromthecenterofthecolony;thesecondisforthree-eggclutchesattheperiphery.SUSCEPTIBILITYTOOILPOLLUTIONRing-billedGullsarenotparticularlysusceptibletooiling,butreportsintheBirdBandingLaboratory,Laurel,Maryland,suggestdeathasaresultofoilinginNorthandSouthCarolina,Florida,andLouisiana,andalsoinmorenorthernlocalities.Oil-smearedRing-billedGullsarefairlycommonat175

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Ring-billedGullgarbagedumpsinFlorida(W.Hoffman,pers.comm.),buttheyprobablycomeintocontactwithwasteoilinthedumps. KingandSanger(1979)consideredthisspeciesasonewithlowriskfromoilinginthePacificNorthwesternstates.AmajorityofthepopulationwintersinthesoutheasternUnitedStates, but theydonotoccuroffshoreinanygreatabundance;theyusuallyroostonlandratherthanonthewater,andtendtooccurinlandwithgreaterfrequencythanothergulls.Consequently,thisspeciesisprobablynotatanygreathazardfromdevelopmentofpetroleumresourcesinthesoutheastandtheeffectofoffshoreoilspillsonthespecieswillbenegligible.BIBLIOGRAPHY1983Southern,L.K.andW. E. Southern.1983.ResponsesofRing-billedGullstocannon-nettingandwing-tagging.J.Wildl.Manage.47:234-237.1982Blokpoel,H.andP.A.Courtney.1982.ImmigrationandrecruitmentofRingbilledGullsandCommonTernsonthelowerGreatLakes.Can.Wildl.ServoProg.Notes133.12pp.Blokpoel,H.andD.V.Weseloh.1982.StatusofcolonialwaterbirdsnestingonLittleGallooIsland,LakeOntario.Kingbird32:149-157.Bos, E. andP.deHeer.1982.Ring-billedGullsonCanaryIslandsinMarch1982.DutchBirding4:90-91.Cade,M.1982.PlumagevariabilityofimmatureCommonandRing-billedGulls.Brit.Birds75:580.Cooper,C.R.,A.P.Gilman andH.Blokpoel.1982.Ring-billedGullsnestingon OttawaRivernearOttawa.OntoFieldBioI.36:11-15.Hayward,J.L.1982a.Effectsofnesthabitat,behavior,andvolcanicashonreproductivesuccessinRing-billedandCaliforniaGulls.Ph.D.thesis,Univ.Washington/Pullman,WA.1982b.Asimpleegg-markingtechnique.J.FieldOrnithol.53:173.Hayward,J.L.,D.E.MillerandC.R.Hill.1982.MountSt.HelensAsh:itsimpactonbreedingRing-billedandCaliforniaGulls.Auk99:623631.Heinz,F.A.tonest177.1982.Growth anddevelopmentofRing-billedGullchicksrelatedplacementinacolony.(Abstractonly.)ColonialWaterbirds5: 176

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Ring-billedGullHoogendoorn,W.1982.Ring-billedGullinMoroccoinAugust1982.DutchBirding4:91-92.King,B.1982.AmericanCootandRing-billedGullfeedingonduckdroppings. Fla. FieldNat.10:80.Mullarney,K.1982.Mysteryphotographs.[Ring-billedGull.]DutchBirding4:20-22.Southern,L.K.andW.E.Southern.1982a.EffectofhabitatdecimationonRing-billedGullcolony-andnest-sitetenacity.Auk99:328-331.1982b.MatefidelityinRing-billedGulls.J.FieldOrnithol.53:170--171.1982c.Short-termeffectsofcannon-nettingandwing-taggingonbreed--ingRing-billedGulls.(Abstractonly.)ColonialWaterbirds5:179.Southern,L.K.,S.R.Patton,andW.E.Southern.1982.NocturnalpredationonLarusgulls.ColonialWaterbirds5:169-172.Southern,W.E.andL.K.Southern.1982.IntensificationofadultRingbilledGullaggressionduringreproductionanditspossibleconsequences.ColonialWaterbirds5:2-10.Wong,P.L.andR. C.Anderson.1982a.RedescriptionofPectinospiruraargentataWehr,1933(Nematoda:Acuariodea)fromLarusdelawarensisOrd(Laridae).Can.J.Zoo1.60:1940-1944.1982b.ThetransmissionanddevelopmentofCosmocephalusobvelatus--(Nematoda:Acuarioidea)ofgulls(Laridae).Can.J.Zoo1.60:1426-1440.1981Burger,J.1981a.Effectsofhumandisturbanceoncolonialspecies,particularlygulls.ColonialWaterbirds4:28-36.Burger,J.andM.Gochfeld.1981a.Age-relateddifferencesinpiracybehaviouroffourspeciesofgulls,Larus.Behaviour77:242-267.Conover,M.R.andD.O.Conover.1981.AdocumentedhistoryofRing-billedGullandCaliforniaGullcoloniesinthewesternUnitedStates.ColonialWaterbirds4:37-43.Conover,M.R.andD.E.Miller.1981.ElicitationofvocalizationsandpeckinginRing-billedGullchicks.Behaviour77:268-286.Fetterolf,P.M.1981.ReproductiveperformanceofminimallydisturbedRingbilledGulls.(Abstractonly.)ColonialWaterbirds4:68.177

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Ring-billedGullKing,B.1981.CaspianTernssun-bathingwithRing-billedGulls.Brit.Birds74:181.Kovacs,K.M.andJ.P.Ryder.1981.Nestsitetenacityandmate fidelity infemale-femalepairsofRing-billedGulls.Auk98:625-627.Petokas,P.J.1981.SnappingTurtlepredationonRing-billedGulls.Bull.Md.Herpetol.Soc.17:111-115.Ryder,P.L.andJ.P.Ryder.1981a.ReproductiveperformanceofRing-billedGullsinrelationtonestlocation.Condor83:57-60.1981b.Minimuminter-nestdistance,isitausefulparamaterinpre--dietingbreedingsuccess?(Abstractonly.)ColonialWaterbirds4:200.Scharf,W.C.1981.Thesignificanceofdeterioratingman-madeislandhabitatstoCommonTernsandRing-billedGullsintheSt.Mary'sRiver,Michigan.ColonialWaterbirds4:15.5-159.Southern,L.K.1981.Sex-relateddifferencesinterritorialaggressionbyRing-billedGulls.Auk98:179-181.Southern,L.K.andW. g. Southern.1981.TheinfluenceofhabitatalterationsonsitetenacityinRing-billedGulls.(Abstractonly.)ColonialWaterbirds4:201.Southern,W.E.andL.K.Southern.1981.Colonycensus.resultsasindicatorsofpre-hatchingperturbations.ColonialWaterbirds4:143-149.Weber,J.W.1981.TheLarusgullsofthePacificNorthwest'sinterior,withtaxonomiccomments forms.(PartII--conclusion).ContinentalBirdlife2:74-91.Weseloh,D. V.1981.AprobableFranklin'sXRing-billedGullpairnestinginAlberta.Can.Field-Nat.95:474-476.Weseloh,D.V.C.andM.T.Myres.1981.SeasonalfluctuationsingullnumbersatCalgary,Alberta,Canadaandamodelforestimatingtheirnumbers.ColonialWaterbirds4:132-137.1980Blokpoel,H.1980.StatusoftheRing-billedGullontheCanadianLowerGreatLakes.(Abstractonly.)Pac.SeabirdGroupBull.7:57.Blokpoel,H.andP.Courtney.1980.Sitetenaci 1:Y inanewRing-billedGullcolony.J. Field Ornithol.51:1-5.Bull,J.1980.NotesonaRing-billedGull.Linn.Newsl.34:3.178

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Ring-billedGullBurger,J.,M.Fitch,G.Shugart,andW.Werther.1980.PiracyinLarusgullsata dumpinNewJersey.Proc.1979Conf.ColonialWaterbirdGroup3:87-98.Conover,M. R. andD.E.Miller.1980.DailyactivitypatternsofbreedingRing-billedandCaliforniaGulls.J.FieldOrnithol.51:329-339.Conover,M. R.,R. Klopfer, 1<". DudleyandD.E.Miller.1980.StimulusfeaturesofchicksandotherfactorsevokingparentalprotectivebehaviourinRing-billedGulls.Anim.Behav.28:29-41.Courtney,P.A.andH.Blokpoel.1980a.BehaviorofCommonTernsnestingnearRing-billedGulls.Can.Field-Nat.94:336-338.Evans, R. M.1980.DevelopmentofindividualrecognitioninyoungRing-billedGulls(Larusdelawarensis):aneffectoffeeding.Anim.Behav.28:60-67.Grace,J.W.1980.KleptoparasitismbyRing-billedGullsofwinteringwaterfowl.WilsonBull.92:246-248.Haymes,G.T.andH.Blokpoel.1980.TheinfluenceofageonthebreedingbiologyofRing-billedGulls.WilsonBull.92:221-228.Koonz,W.H.1980.LargeclutchesofRing-billedGullsinManitoba.BlueJay38:46-47.Lauro,A.J.andB.J.Spencer.1980.AmethodforseparatingjuvenalandfirstwinterRing-billedGulls(Larusdelawarensis)andCommonGulls(Larus Am.Birds34:111-117.Mullarney,K.1980.Ring-billedGullsinCo.Mayo.AspeciesnewtoIreland.IrishBirds1:517-525.Ryder,J.P.1980.Apreliminaryinvestigationofthesecondarysex-ratioofRing-billedGullsonGraniteIsland,1978-79.(Abstractonly.)Proc.1979Conf.ColonialWaterbirdGroup3:256.Southern,L.K.andW.E.Southern.1980.PhilopatryinRing-billedGulls.Proc.1979Conf.ColonialWaterbirdGroup3:27-32.Southern,W.E.1980a.ComparativedistributionandorientationofNorthAmericangulls.Pp.449-498inJ.Burger,B.C.OllaandH.E.Winn(eds.).BehaviorofMarineAnimals.Vol.4:MarineBirds.PlenumPressNY.xviiand515pp.__1980b.Theimpactoffoxpredationongullbreedingsuccess,SleepingBearDunesNationalLakeshore.Pp.111-129inProc.2ndConi.Sci. Res. Natl.Parks,Vol.12:TerrestrialBiology:Zoology.Natl.ParkServ.,Washington,D.C.179

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Ring-billedGullSouthern,W.E.,S.R.PattonandL.A.Hanners.1980.DifferentialresponseofRing-billedGullsandHerringGullstofoxpredation.Proc.1979Conf.ColonialWaterbirdGroup3:119-127.1979Blokpoel,H.andG.T.Haymes.1979a.SmallmammalsandbirdsasfooditemsofRing-billedGullsontheLowerGreatLakes.WilsonBull.91:623-625.1979b.OriginsofRing-billedGullsatanewcolony.Bird-Banding50:--210-215.Burger,J.andM.Gochfeld.1979.AgedifferencesinRing-billedGullkleptoparasitismonStarlings.Auk96:806-808.Conover,M.R.andD.E.Miller.1979.ReactionofRing-billedGullstopredatorsanddisturbancesattheirbreedingcolonies.Proc.1978Conf.ColonialWaterbirdGroup2:41-47.Conover,M.R.,D.E.MillerandG.L.Hunt,Jr.1979a.Female-femalepairsandotherunusualreproductiveassociationsinRing-billedandCaliforniaGulls.Auk96:6-9.Conover,M.R.,B.C.Thompson,R.E.FitznerandD.E.Miller.1979b.IncreasingpopulationsofRing-billedandCaliforniaGullsinWashingtonState.West.Birds10:31-36.Courtney,P.A.1979b.EffectsofarabbitonnestingCommonTerns.Can.J.Zoo1.57:2457-2460.Fetterolf,P.M.1979a.CommonGarterSnakepredationonRing-billedGullchicks.Can.Field-Nat.93:317-318.1979b.NocturnalbehaviourofRing-billedGullsduringtheearlyincu--bationperiod.Can.J.Zoo1.57:1190-1195.Grant.G.S.1979.Ring-billedGullsfeedingondatefruits.Condor81:432433.Grant,P.J.1979.FieldidentificationofwestPalearcticgulls.Part2.Common,Mediterranean,Ring-billed,LaughingandF.ranklinIsGulls.Brit.Birds72:142-182.Hogg,R.H.1979.Ring-billedGullinAberdeenshire.Scott.Birds10:275276.Kirkham, 1. R.andR.D.Morris.1979.FeedingecologyofRing-billedGull(Larusdelawarensis)chicks.Can.J.Zoo1.57:1086-1090.180

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Ring-billedGullMiller,D.E.andM.R.Conover.1979.Differentialeffectsofchickvocalizationsandbill-peckingonparentalbehaviorintheRing-billedGull.Auk96:284-295.Nunnally,S.,D.Nunnally,"R.NeedhamandR.Lennon.1979.Nocturnalfeedingofgullsatalightedpier.Chat43:63.Patton,S.R.1979.ShortandlongtermeffectsofRedFoxnocturnalpredationonthenestingsuccessofcolonialbreedingHerringandRing-billedGulls.M.S.thesis,N.Ill.Univ./Dekalb,IL.viand107pp.Ryder,J.P.1979.ingingulls.Possibleoriginsandadaptivevalueoffemale-femalepairProc.1978ConLColonialWaterbirdGroup2:138-145.Ryder,J.P.andP.L.Somppi.1979.Female-femalepairinginRing-billedGulls.Auk96:1-5.Southern,L.K.andW.E.Southern.1979.Absenceofnocturnalpredatordefensemechanismsinbreedinggulls.Proc.1978Conf.ColonialWaterbirdGroup2:157-162.Weseloh,D.V.andH.Blokpoel.1979.HinterlandWho'sWho:Ring-billedGull.Environ.Can.Wildl.ServoCat.CW69-4/63.4pp.1978Bishop,M.M.1978.ImprintingofaRing-billedGull.Auk95:196-197.Blokpoel,H.andp.M.Fetterolf.1978.ColonizationbygullsandternsoftheEasternHeadland,TorontoOuterHarbour.Bird-Banding49:59-65.Blokpoel,H.andG.B.McKeating.1978.ianLakeErieandadjacentwaters.12pp.Fish-eatingbirdsnestinginCanadCan.Wildl.Serv.Prog.Notes87.Brackbill,H.1978.PlaybyRing-billedGulls.Bird-Banding49:282-283.Chardine,J.W.1978.SeasonalvariationinthereproductivebiologyoftheRing-billedGull(Larusdelawarensis).M.S.thesis,BrockUniv./St.Catharines,ON.102pp.Conover,M.R.andD.E.Miller.1978.Acousticalpropertiesoftheswoopand-soarcalloftheRing-billedGull.Auk95:599-602.Elston,S.F.,C. D. RymalandW.E.Southern.1978.IntraspecifickleptoparasitisminbreedingRing-billedGulls.Proc.1977ConLColonialWaterbirdGroup1:102-109.181

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Ring-billedGullHaymes,G.T.andH.Blokpoel.1978a.ReproductivesuccessoflaridsnestingontheEasternHeadlandoftheTorontoOuterHarbourin1977.Ont.FieldBioI.32:1-17.1978b.FoodofRingO-billedGullchicksattheEasternHeadlandof--theTorontoOuterHarbourin1977.Can.Field-Nat.92:392-395.Kinkel,L.K.andW.E.Southern.1978.AdultfemaleRing-billedGullssexuallymolestjuveniles.Bird-Banding49:184-186.Moulton,M.andM.Ports.1978.PiracybyRing-billedGullonCommonGolden-oeye.Bull.KansasOrnithol.Soc.29:18.Patton,S.R.andW.E.Southern.1978.TheeffectofnocturnalRedFoxpredationonthenestingsuccessofcolonialgulls.Proc.1977.Conf.ColonialWaterbirdGroup1:91-101.Ryder,J.P.1978.SexingRing-billedGullsexternally.Bird-Banding49:218-222.Ryder,R.A.1978.GullsinColorado:theirdistribution,status,andmovements.Proc.1977Conf.ColonialWaterbirdGroup1:3-9.Somppi,p.L.1978.ReproductiveperformanceofRing-billedGullsinrelationtonestlocation.M.S.thesis,LakeheadUniv./ThunderBay,ON.Southern,W.E.1978a.Ring-billedGullpairwith2nests.WilsonBull.90:299-301.1978b.OrientationresponsesofRing-billedGullchicks:are-evalua---tion.Pp.311-317inK.Schmidt-KoenigandW.T.Keeton,(eds.).Pro-ceedingsinLifeSciences,AnimalMigration,NavigationandHoming.Springer-Verlag,Berlin.Weseloh,D.V.,S.BrechtelandR.D.Burns.1978.RecentpopulationchangesinDouble-crestedCormorantsandCaliforniaandRing-billedGullsinAlberta,Canada,withanoteonWhitePelicans.Proc.1977Conf.ColonialWaterbirdGroup1:10-18.1977Allen,L.J.1977.ThecomparativefeedingecologyrelativetopopulationdynamicsoftheHerringandRing-billedGulls(LarusargentatusandL.delawarensis)andsomecomparisonswiththeCaspianTern.M.S.thesisQueensUniv./Kingston,Ontario.196pp.Blokpoel,H.1977.GullsandternsnestinginnorthernLakeOntarioandtheupperSt.LawrenceRiver.Can.Wildl.ServoProg.Notes75.12pp.182

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Ring-billed Gull Evans,R.M.1977.Auditorydiscrimination-learninginyoungRing-billedGulls(Larusdelawarensis).Anim.Behav.25:140-146.Fetterolf,P.M.andH.Blokpoel.1977.TernsandgullsnestingonToronto'seasternheadland. Out.. FieldBioI.31:51-52.Gollop,J.B.1977.WhiteRing-billedGullatSaskatoon.BlueJay35:166.Hall,R.F.,D.G.Waldhalm,W.A.MeinershagenandD.A.Dubose.1977.IsolationofSalmonellaspp.fromdeadgulls(LaruscalifornicusandLarusdelawarensis)fromanIdahoirrigationreservoir.AvianDis.21:452-454.Lauro,A.J.1977.Gullpredationonanantswarm.Kingbird27:87-88.Penland,S.T.andS.J.Jeffries.1977.NewbreedingrecordsfortheRingbilledGullinWashington.Murrelet58:86-87.Ryder,J.P.,D.E.OrrandG. H.Saedi.1977.EggqualityinrelationtonestlocationinRing-billedGulls.WilsonBull.89:473-475.Ryder,J.P.andL.Somppi.1977.Growthanddevelopmentofknown-ageRingbilledGullembryos.WilsonBull.89:243-252.Schreiber,E.A.andR.W.Schreiber.1977GullswinteringinFlorida:ChristmasBirdCountanalysis.Fla.FieldNat.5:35-40.Scott,C.M.1977.FoodstealingbehaviorintheRing-billedGull.Bull.Okla.Ornithol.Soc.10:33.Shugart,G.w.1977a.Amethodforexternallysexinggulls.Bird-Banding48:118-121.1977b.ResidentRedFoxpredationuponanislandgullcolony.Jack--PineWarbler55:199-205.Sileo,L.,L.Karstad,R.Frank,M.V. H.Holdrinet,E.AddisonandH.H.Braun.1977.OrganochlorinepoisoningofRing-billedGullsinsouthernOntario.J.Wildl.Dis.13:313-322.Southern,W.E.1977.ColonyselectionandcolonysitetenacityinRing-billedGullsatastablecolony.Auk94:469-478.Southern,W.E.,W.L.JarvisandL.Brewick.1977.FoodhabitsandforagingecologyofGreatLakesregionRing-billedGulls.Proc.Symp.FisheatingBirdsoftheGreatLakes&EnvironmentalContaminants1977:100-144.Stevens,V.J.andF.D.Klopfer.1977.Interoculartransferofconditioningandextinctioninbirds.J.CompoPhysiol.Psychol.91:1074-1081.183

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Ring-billedGullVoous,K. H.1977.NortherngullsinAruba,NetherlandsAntilles.Ardea65:80-82.1976Baird,P.A.1976.ComparativeecologyofCaliforniaandRing-billedGulls(LaruscalifornicusandL.delawarensis).Ph.D.thesis,Univ.Montana/Missoula,MT.xand 183 pp.Dawson,W.R.,A.F.BennettandJ.W.Hudson.1976.MetabolismandthermoregulationinhatchlingRing-billedGulls.Condor78:49-60.Hunter,R.A.,H. A.RossandA.J.S.Ball.1976.Alaboratorysimulationofpredator-inducedincubationinterruptionusingRing-billedGulleggs.Can.J.Zool.54:628-633.Jarvis,W.L.andW.E.Southern.1976.FoodhabitsofRing-billedGullsbreedingintheGreatLakesregion.WilsonBull.88:621-631.Morris,R.D.andR. A.Hunter.1976.FactorsinfluencingdesertionofcolonysitesbyCommonTerns(Sternahirundo).Can.Field-Nat.90:137-143.Payne,R.B.andH.F.Howe.1976.Cleptoparasitismbygullsofmigratingshorebirds.WilsonBull.88:349-351.Ryder,J.P.1976.TheoccurrenceofunusedRing-billedGullnests.Condor78:415-418.Shugart,G.W.1976.EffectsofRing-billedGullnestingonvegetation.JackPineWarbler54:50-53.Southern,W.E.1976a.MigrationalorientationinRing-billedGullchicks.Auk93:78-85.1976b.RetractionoflongevityrecordsforRing-billedGulls.Auk93:--213.1975Cuthbert,F.J.andW.E.Southern.1975.Amethodformarkingyounggullsforindividualrecognition.Bird-Banding46:252-253.Evans,R.M.1975.ResponsivenesstoadultmewcallsinyoungRing-billedGulls(Larusdelawarensis):effectsofexposuretocallsaloneandinthe presenceofvisual imprintingstimuli.Can.J.Zool.53:953-959.Hanzely,L.,W.E.SouthernandD.Molsen.1975.UltrastructureoftheRingbilledGulleyepecten.Cytobios12:191-201.Harrington,B.A.1975.PelagicgullsinwinteroffSouthernCalifornia.Condor77:346-350.184

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Ring-billedGullHarris,J.T.andS.W.Matteson.1975a.Gullsandternsasindicatorsofman'simpactuponLakeSuperior.Dniv.Wis.SeaGrantCollegeProgramTech.Rep.No.227.45pp.1975b.GullsandternsonWisconsin'sLakeSuperiorshore.Passenger--Pigeon37:99-110.1975c.GullsandternsnestingatDuluth.Loon47:70-77.Kleen,V.M.1975.Thechangingseasons;thenestingseason:JuneI-July31,1975;MiddlewestPrairieRegion.Am.Birds29:978-982.Miller,D. E. andJ.T.Emlen.1975.IndividualchickrecognitionandfamilyintegrityintheRing-billedGull.Behaviour52:124-144.Ryder,J.P.1975.Egg-laying,eggsize,andsuccessinrelationtoimmaturematureplumageofRing-billedGulls.WilsonBull.87:534-542.Southern,W.E.1975a.LongevityrecordsforRing-billedGulls.Auk92:369.1975b.OrientationofgullchicksexposedtoProjectSanguine'selectro--magneticfield.Science189:143-145.Waide,R.B.,andJ.P.Hailman.1975.Differenthead-scratchingattemptsinaone-leggedgullandparrot.Condor77:350.1974Citron,J.D.1974.GullsstealfishfromCommonMergansers.DelmarvaOrnithoI.9:71-72.Dexheimer,M.andW.E.Southern.1974.BreedingsuccessrelativetonestlocationanddensityinRing-billedGullcolonies.WilsonBulL86:288-290.Forsythe,D.M.1974a.Anecologicalstudyofgullpopulationstoreducethebird/aircraftstrikehazardatCharlestonAirForceBase.AirForceWeaponsLab.Tech.Rep.73.142pp.1974b.Gulls,solidwastedisposal,andthebird-aircraftstrikehaz--ard.Pp.17-25inS.A.Gauthreaux,Jr.(ed.).Aconferenceonthebiologicalaspectsofthebird/aircraftcollisionproblem.ClemsonUniver-sity,Clemson,SC,5-7February1974.xiand513pp.Freitag,R.,J.P.RyderandP.Wanson.1974.Mite(acarina)populationsinRing-billedGullnests.Can.Entomol.106:319-327.Janssen,R.B.1974.Ring-billedGullnestingatthePortTerminal,Duluth.Loon46:175-176.185

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Ring-billedGullLudwig,J.P.1974.RecentchangesintheRing-billedGullpopulationandbiologyintheLaurentianGreatLakes.Auk91:575-594.Merilees,W.J.1974.Ring-billedandCaliforniaGullnestingcolonyinsouthcentralBritishColumbia.Can.Field-Nat.88:484-485.Ponshair,J.F.1974.Ring-billedGullsnestinginMuskegonCounty.JackPineWarbler52:194-195.Ryder,J.P.1974.OrganochlorineandmercuryresiduesingullsIeggsfromwesternOntario.Can.Field-Nat.88:349-352.Southern,W.E.1974a.GreatLakesregion.FloridadistributionofRing-billedGullsfromtheBird-Banding45:341-352.1974b.SeasonaldistributionofGreatLakesregionRing-billedGulls.--Jack-PineWarbler52:155-179.1974c.TheannualrangeofRing-billedGullsintheeasternUnited--States: with commentsonpotentialbird/aircraftcollisionproblems.pp.149-190inS.A.Gauthreaux,Jr.(ed.).Aconferenceonthebiologicalaspects Of thebird/aircraftcollisionproblem,ClemsonUniversity,Clemson,SC,5-7February1974.xiand513pp.1974d.Theeffectsofsuperimposedmagneticfieldsongullorienta--tion.WilsonBull.86:256-271.1974e.Copulatorywing-flagging:asynchronizingstimulusfornesting--Ring-billedGulls.Bird-Banding45:210-216.Southern,W.E.andF.R.Moore.fromtheGreatLakesregion.1974.RangeextremesforRing-billedGullsInl.BirdBanding News 46:83-87.1973Briggs,D.1973.Gullisland.Pac.Discovery26:22-25.Cuthbert,F.J.andW.E.Southern.aresultofeggdisplacement.1973.Ring-billedGullrelocatesnestasBird-Banding44:225-227.Evans,R.M.1973.DifferentialresponsivenessofyoungRing-billedGullsandHerringGullstoadultvocalizationsoftheirownandotherspecies.Can.J.Zoo1.51:759-770.Forsythe,D.M.1973.GullpopulationsatCharleston,S.C.,June1971toJune1972.Chat37:57-62.Grant,P.S.1973.FieldidentificationofRing-billedGulls.Brit.Birds66:115-118.186

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Ring-billedGullHume,P.A.1973.Ring-billedGullinGlamorgan:aspeciesnewto Britain andIreland. Brit. Birds66:509-512.Leek,C.F.1973.AttemptsofgullstorobaCommonEgretandanOsprey.Cassinia54:32.Vermeer,K.1973c.Comparisonofegg-layingchronologyofHerringandRingbilledGullsatKawinawLake,Manitoba.Can.Field-Nat.87:306-308.Vinicombe,K.E.1973.AsecondRing-billedGullinGlamorgan. Brit. Birds66:513-517.1972Dwernychuk,L.W.andD.A. Boag. 1972.Ducksnestinginassociationwithgulls--anecologicaltrap?Can.J.Zool.50:559-563.Forsythe,D.M.1972.Ring-billedandHerringGullrecoveriesfromSouthCarolina.Bird-Banding43:276-281.Gibson,D.D.1972.SightrecordsoftwobirdsnewtointeriorAlaska.Murrelet53:31-32.Miller,D.E.1972.ParentalacceptanceofyoungasafunctionofincubationtimeintheRing-billedGull.Condor74:454-461.Moos,L.M.1972.GullbandinginMontana.Proc.Mont.Acad. Sci. 32:20-23.Ryder,J.P.andD.J.Chamberlain.1972.Congenitalfootabormalityinthe Ring'-billed Gull.WilsonBull.84:342-344.Seymour,N. R.1972.SuccessofthreegullspeciesfeedingonswarmingantsinAntigonishCounty,NovaScotia.Can.Field-Nat.86:391-392.Siegfried,W.R.1972.Ring-billedGullsrobbingLesserScaupoffood.Can.Field-Nat.86:86.Southern,W.E.1972a.MagnetsdisrupttheorientationofjuvenileRingbilledGull.BioScience22:476-479.1972b.Influenceofdisturbancesintheearth'smagneticfieldon--Ring-billedGullorientation.Condor74:102-105.1972c.Useofthecannon -net inRing-billedGullcolonies.Inl.Bird--BandingNews44:83-93.Southern,W.E.andF.J.Cuthbert.1972.Ring-billedGullchick(Larusdelawarensis)withheaddeformities.Jack-PineWarbler50:92-93.----187

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Ring-billedGull1971Brackbill,H.1971.HerringandRing-billedGullspairedorcourtinginMarylandinJanuaryandFebruary.Auk88:438-440.Hardin,W.D.1971.Ring-billedGullstealsfishfromcoot.Bull.Okla.Ornitho1.Soc.4:5-6.Ivanovs,M.1971.NestingofRing-billedandHerringGullsatMilleLacsLake,1970.Loon43:58-59.Leek,C.F.1971.NocturnalhabitsofRing-billedGulls(Larusdelawarensis)atThimbleShoal,Virginia.Chesapeake Sci. 12:188.--Parmalee,D.F.1971.NestingofRing-billedGullsonLeechLake,CassCounty,Minnesota.Loon43:72-74.Southern,W.E.1971.Evaluationofaplasticwing-markerforgullstudies.Bird-Banding42:88-91.1970Evans,R.M.1970.ImprintingandcontrolofmobilityinyoungRing-billedGulls(Larusdelawarensis).Anim.Behav.Monogr.3:193-248.Southern,W.E.1970.Orientationingulls:effectofdistance,directionofrelease,andwind.LivingBird9:75-95.Tolonen,K.E.1970.Ring-billedGullandLaughingGullcatchfishby"ploughing"and"skinnning".WilsonBull.82:222-223.Vermeer,K.1970.BreedingbiologyofCaliforniaandRing-billedGulls:astudyofecologicaladaptationtotheinlandhabitat.Can.Wild1.Serv.Rep.Ser.12.52pp.1969Emlen,J.T.andD.E.Miller.1969.Pace-settingmechanismofthenestingcycleinaRing-billedGullcolony.Behaviour33:237-261.Southern,W.E.1969a.OrientationbehaviorofRing-billedGullchicksandfledglings.Condor71:418-425.1969b.Gullorientationbehavior:influenceofexperience,sex,age,--andgroupreleases.Jack-PineWarbler47:34-43.1969c.SkyconditionsinrelationtoRing-billedandHerringGull--orientation.Trans.Ill.StateAcad.Sci.62:342-349.188

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Ring-billedGullVermeer,K.1969a.EggmeasurementsofCaliforniaandRing-billedGulleggsatMiquelonLake,Alberta,in1965.WilsonBull.81:102-103.1969b.EndoparasiticvariationbetweenCaliforniaGullsandRing--billedGullsLaruscaliforniusandL.delawarensis.IbisIll:393-395.---1968Berndt, R. andU.Rahne.1968.ErstnachweissderRingschabelmowe(Larusdelawarensis)inEuropa.J.Ornithol.109:438-440.[In Kadlec,J.A.andW.H.Drury.1968a.StructureoftheNewEnglandHerringGullpopulation.Ecology49:644-676.Ludwig,J.P.1968.DynamicsofRing-billedGullandCaspianTernpopulationsoftheGreatLakes.Ph.D.thesis,Univ.Michigan/AnnArbor,MI.Rabinowitch,V.E.1968.Theroleofexperienceinthedevelopmentoffoodpreferencesingullchicks..Anim.Behav.16:425-428.Southern,W.E.1968.AgecompositionofabreedingRing-billedGullpopulation.Inl.BirdBandingNews40:166-167.1967Dick,J.A.1967.AnalbinoRing-billedGullfromOntario.Can.Field-Nat.81:224-225.Hart,J.A.1967.Ring-billedGullsSummerinwesternMinnesota.Loon39:139.LudWig,J.P.1967.Band-loss--itseffectonbandingdataandapparentsurvivorshipin.theRing-billedGullpopulation.Bird-Banding38:309-323.Ryder,R. A.1967.MigrationandmovementsofsomegullsfromColorado.Colo.FieldOrnithol.2:16-20.Southern,W.E.1967a.DispersalandmigrationofRing-billedGullsfromaMichiganpopulation.Jack-PineWarbler45:102-111.1967b.Colonyselection,longevity,andRing-billedGullpopulations:--preliminarydiscussion.Bird-Banding38:52-60.1967c.TheroleofenvironmentalfactorsinRing-billedandHerring---Gullorientation.Ph.D.thesis,CornellUniv./Ithaca,NY.Vermeer,K.1967.Astudyoftwospeciesofgulls,LaruscalifornicusandL.delawarensis,breedinginaninlandhabitat. Univ.Alberta/Edmonton,AB.128pp.189

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Ring-billedGull1966Bendell,J.F.1966.AnunusualnumberofdeadRing-billedGulls.Can.Field-Nat.80:62.Buckley,P.A.1966.Foot-paddlinginfourAmericangulls,withcommentsonitspossiblefunctionandstimulation.Z.Tierpsychol.23:395-402.Emlen,J.T.,D.E.Miller,R.M.EvansandD.H.Thompson.1966.PredatorinducedparentalneglectinaRing-billedGullcolony.Auk83:677-679.Evans,R.M.1966.ThedevelopmentofmobilityandapproachresponsesinyoungRing-billedGulls.Ph.D.thesis,Univ.Wisconsin/Madison,WI.Hickey,J.J.,J.A.KeithandF.B.Coon.1966.AnexplorationofpesticidesinaLakeMichiganecosystem.J.Appl.Ecol.3(suppl.):141-154.Ludwig,J.P.1966.HerringandRing-billedGullpopulationsoftheGreatLakes1960-65.GreatLakesRes.Div.,Univ.Mich.Pub!.15:80-89.Ryder,R.A.1966.SomemovementsofRing-billedGulls color""1Ilarked inColorado.BlueJay24:73-75.1965Anderson,W.1965.Waterfowlproductioninthevicinityofgullcolonies.Calif.FishGame51:5-15.Holroyd,G.L.1965.Ring-billedGullsonaLakeEriecolony.Ont.BirdBanding2:19-33.LeFebvre,E.1965.Ring-billedGullnestingrecordsforMinnesota.Loon37:79.Ludwig,J.P.1965b.Ring-billedGullnestingwithinCaspianTerncolony.Jack-PineWarbler43:61.Meyerriecks,A.J.1965.Ring-billedGullsgorgeonfiddlercrabs.WilsonBull.77:402-403.Mueller,H.C.andD.D.Berger.1965.Ring-billedGullsfeedonflyingants.Auk82:504.1964Hiemenz,N.M.1964.Ring-billedGullnestinginMinnesota.Loon36:133.Tomkins,I. R. 1964.TheRing-billedGull"skimming"onfoot.Oriole29:62.190

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Ring-billed (,ull 1963Hanson,W.C.1963.Censusofking-billedandCaliforniaGullcoloniesineasternWashington.Condor65:163-164.Holroyd,G.L.1963.DispersalofyoungRing-billedGullsfromMohawkIsland.Ont.FieldBioI.17:27-29.1962Ludwig,J.P.1962b.AsurveyofthegullandternpopulationsofLakesHuron,Michigan,andSuperior.Jack-PineWarbler40:104-119.1961Broadbrooks,H.E.1961. Ring-billed GullsnestingonColumbiaRiverIslands.Murrelet42:7-8.Dickerman, k. W.andE.LeFebvre.1961.Ring-billedGullnestinginMinnesota.Flicker33:23.Eastman,W.1961. Ring-billed Gullsfeedingonblueberries.Flicker33:57.Nero,R.W.1961.Drylandgullcolony.BlueJay19:166-168.1960Burton,D.E.1960. king-billed Gullsflycatching.Ont.FieldBioI.14:31.Railman,J.P.1960a.R.ing-billedGullsfollowingtheplow.Raven31:109.Rothweiler, k. A.1960.Foodhabits,movementsandnestingofgullsonawaterfowlarea,FreezeoutLake,TetonCounty,Montana.M.S.thesis,MontanaSt.College/Bozeman,MT.29pp.Turcotte,W.H.1960.Ring-billedGulls.Miss.Ornithol.Soc.Newsl.5:2.1959Janssen, R. B.1959.Ring-billedGullabundance.Flicker31:136.Moynihan,M.1959.NotesonthebehaviorofsomeNorthAmericangulls.IV:Theontogenyofhostilebehavioranddisplaypatterns.Behaviour14:214239.1958Moynihan,M.1958a.NotesonthebehaviorofsomeNorthAmericangulls.II:Non-aerialhostilebehaviorofadults.Behaviour12:95-182.191

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Ring-billedGullMoynihan,M.1958b.NotesonthebehaviorofsomeNorthAmericangulls.III.Pairingbehavior.Behaviour13:112-130.Pettingill,O.S.,Jr.1958a.Ring-billedGullshawkingmay-flies.Jack-PineWarbler36:154.1958b.NestingoftheCaspianTernwithina lUng-billed Gullcolony.--Jack-PineWarbler36:183-184.Sedwitz,W.1958.FiveyearcountoftheRing-billedGull(Larusdelawarensis)onwesternLongIsland.Proc.Linn.Soc.N.Y.66-70:71-76.1957Bartlett,L.M.1957.Ring-billedGullstealsfoodfromCoot.WilsonBull.69:182.Thiessen,G.J.andE.A. G. Shaw.1957.AcousticirritationthresholdofRing-billedGulls.J.Acoust.Soc.Am.29:1307-1309.1956Emlen,J.T.,Jr.1956.JuvenilemortalityinaRing-billedGullcolony.WilsonBull.68:232-238.Moynihan,M.1956.NotesonthebehaviorofsomeNorthAmericangulls.I.Aerialhostilebehavior.Behaviour10:126-178.1955Belknap,J.B.1955.TheexpandingrangeoftheRing-billedGull.Kingbird5:63.Emlen,J.T.,Jr.1955.Ring-BillCity...PassengerPigeon17:139-143.1954Johnston,D.W.andM.E.Foster.1954.InterspecificrelationsofbreedinggullsatHoneyLake,California.Condor56:38-42.1953Lamore,D.1953.Ring-billedGullsstealingfishfromfemaleAmericanMergansers.WilsonBull.65:210-211.Rapp,W.F.,Jr.1953.FieldseparationoftheHerring,Californiaand Ring billedGulls.Nebr.BirdRev.21:32-33.192

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Ring-billedGull1952Ganier,A.F.1952.Ring-billedGullintheGreatSmokyMountainsNationalPark.Migrant23:30.1951Savile,D. O.1951.TheRing-billedGullatOttawa,Ontario,anditsfieldrecognition.Can.Field-Nat.65:109-112.1950Cruickshank,A. D.1950.Ring-billedGullsandcabbagepalmettos.Auk67:237.Harper,S.A.1950.Ring-billedGullsfeedingonfruitofcabbagepalmetto.Auk67:236-237.Imhof,T.A.1950.Ring-billed'GullchasesGreatBlueHeron.WilsonBull.62:210.1949Southam,H.H.1949.CaspianTernandRing-billedGullcolonies,GreenIsland,PrinceEdwardCounty,Ont.Can.Field-Nat.63:115-116.1948Hyde,A.s.1948.BreedingoftheRing-billedGullinNewYorkin1936.Auk65:317.Nicholson,D.J.1948c.Berry-feedingoftheRing-billedGull.Auk65:601.1947Cooke,M.T.1947.Anothertransatlanticspecies.Bird-Banding18:170.1946Bunker,A.1946.Gullstakingfishfrommergansers.Can.Field-Nat.60:115.Kutz,H.L.1946.BreedingoftheRing-billedGullinNewYork.Auk63:591.1945Norris-Elye,L.T.S.1945.Symbiotictendenciesamongbirds.Can.FieldNat.59:174.Sheppard, R. W.1945.Gullsandternshawkingflyinginsects.WilsonBulL57:204.193

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Ring-billedGullTrautman,M.B.andM.A.Trautman.1945.Ring-billedGullsfly-catching.WilsonBull.57:77.1943Alcorn,J.R.1943.Flight-feedingoftheRing-billed Gull. Condor45:199-200.LudWig,F.E.1943.Ring-billedGullsoftheGreatLakes.WilsonBull.55:234-244.1942Moffitt,J.1942.AnestingcolonyofRing-billedGullsinCalifornia.Condor44:105-107.Shortt,T.M.1942.AbnormalprimaryofRing-billedGull.Auk59:438.1941Brown,M.1941.Gullseatfruitofthecabbagepalmetto.Auk58:579.Lewis,H.F.1941.Ring-billedGullsoftheAtlanticcoast.WilsonBull.53:22-30.Lewis,H.F.andT.S.Hennessy.1941.AnestingcolonyofRing-billedGulls(Larusdelawarensis)intheSt.LawrenceRivernearGananoque,Ontario.Can.FieldNat.55:77.1939Norris,R.1939.Ring-billedGullsatSt.SimonsIslandduringJune.Oriole4:31.1936Munro,J.A.1936.AstudyoftheRing-billedGullinAlberta.WilsonBull.48:169-180.Thompson,M.1936.MinnesotanestingrecordoftheRing-billed Gull. Flicker8:50.1935McCabe,T.T.andE.B.McCabe.1935.Ring-billedGullkilledbyaCanadaGoose.Condor37:79-80.194

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Ring-billedGull1934Bailey,A.M.1934.Ring-billedGullfromBarrow,Alaska--acorrection.Condor36:248.Shelley,L.O.1934.GullnotesfromtheNewHampshireandMainecoasts.Auk51:83.1933Tomkins,1.R.1933.Ring-billedandHerringGullsattheSavannahRivermouthinJulyandAugustoAuk50:103.1932Shelley,L.O.1932.Ring-billedGull(Larusdelawarensis)ontheNewHampshireandMainecoastsinJuly.Auk49:476.1929Komarek,E.V.1929.Strangulationofgulls.WilsonBulL41:41.1926Lano,A.1926.Ring-billedGull(Larusdelawarensis)inArkansas.Auk43:87.1910Bishop,L.B.1910.Laruscanus:acorrection.Condor12:74.1907Saunders,W.E.1907.Ring-billedGull.WilsonBulL19:73-74.195

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Second-winterHerringandRing-billedGull13.PhotoguphbyRogerB.Clapp.

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HERRINGGULL(Larusargentatus)[DA:Solvmage,DU:Zilvermeeuw,F:Harmaalokki,FR:Goelandargente,GE:Silbermowe,Ie:Silfurmafur,IT:Gabbianoreale,JA:Segurokamome, NW: Gramake,po:Mewasrebrzysta, PR: Gaivota, KU: (SilverGull),SP:Gaviotaplatena,SW:Gratrut] GENERALDISTRIHUTION NorthAmericaHerringGullsbreedcasuallyonSt.LawrenceIslandandelsewhereintheBeringSea(Sowlset aL 1978),inlandinborealAlaska(PattenandWeisbrod1974),andacrossmainlandnorthernCanadatotheAtlanticOcean.InlandtheynestsouthonallfiveGreatLakes(Gilmanetal.1977,Shugart1977b),andnorthtosouthernBaffinIsland(Smith1966).OntheAtlanticcoastthisspeciesbreedsfromLabradorsouththroughtheMaritimeProvincesandNewEnglandtoNorthCarolina(Map5).TheywinterintheirbreedingrangeasfarnorthasopenwaterremainsandsouthonthePacificcoasttoMexico(sparsely).TheyalsoareabundantontheAtlanticcoastofthesouthernUnitedStatesandcommonontheGulfofMexicotoVeraCruz.SomewinterinsmallnumberssouthtoCentralAmerica(Cooke1940,KadlecandDrury1968a),butthespeciesisaccidentalinnorthernSouthAmerica(Donahue1977).IJorldDistributionHerringGullsbreedacrossnorthernEurasiafromtheBeringSeatoNorwayandsouthtoLakeBaikal,andtheCaspian, Hlack, andMediterraneanSeas.TheyalsobreedontheAtlanticcoastofMoroccoandontheCanaryandAzoresislands(Vaurie1965).TheEurasiansubspeciesaremuchmoresedentarythantheNorthAmericanform,althougheasternSiberianbirdsmigratesouthtoJapan,China,andKorea,andoccasionallystraggletothetropicalPacific(ClappandWoodward1968).MostEuropeanHerringGullswinterwithintheirbreedingrange. DISTRIHUTION ANDABUNDANCEIN THE COASTALSOUTHEASTERNUNITED STATES NorthCarolinaHerringGullsarecommoninNorthCarolinafromDecembertoMay(WrayandDavis1959).Mostoccuronthecoast,buttheyarealsofoundinlandalongwatercourses.FlocksofthousandsofHerringGullsareoccasionallyfoundontheOuter Hanks inwinter(LeGrand1977a).Althoughtotalwinterpopulationsareunknown,datafromafewChristmasCourits(Map6)suggestthatthewinteringpopulation nOli( numbersatleastinthetensofthousands.HerringGullshavebredregularlyinHorthCarolinasinceabout1972;thegrowthofthebreedingpopulationissummarizedinTable26andrecentcolonysitesareshowninMap5.BreedingpopulationsofHerringGullsinthecen-197

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ae-----+-------GULFOFMEXICOsAEXTDALLASBreedingRanleMapfarSoutheastern Unbed States Number.inboxesdenotemaximume.timate.of breedingbird.atcolonie.in recent years. Fir.t figure indicate. maximum number of bird.Second figure indicate. year inwhich estimatewasobtained. G:;XAMPLE320-77, MALNO.YEAR< "/"/ Map5

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INDIVIDUALSOBSERVEDDURINGCHRISTMASBIRDCOUNTS,1973-1977(ARITHMETICMEAN) @ Numberof individuals 8 Lessthanoneindividual NoneobservedWinterDistributianMapforSoutheasternUnitedStatesBIRD NAME'88" GULFOFMEXICO96X'ASEBIRDSPER10PARTY-HOURSIILess t"an oneliJllllltlliliiitll!111-5 err-M 5-20 ro)ntl Morethan20(Adapt.dfrom IYltrall. 1974) TMap6

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HerringGulltralAtlanticstateshavebeenincreasingrapidly(AOU1957,Armistead1975, Burger 1977a).ItisapparentthattheNorthCarolinacoloniescontinuetoreceiveimmigrantsfromthenorthbecausegrowthhasbeentoofasttostemfromasinglecolonizationintheearly1960's.MostHerringGullsbeginnestinginPamlicoSound(Map5)inMay;youngareoftenstillpresentintolateAugust(ParnellandSoots1979ms).HerringGullcoloniesinNorthCarolinavaryinsizefromsmalltomoderatelylarge,buttheyaresmallerthanthosefoundfurthernorth(1)inNorthAmerica.AnaveragecolonyfoundinNorthCarolinain1976(Portnoyet al. 1981)contained91nests(n=7,range=16-462),butthisfigureprobablyoverstatesthemean;20coloniesexaminedthefollowingyearheldanaverageof24nests(range=1-180)(ParnellandSoots1979ms).Fifteenofthesecoloniesheldlessthan10pairsand10hadonlyonepair.ThelargestcoloniesinNorthCarolinainrecentyearswereoneonHerringGullIslandthatheld462nestsin1976(Portnoyet al. 1981)andtwoondredge-spoilislandsin H.oanoke Soundin1977;oneoftheseheld180nests,theother114(ParnellandSoots1979ms).SouthCarolinaHerringGullsarecommontoabundantmigrantsandwinterresidentsincoastalSouthCarolina.TheyarecommonfromOctobertoMay(SpruntandChamberlain1949),andwinterpopulationshavebeenincreasingsteadily (Burton 1970).Althoughtheyapparentlyhavenotbredinthestate,HerringGullsrecentlyhaveextendedtheirbreedingrangesouthtotheCapeFear H.i ver,lessthan50km(30mi)northoftheSouthCarolinaborder(ParnellandSoots1975).ThespeciesprobablywillbefoundbreedinginSouthCarolinainthenearfuture.JudgingfromrecentChristmasCounts(Map6),HerringGullsarelessabundantinSouthCarolinainwinterthantheyareinNorthCarolina.However,thousandswinterinSouthCarolina.Forsythe(1973)indicatedthatwinterpopulationsnearCharlestonnumberedover2,500birdsin1971/72.Bandrecoverydata(Forsythe1972)indicatethatmostofthewinteringpopulationoriginatesfromcoloniesinnortheasternNorthAmerica,particularlythoseinNewBrunswick, Haine, Massachusetts,and l<.hode Island.OnlyasmallproportionofthebirdswinteringinSouthCarolinacomefrombreedingpopulationsintheGreatLakes.Mostoftherecoveries(77%)wereyoung-of-the-year;onlyasmallproportion(ca.2%)wereadults.GeorgiaDentonet al. (1977)summarizedthestatusoftheHerringGullsinGeorgia,statingthattheyare"residentoncoast,abundantexceptinsummer,nonbreeding."Burleigh(1958)suggestedthatthelargestnumberswerepresentfromaboutmid-OctoberthroughearlyMay. Recent ChristmasCounts(Map6)suggestthatwinteringpopulationsarelargestinthesouthernpartofthestate.(1)TheaveragenumberofnestsfoundincoloniesalongtheNorthAtlanticcoastoftheUnitedStatesin1977wassometimessmall,buttherewasanaverageofmorethan100nestspercolonyinallstatessurveyedexceptDelaware(Erwin1979a,Korschgen1979).Coloniescontainingthelargestaveragenumberofnestswerefoundin Rhode Island(401),Massachusetts(424),Maryland(459),andLongIsland, New York(563)(Erwin1979a).200

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HerringGullTable26.HerringGullsnestinginNorthCarolina(a).NumberofNestsnestingYearfoundsitesSouthernmostsitesSource1962GullIsland(35'N)Hailman19631963 Heacon Island(35.9'N) Ames1963197281lowerCapeFearRivernearParnellandSoots1975Southport(ca.34'N)197398lowerCapeFearRivernearParnellandSoots1975Southport(ca.34'N)1976640 Dump Island(34.5'N)Portnoyetal.1981197749023OldDrumInlet(34'N)ParnellandSoots1979ms(a)Onlyrecordsfor1972,1973,1976,and1977maybeassumedtobecompleteastherewerenoadequatesurveysofcoastalpopulationsofnestingmarinebirdsbeforethattime.FloridaHerringGullsarecommonwinterresidentsontheAtlanticcoastandintheFloridaKeys(Sprunt1954,HundleyandHames1960-1962);theyaremostabundantonthenorthernportionofthecoast(Kale1979msa;Map6). Hirds arriveinnumbersinmid-October,andmostaregonebymid-Hay.HerringGullsarealsocommoninwinterandveryrareinsummerontheFloridaGulfcoast.AsontheAtlanticcoast,populationsarelargertothenorth(Map6,SchreiberandSchreiber1977).ThelargestconcentrationsinrecentyearshavebeeninPinellasCounty,primarilyattheToytowngarbagedump.Inmid-February1978,thisdumpwasusedby12,000-15,000HerringGulls(W.Hoffman,pers.comm.)Alargeproportionofthewinteringpopulationconsistsofyoung-ofthe-year(KadlecandDrury1968a,Moore1976).AlabamaHerringGullsarecommonfromOctobertoAprilontheAlabamacoast concentrationof3,000birdswasnotedonDauphinIslandinNovember1970(Imhof1976b).HerringGullsarealsoabundantinlandinmuchofthestate.Althoughsomebirdsremainthroughthesummer,noneareknowntobreed. Hand recoveriesfromAlabamarevealthatbirdswinteringalongthecoastaremainlyfromAtlanticcoastbreedingpopulations,whereasmostofthosewinteringinlandcomefromcoloniesintheGreatLakes(Imhof1976b);Michigan, l,ew Hrunswick,andMainewerethesourceofoverathirdofallrecoverieslistedbyImhof.AlargeproportionofwinteringHerringGullsrecoveredinAlabama(andMississippi)wereyoungbirds,butanaerialcensusrevealedthatmoreadultsarepresentthanthenumbersfromrecoverydatasuggest(DruryandNisbet1972).AmajorityoftherecoveriesinAlabamaofbirds201

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HerringGullbandedintheGreatLakesarefromcoloniesinthewesternportion ofr thatarea(Moore1976).MississippiBurleigh(1944)consideredtheHerringGulla commonwinterresidentontheMississippicoastfromlateSeptemberuntillateApril.MostofthebirdsseenbyBurleighwereimmatures.Theseweresometimes.seeninflocksof200birdsormore. ]{ecent ChristmasBirdCountsinJacksonCountyrecordedfewerthan100HerringGullseachyear,suggestingthatfewerbirdswinterinlandthanonthecoast.JacksonandCooley(1978b)alsonotedthatthespeciesisrareinsummerinMississippi.LouisianaHerringGullsarecommononthecoast,alongcoastalwaterways,andalongtheMississippi ]{iver frommid-NovembertolateApril(Lowery1974).Somebirds,mainlyimmatures,remainalongtheGulfcoastinthesummer.AnaerialcensusconductedbyKadlecandDrury(1968a)revealedlargerWinteringpopulationsthanweresuggestedbyChristmasCountsbecausetheysurveyedtheoutersandyislandsoftheMississippi ]{iver Deltawhicharesparselyinhabitedbyhumans.KadlecandDrurynotedparticularlylargeconcentrationsonislandsatthemouthsofAtchafalaya,Caillou,TerrebonneandTimbalierBays.TexasAlongtheTexascoasttheHerringGulliscommontouncommon;inlanditiscommontoscarce(Oberholser1974).Thevariableabundancemaybeeitheramatterofdistributionbyflocksorofannualvariation.WinteringbirdsoccurfromSeptembertoApril,andSomenonbreedingbirdsremainthroughthesummer.SYNOPSISOFPRESENT DISTRIBUTIONANDABUNDANCEBreedingHerringGullsbreedacrossthenorthernportionsofbothNorthAmericaandEurasiatoabout80N(Southern1980a).InNorthAmericatheybreedsouthtotheGreatLakesandalongtheAtlanticcoasttosouthernNorthCarolina.MostofthosebreedingineasternNorthAmericaarefoundnorthof35N,butthoseinwesternNorthAmericabreedmainlynorthof48N.HostoftheNorthAmericanbreedingpopulationisconcentratedinnortheasternCanada,thenortheasternUnitedStatesandaroundtheGreatLakes(Southern1980a).TherangeofthespeciesinbothNorthAmericaandEuropehasexpandedinrecentyears.Theoverallsizeoftheworldpopulationisunknown,butbecausethespeciesisoneofthemostabundantgulls,thetotalpopulationprobablynumbersinthelowmillions.KadlecandDrury(l968a)documentedanincreasefrom4,000-8,000breedingpairsin1901to110,000-120,000pairsin1966inNewEngland(l).Approximately89,900pairsbredinanareaextendingfrom Haine toLongIsland,NewYork,in1972,andanother14,100bredintheGrandMananArchipelagothesameyear(DruryandKadlec1974).Theformerareacontainedapproximately78,800breed-(1)KadlecandDrury'sestimateapparentlyincludednotonlyNewEnglandbutpartofNewYorkandnorthfromMainetotheGrandMananArchipelago, New Brunswick.202

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HerringGullingpairsin1977;another13,200pairsnestedfromNewJerseytoVirginia (J<:rwin 1979a,Korschgen1979).Whetherornottheresultsofthelattercensusindicateadeclineinthebreedingpopulationisquestionable.IncreasingnumbershavebeenseenonChristmasCountsandthebreedingrangeofthisspeciescontinuestoexpandsouthward.In1977,HerringGullswerethemostabundantbreedingcolonialwaterbirdfromMainetoVirginia (J<:rwin 1979a).ThebulkofthepopulationwasconcentratedinMaine(26,000pairs)(Korschgen1979)andinMassachusetts(26,000pairs),butsubstantialnumbers(17,500pairs)werefoundnestingonLongIsland(Erwin1979a).LargenumbersofHerringGullsalsobreedintheGreatLakesregion.Scharf(1978)reportedthatapproximately28,000pairsbredinthevicinityoftheU.S.GreatLakesin1976-77,butthenumbersbreedingintheCanadianportionofLakeOntariohavedecreasedinrecentyears(BlokpoelandFetterolf1978).Scharf's(1979)1976censusofcolonialseabirdsontheU.S.GreatLakesfoundatotalof26,719nests.Thefivelargestcolonies,togethercontainingalittIemorethanone-fifthoftheentirepopulation,wereasfollows:Hat(800-1,000nests)andGull(1,426)IslandsinLakeMichigan;Gull(1,510)andBlack lli ver(1,064)IslandsinLakeHuron;andacolonynearthemouthofSanduskyBay(983)inwesternLakeErie.WelackadequateinformationonthenumbersofHerringGullsbreedinginCanada,butthetotalisprobablysubstantial.NumbersnestingontheAlaskancoastaresmall,probablylessthanafewhundredbirds(Sowlset al. 1978).LargepopulationincreaseshavebeenreportedintheHetherlands,Germany,Sweden,andBritain(Crampetal.1974,andreferencestherein).About333,000pairsbredinBritainandIrelandin1969-70(Crampetal.1974),about60,000pairsbredinDenmarkin1970-72(Salomonsen1979),about11,000pairsbredonthenorthcoastofFinlandin1980(Kilpietal.1980),andatleast260,000pairsbredalongtheNorwegiancoastin1970-74(Brun1979).WinterHerringGullswinterbothin,andtothesouthof,theirbreedingrange(Map6).InNorthAmerica,theyarecommonsouthtoFloridaandTexas,lesscommonsouthtoVeraCruz,Mexico,andcasualsouthtoPanama.BirdsbreedinginNorthCarolinaprobablywinterinthesoutheast,buttheymustrepresentaminorfractionofthetotalwinter;i.ngpopulationofthatregion.Infavorablelocalities,flocksofvariousspeciesofgulls,includingalargecontingentofHerringGulls,maycontainseveralthousands(LeGrand1977a).KadlecandDrury(1968a)countednearly572,000HerringGullswinteringfromMainetoTexasin1965;thetotalpopulationisundoubtedlylargernow.MapsofmonthlyrecoveriesplottedbySouthern(1980a)showedthatthemostsignificantwinteringground(January-March)forNorthAmericanHerringGullsisontheNorthAtlanticcoastbetween42and36N(fromBoston,Massachusetts,totheOuterBanksofNorthCarolina).Aboutaquarterofallrecoveriesobtainedduringthisperiodwerefromthisarea.203

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HerringGullHigrationRecentbandingstudiesdocumentingmigrationofyoungHerringGullsfromnortherncolonieshavebeenconductedatWitlessBay,Labrador(Threlfall1978),CapeAnn,Hassachusetts(Dexter1978),ChimonIsland,Connecticut(DeWire1981)andJamaicaBay,NewYork(Burger1981c). Host youngfromWitlessBaymigratesouthtowinterinthenortheasternUnitedStates,butthosefromMassachusetts,Connecticut,andNewYorklargelywinterlocallyandforrelativelyshortdistancestothesouth.Of 227recoveriesfromNewfoundland,5first-winterbirdswereencounteredintheCarolinasandonlyonewasrecoveredfarthersouth,inCuba;asecond-yearbirdwasobtainedinFlorida.Only3of126birdsbandedatCapeAnnwererecapturedintheCarolinas,andanotherwasfoundontheGulfCoastofFlorida.YoungbandedinConnecticutwinteredlocallyandsouthtoNewYorkandNewJersey,butafewwererecoveredinNor.thandSouthCarolina,GeorgiaandFlorida;individualsstraggledtoMississippiandVeraCruz, Hexico. Sevenof158sightingsofbirdstaggedatJamaicaBaywererecoveredinthesoutheast,oneinSouthCarolina,andthreeeachinGeorgiaandFlorida.AgreatmanyearlystudieshaveprovidedinformationonmovementsfrombreedingcoloniesinthenortheasternUnitedStatesandtheGreatLakesarea(Lincoln1928a;Eaton1933,1934a,1934b;Hinchman1934;Shelley1934a;Gross1940,1944;Poor1943;Paynter1947a;Hofslund1959;Smith1959;Gillespie1961;LudwigandLudwig1961;DennisandPepper1962;Ludwig1962a;Drury1963a,1965a;Southern1968;KadlecandDrury1968a;DruryandNisbet1972;Forsythe1972;Moore1973,1976;Southern1980a).ThesestudiesmakeitapparentthatGreatLakesHerringGullswinterpredominantlyalongtheGulfcoastandinFlorida,moresothanonthecoastsofGeorgiaandtheCarolinas.Alargeproportionofthemigrantsthroughoutthesoutheastareyoungbirds,asadultstendtowinterinthevicinityoftheirbreedingcolonies.Southern(l980a)foundthatsomeHerringGullsmayleavetheirbreedingcoloniesinlateJuly,butothersremainforsometimeinornearthenestingareas.Thesegullsmaywanderforseveralmonthsthereafter,onlygraduallymovingsouth,althoughsomearriveinthesouthearlyinSeptember.MostofthemigrantpopulationsreachthesouthernmostpointofdispersalinJanuaryandFebruary,andreturntotheirbreedingrangeinFebruaryandearlyMarch(Southern1980a).HABITATNestingHerringGullsnestinawidevarietyofhabitats,includingsandyislands(Shugart1977b),stabilizedsanddunes(Tinbergen1961,Harris1964a),themarginsoftundralakes(Smith1966),Spartinamarshes(Burger1977a),cliffs(Harris1964a,Andrle1976),woodedislands(Chamberlin1975;Patten,pers.comm.),andevenonbuildings(MonaghanandCoulson1977).InsomeareassuchasNewEngland,coloniesareoftensituatedneardumpsorlandfillsthatserveasthemainsourceoffood.Theprimaryrequirementsforanestingsitemaybefreedomfromterrestrialpredatorsandproximitytofoodandwater.MostHerringGullsnestinexposedsites,butChamberlin(1975)reportedthatsomenestincedarstands.204

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HerringGullHerringGullsprefertonestinlowsitesandareoftenpresentontheslopesanddomesofdredge-spoilislands.Thenestingsubstratemaydeterminethenestingdensity.NestsatGullIsland,Newfoundland,weresomewhatmoredenselypackedonrock(3.98nests/l00sqm[0.37nests/lOOsqft])thanongrass(3.07nests/l00sqm[0.29nests/lOOsqft]).However,nestdensitywasgreatest(7nests/lOOsqm[0.65nests/lOOsqft])attheinterfacebetweenthesetwosubstrates(HaycockandThrelfall1975).HaycockandThrelfallalsonotedthatnestdensitywaslowerinmoreopenareasandthatHerringGullsongrassyslopestendedtoplacetheirnestsatthebaseofemergentfeaturessuchasbouldersorstumps.AtGullIslandnogullsnestamongheavyshrubs,butafewnestupto4m(13ft)insideopenwoods;thenestshereareusuallyplacedunderatree.Pierotti(1982)reportedthatnestdensityofHerringGullsatGreatIslandisgreatestinrockyareaswherethereisthemostshelter.HerringGullsnestingintheopenatGooseIslandinnorthernLakeHuronweresignificantlyfartherapart(meandistance=17.2ft[5.2m])thanwerethoseinhabitatsdominatedbynorthernwhitecedar(Thujaoccidentalis)andred-osierdogwood(Cornusstolonifera)(meandistance=10.1ft[3.1m])(Chamberlin1975).---. llurger andLesser(1980)examinedcolony-andnest-siteselectionfor28HerringGullcoloniesonsalt-marshislandsin llarnegat Bay,NewJersey.Thecolonieswerefoundonislandsatleast5ac(2ha)insize,andmost(85.7%)ofthecolonieswerelocatedinthecenteroftheislands.Almostfour-fifthsofthecolonieswerefoundonislandswithbushes(Iva,Baccharis).Islandswithnestscontained,ontheaverage,50%smoothcordgrass(5partinaalterniflora),33%saltmeadowcordgrass(5.patens),2%common australis),and7%standingwater. On theseislands,HerringGullsnestedprimarilyin5.patensandunderbushes(39%ofthenestswerefoundunderbushes,althoughbushesmadeuponly5%ofthevegetativecover).Gullsnestinginmainlandcolonieschoseareaswithsimilarvegetationandwerefoundinsitesseparatedfromthemainlandbysmallcreeks.AdditionalinformationonsitesusedinNewJerseyandthevariousfactorsinvolvedinnest-siteselectioninthatareaisgivenbyBurger(l977a,1977b,1980a),andBurgerandShisler(l978a,1978b,1979,1980b).All1976coloniesofHerringGullsinNorthCarolinawerefoundoncoastalislandsthatoriginatedfromdredge-spoil(Portnoyetal.1981),and99%and97%ofthepopulationoccupieddredge-spoilsitesin1973(SootsandParnell1975a)and1977(ParnellandSoots1979ms),respectively.Nestsareoftenfoundondriftadjacenttosaltmarshesandarealmostalwaysplacednearorbetweenlarge,tallclumpsofvegetation(SootsandParnell1975a,ParnellandSoots1979ms).HerringGullsnestedinmoreheavilyvegetatedcolonies(meancover=45%)thananyotherseabirdstudiedexcepttheLaughingGulland Fors ter'sTern;theaverageheightofnestcover(ca.0.75m[2.5ft])wasalsohigherthanforotherspeciesstudied(SootsandParnell1975a).PlantsmostfrequentlyfoundinHerringGullcoloniesinNorthCarolinawereseasidegoldenrod(Solidagosempervirens),apanicgrass(Panicumamarulum),andbeachgrass (AmiiiO]>hila breviligulata)(SootsandParnell 197sar:-205

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HerringGullFeedingHerringGullsareopportunisticfeedersingarbagedumps,aroundseafood-processingoperations,inpasturesandcultivatedfields,onlawns,tundra,andbeaches,andatsea.Theonegeneralrequirementisthattheirfeedinghabitatberelativelyopen.Drury(l963a)describedfeedinghabitatsinNewEnglandinsome.detail.Foodof"natural"originisobtainedfromsandyandrockyshoresandshallowinshorewaters.Inland,HerringGullsfeedonberriesandinvertebratesinfields.Foodofman-madeoriginisobtainedatthebackoffishingboats,atfish-cleaningoperationsatpiersandwharves,fromtruckscarryingwastefishtofoodandfertilizerplants,atpigfarms,atseweroutflows,andatthemanyotherplaceswhereorganicwastesmaybefound.HerringGullsinMaineforageprimarilyondumpsandtidalmudflats,butarealsofoundinareaswherewastesaredischargedfromsewersandfood-processingplants,andalongtheshoresinareaswheremusselsareparticularlyabundant(HuntandHunt1973).Drury(1963a)characterizedtheidealdumpforfeedingas"anopentreelesssitewithalarge,flatloafingground,andwithinhalfamileoffreshwater."Henotedthatthenumberofgullsusingdumpsincreasedwithhightides,onshorewinds,adverseweatherconditions(suchaswindandfog),andwiththepresenceoficeontidalflats.HealsoremarkedthatHerringGullsfeedonlyonfreshgarbageandareattractedtodumpswherebulldozersuncovermaterial.Her-ringGullsfeedingina dumpinNewJerseyfedlargelywherefreshgarbagewasdepositedandtoalesserdegreeonaslopewherethisgarbagewaspushedbybulldozers(Burger1981b);adultsfedmoreoftenatthebottomofthisslopethandidyoungergulls.DruryandNisbet(1972)marked4,900breedingHerringGullsatcoloniesinMassachusettstoexaminetheirforagingmovements.TheysummarizedtheirworkbystatingthatHerringGulls"wentnofartherthanthenearestdependablefoodsourcebutcommutedreadilyasfaras40 km[25miJtoagoodfoodsupply."Theypointedoutthatforagingrangesfromdifferentcoloniesareverydifferentandthatthereislittleoverlapinfeedingrangesbetweencolonies.NonbreedingandOffshoreWinterhabitatsareessentiallythesameasthoseusedduringthebreedingseasonforfeeding,loafingandroosting.HerringGullsarefoundprimarilyalongtheshoreoftheoceanorotherbodiesofwater,concentratingonbeachesandinareaswherefoodislikelytobeabundant(Oberholser1974,Imhof1976b).HerringGullsalsoventureinlandtogarbagedumpsandfields.Winteringbirdsroostonisolatedsandbars,onprotectedwaterways,andinisolatedfields(Operholser1974)Gullsregularlyfollowboats,particularlyfishingboats,tofeedonoffal.Slightlynorthofourstudyarea,Rowlett(1980)reportedthatchummingattractslargenumbersofHerringGullsfromNovemberthroughMarch;flocksreachedmaximumnumbersneartheedgeoftheContinentalShelf.Suchflocksmaybelarge.Rowlettreportedasmanyas20,000aroundfishingvessels,75and90km[47and56mi1eastsoutheastofOceanCity,MarylandinearlyDecember1977;Pimlott(1952)notedoneflockofabout4,000HerringGullsfeedingnearthesouth206

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Herring Gull shoreofDeerIsland,NewBrunswickinlateJuly1949. During sixhoursofobservationsome3,700gullswereseenupto120km[75miJeastofOceanCityinearlyFebruary1977(Rowlett1980).FOODAND FEEDINGBEHAVIOK HerringGullsfeedlargelyonanimalmatter,includingsmallmammals,birdsandbirdeggs,amphibians,fish,andagreatvarietyofinvertebrates;theyalsoeatberriesandsomefruit.Carrionisamajorportionofthedietandinsomeareasorseasonsgarbageisveryimportant.AsSpruntandChamberlain(1949)stated,theHerringGull"takespracticallyanyanimalfooditcanfind."Thespeciesisnotedforitshabitofdroppingmolluscsandotherhardfoodontobeaches,roadways,roofs,orparkinglotstoshatterthefoodandobtainaccesstothesoftinterior.Tolonen(1976)notedthatHerringGullsobtainfoodfrommusselsbytwoothermethodsaswell:(1)agullmayinsertitsbeakintoanopenmusselandseizetheflesh;or(2)thebirdmayforceitswayintoaclosed shell. SeveralauthorshavestudiedtheeffectofageonforagingefficiencyofNorthAmericanHerringGulls.Tolonen(1976)reportedthatadultHerringGullsfeedingatConowingoDam,Maryland,capturedfishonasignificantlyhigherproportion(93%)oftheirattemptsthandidimmaturebirds(79%).Adultswerealsomoresuccessfulthanimmaturesatfeedingonmoonsnails(Naticidae),musselsandquahogsinConnecticut,andonclamsinNorthCarolina.IngolfssonandEstrella(1978)reportedthatadultsweremoresuccess-fulatfeedingonmusselsinsoutheasternMassachusettsthanwereyoungerbirds.Ata dumpinNewJersey,HerringGullsfedprimarilybywalkingoverthesurfacepickingupfooditemsandbyturningovergarbage(Burger1981b).Most(67%)ofthebirdswereyoung-of-the-year,butsomeadultsandsubadults(birds16-18monthsold)werealsopresent.HerringGullskleptoparasitizebirds,stealingpreyfromtheirown(Tolonen1976)andotherspecies.InNorthAmerica,HerringGullshavebeenseenstealingfoodfrom lting-billed Gulls,GreatBlack-backedGulls(Tolonen1976),ParasiticJaegers(Stercorariusparasiticus)(Morrisson1978),andCommon Her gansers (Hergus merganser)(Lovell1945).IntheOldWorldHerring Gulls havebeenseenstealingfoodfromAtlanticPuffins(Fraterculaarctica) (Hylne 1960)andCommonEiders(Somateriamollissima)(Ingolfsson1969a).TheyhavebeenseenattemptingtoseizefoodfromGreatBlueHerons(Ardeaherodias)inNorthAmerica(Tolonen1976,Quinneyet al. 1981)andGreyHerons(Ardeacinerea)intheOldWorld(Karmborg1979).FurtherinformationonotherreportedorpotentialvictimsofkleptoparasitismbyHerringGullsmaybefoundinother207

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HerringGullpaperscitedintheterminalspeciesbibliography(e.g.,Steiniger1952,Schmidt1954,Duchrow1958,Bergman1960).HerringGullsalsofeedonbirdsandmammals.Theyfrequentlyfeedontheyoungofothermembersoftheirownspecies,aswellasontheeggsandyoungofotherlaridsnestinginassociationwiththem.LaridseatenbyHerringGullsinNorthAmericaincludeeggs(Mendall1935)andyoungofCommonTerns(Mendall1935,CrowellandCrowell1946,Ligas1952,Houde1977b),youngRoseateTerns(Houde1977b),youngRing-billedGulls(Emlen1956,FetterolfinSchnell1981),andeggsandyoungofCaliforniaGulls(Laruscalifornicus)(Nero1961).OthernonpasserineavianpreyrecordedinthedietofNorthAmericanHerring Gulls includeeggsandyoungofDouble-crestedCormorants(Phalacrocoraxauritus)(Mendall1935),Wilson'sStorm-Petrel(Oceanitesoceanicus)(Houde1977b),Leach'sStorm-Petrel(Oceanodromaleucorhoa)(Gross1935),Oceanodromastorm-petrels(Houde1977b),youngRed-breastedMergansers(Mergusserrator)(Braunetal.1980),youngCommonEiders(MinotinBraunetal.1980),adultOldsquaws(Clangulahyemalis)(Snell1981),AmericanKestrels(Falcosparverius)(Houde1977b),adultSpottedSandpipers(Aetitismacularia),adultSemipalmatedSandpipers(Calidrispusilla),youngBlackGuillemots(Cepphusgrylle)(Mendall1935),andMourningDoves(Zenaidamacroura)(Houde preyeateninNorthAmericaincludeBlueJays(Cyanocittacristata)(Houde1977b),PurpleMartins(Prognesubis)(Ligas1952),CommonGrackles(Quiscalusquiscula),Red-wingedBlackbirds(Agelaiusphoeniceus)(Ludwig1966,Houde1977b),Brown-headedCowbirds(Molothrusater),AmericanRobins(Turdusmigratorius),GrayCatbirds (Dumeteila-carorrnenB'is), Starlings(Sturnusvulgaris)(Houde1977b),adultBlack-throatedGreenWarblers(Dendroicavirens)(Mendall1935),unidentifiedwarblers(Dendroicasp.)(Houde1977b), and SongSparrows(Melospizamelodia)(Mendall1935).ApartiallistofbirdsreportedeatenintheOldWorld,excludingthosementionedabove,isasfollows:adultandyoungManxShearwaters(Puffinuspuffinus),eggsand/oryoungNorthernGannets(Sulabassanus)(Harris1965),adultMallards(Anasplatyrhynchos)(Andersson ---r9"70), youngShelducks(Tadornatadorna) ('VanDobben 1934inTinbergen1953),fledglingGadwall(Anasstrepera)(Sabinevskii1958inBianki1967),GrayPartridge(Perdixperdix)(Spitzer1976),youngCoots(Fulicaatra)(Lugovoi1961inBianki1967),eggsand/oryoungNorthernLapwings(Vanellusvanellus)(Harris1965),youngOystercatchers(Haematopusostralegus)(Harris1965,Bianki1967),Avocets(Recurvirostraavosetta), lUnged Plovers(Charadriushiaticula),Kentish[= Snowyralexandrinus)(Tinbergen1953),eggsand/oryoungofGreatBlack-backedGulls,LesserBlack-backedGulls(Larusfuscus),andBlack-headedGulls(Larusridibundus)(Harris1965),eggsandyoungofCommonGulls(Laruscanus)(Bianki1967),youngBlack-leggedKittiwakes(Rissatridactyla) (BeID="""" pol'skii1957),eggsandyoungofArcticTerns(Bianki1967),Razorbills(Alcatorda)andCommonMurres(Uriaaalge)(Harris1965),youngMeadowPipits-(Anthuspratensis) (Bianki1%7f;-Yellow Wagtails(Motacillaflava)(Spitzer (Alaudaarvensis)(Borodulina1960),Redwings(Turdusmusicus),andBlackbirds(Turdusmerula)(Hobbs1959).-------208

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HerringGullHarris(1965)citedotherauthorsfortheeggsand/oryoungofNorthernFulmars(Fulmarisglacialis),Shovelers(Anasclypeata),Ring-neckedPheasants (PhasianuSCOI'cliIcus), EurasianCurlews(Numeniusarquata),Black-tailedGodwits(Limosalimosa)andRedshanks(Tringa fullygrownCornCrakes(Crexcrex),Dunlin(Calidris Swallows(Hirundorustica). HerringGUlls havealsobeen seenattacking migratingMerlins COlumbarius) (Dyck1969),Jays(Garrulusglandarius)(Edholm1978),andthrushes(Turdinae)(Hobbs1959,Karmborg1979),aswellasavarietyofothermigrants(MacDonaldandMason1973,G.Taylor1979).G.Taylor(1979)thoughtsomeofthisbehaviorcouldbeinterpretedasplay,ratherthanasseriousattemptstoobtainfood.MannnalseateninNorthAmericaincludeNorwayrats (Hattus norvegicus),meadowvoles(Microtuspennsylvanicus)(Ludwig1966.Houde1977b),redsquirrels (Mendall1939),andanunidentifiedinsectivore(Mills1957).MannnalianpreyeatenintheOldWorldincludemoles(Talpaeuropaea)(Harris1965),twoweasels(Mustelanivalis,M.erminea),hares (LepUSeuropaeus) (authorscitedinHarris1965),rabbits cuniculliS) (Harris1965),commonvoles(Microtusarvalis),commonhamsters(Cricetuscricetus)(Spitzer1976),watervoles large-toothed baC'k'ed voles(Clethrionomysrufocanus),rootvoles(Microtusoeconomus)(Bianki1967),Norwaylemmings(Lemmuslemmus)(Belopol 1967),shrews(Melville1974; ::lorex 1970]),andmice(Melville1974).OthervertebratepreyeateninNorthAmericaincludenorthernwatersnakes(Nerodiasipedon)(Goldman1971)andmudpuppies(Necturusmaculosus)(Ludwig1966).HerringGullshavealsobeenreportedeatingfrogs (Hana temporaria),toads(Bufobufo),lizards(Lacertasp.).andanOldWorldviper(Viperusberus) (HarriST965 andauthorscitedtherein).---WesummarizebelowsomeoftheextensiveliteratureonthefoodhabitsoftheHerringGullinNorthAmerica:Manitoba J.
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HerringGulledulis)werethemostimportantiteminthedietinMayandJune;capelin(Mallotusvillosus)wereeatenmostofthesummer.PelletscontainedmanyremnantsofLeach'sStorm-PetrelsandtheeggsofCommonMurres.ThepelletswereregurgitatedbybothHerringandGreatBlack-backedGulls,anditisnotclearfromThrelfall'saccountwhetheroneorbothspecieswerelargelyresponsibleforthispredation.HaycockandThrelfall(1975)gaveamoredetailedaccountofthefoodseatenbythesegullsinasubsequentstudy(1969-71)madeinthesamearea.NewBrunswickPimlott's(1952)reportonthefoodhabitsofHerringGullsintheGrandMananArchipelagoin1949wasbasedonfieldobservations,foodregurgitatedbyyoungbirds,andfooditemsfoundinthestomachsofadults.Of 12stomachs,9heldAtlanticherring(Clupeaharengus)andoneeachcontainedAtlanticmackerel(Scomberscombrus),pollock(Pollachiusvirens),andshrimp.Of40regurgitations,34wereherring,twoeachweremacker er-and pollock,andoneeachcontainedinsectsandarockgunnel(Pholisgunnellus).OtherfoodseatenbyHerringGullsinthisareaincludedfishoffalusedasfertilizeronfieldsandfounddumpedintoharbors,crowberries(Empenigrum),andseaurchins(Echinoidea).--Mills(1957)examinedthestomachcontentsof36HerringGullscollectedoverwaterinAugustandSeptember1956intheNorthumberlandStraitbetweenNewBrunswickandPrinceEdwardIsland.Foodfoundmostfrequentlyin31stomachsthatcontainedfoodwereinsects(in67.7%ofthestomachs),crustaceans(35.5%),andfish(25.8%).Fourstomachscontainedtheremainsofbirds,twocontainedsquid,andonecontainedasmallmammal.Millsidentifiedfewofthestomachcontentstospecies,reportingonlyonefish(Atlanticcod[GadusmorhuaJ).Insectseaten,inorderoffrequencyofoccurrenceinthestomachs-, were Coleoptera(45.1%),Diptera(16.1%),Hymenoptera(12.9%),andLepidoptera(9.7%).Tolonen(1976)reportedthatHerringGullsdroppedandfedonsnails(Buccinumundatum)nearBlack'sHarbor.MichiganFishmadeupmostofthedietofHerringGullsonLakesHuronandMichiganin1963-1965(Ludwig1966).LudWigestimatedthatalewives(Alosapseudoharengus)madeup80%ofthediet;rainbowsmelt(Osmerusmordax)werealsoimportant.Otherspeciesoffisheaten(yellowperch,sunfish[Lepomissp.J,androckbass[AmbloplitesrupestrisJ)accountedforasmallproportionofthediet.AtGooseIslandinnorthernLakeHuron,Chamberlin(1975)discoveredthatfishfoundnearHerringGullnestswerelargelysmeltandalewives,althoughminnows,perch,bass,andbullheadswerealsofound.HerringGullsalsogorgedthemselvesonhatchingmayflies(Ephemeroptera).MaineMendall(1935)reportedonthestomachcontentsof838HerringGullscollectedalongthecoastofMainein1933-34.Mollusks,fish,andcrustaceanswerefoundmostfrequently;forbothyearscombinedthesefoodswerefoundin42.8,36.3and26.8%ofthestomachs,respectively.Vegetation,echinoderms,younglobsters,insects,andshrimpwerefoundinmorethan10%ofthestomachs.Birdsandwastewereinfrequentlyencountered.Fewofthefoodswereidentifiedtospecies,butMendall(1935)remarkedthatmostofthefishwereherringandthatafewpollockandmackerelweretaken.Mendallalso210

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HerringGullreportedthatHerringGullsoccasionallyateberries,primarilyblueberries(Vacciniumsp.).Mendall(1939)reportedonthestomachcontentsof62HerringGullscollectedinland.Mostoftheirfoodconsistedoffish(76.9%bybulk);otheranimals,vegetablefood,andrefusemadeup6.9,8.0,and8.2%ofthediet,respectively.Themostimportantfoodfishwerewhiteperch(Moroneamericana),pumpkinseed(Lepomisgibbosus),redbreastsunfish(Lepomisauritus),andyellowperch,whichtogethermadeupmorethanhalfofthediet.Otherfishidentifiedweresmallmouthbass(Micropterusdolomieui),whitesuckers(Catosomuscommersoni),fallfish(Semotiluscorporalis),goldenshiner(Notemigonus crysOIffiiCas), commonshiners(Notropiscornutus),andtroutorsalmon(Salmonidae).Drury(1963a)reportedthatHerringGullsintheIslesofShoalsareaofMaineandNewHampshirefedprimarilyonsmallfishandinvertebrates,andthatincubatingadultsatemoregarbagethantheirchicksdid.BothadultandyoungHerringGullsinthePenobscotBayareafedmainlyonrefuse,whichtheyobtainedfromdumpsandchicken-processingplantsclosetotheircolonies(Hunt1972a).Thesesourcesaccountedfor42-61%(byvolume)ofallfoodbroughttochicksin1969-1970.Mostoftherefuseconsistedofgarbage,chickenremains,andfishwastesEarthworms(Annelida)wereimportantatonecolony,butfishtakenfromthewildandfishofunknownoriginalsowereimportantatallcoloniesstudied.MassachusettsDrury(1963a)summarizedfoodhabitsatanumberofcoloniesinMassachusetts,showingthatfoodseatenbyadultsandyoungvariesconsiderablywithlocality.AdultsatCapeAnnfedonagreatvarietyoffoods,butfoodbroughttoyoungwasprimarilywastesfromhumanactivitites.Fishfedtoyounginthisareaincludedmackerel,pollock,whitingorsilverhake(Merlucciusbilinearis),andoceanperchorredfish(Sebastesmorinus).Gullsfromthisareafedmoreonstarfishthanthoseinotherregions.HerringGullsinBostonHarborfedextensivelyongarbagefromrestaurants,whereasthoseatMuskegetIslandfedmoreonnaturalfoods,includingawidevarietyofmarineinvertebrates.YoungbirdsinBostonwerefedextensivelyoninsectsandearthworms.YoungatcoloniesinNarragansettBaywerefedmostlyonnaturalfoods.IngolfssonandEstrella(1978)describedtheshell-crackingbehaviorofHerringGullsinsoutheasternMassachusettsnotingthatthemostcommonlyeatenmusselwasthebayscallop(Pectenirradians).ConnecticutTolonen(1976)foundthatthemainpreyofHerringGullsatMilfordwerefishinsummer,crabsinfall,andmolluscsinwinter.PreyremainsandregurgitatedboliatFortTrumbullfromNovemberthroughearlyDecembercontainedmostlycrabsofthegenusCancer,butgreencrabs(Carcinusmaenas),bluecrabs(Callinectessapidus)andbeachcrabs(Ovalipes oceII'ii'tiiST werealsorepresented.Molluscseatenincludedpelecypods(quahogs[Mercenariamercenaria],blueorediblemussels[Mytilusedulis],ribbedmussels[Modiolusdemissus],razorclams[Ensisdirectus]),andgastropods(thenorthernandlobedmoonshells[Lunatiaheros and""'POI'IIiIces duplicatusI,andthecommonandflatslipper shellS"["CrepidiiIilforniculata andC.planaJ)(Tolonen1976).Modioluswaseatenextensivelyinwinter,when gulls openedthembydroppingthemonice.Thesoft-shelledclam(Myaarenaria)wasalsoimportant.GullsatMilfordhave211

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HerringGullbeenobservedfeedingonhermitcrabs(Pagurus)andonthefleshofadeadcunner(Tautogolabrusadspersus).Americaneels(Anguillarostrata)(Tolonen1976)andstarfish(Asteriasforbesi)(ThomasandThomas1965)havealsobeenrecordedaspreyitemsinConnecticut.NewYorkHoude(1977b)reportedonthedietofHerringGullsatHicksIsland,LongIsland,inJuly1975,basinghisresultsonobservedfeedingsregurgitatedpellets,andstomachcontents.Awidevarietyofavianpreyandtwomammals(seeabove)werefoundinthediet,inadditiontothreemolluscs(oysters[Ostreavirginica],razorclams,andsurfclams[Spisulasolidissima]),twocrabs(amudcrab[Panopeusherbstii]andaspidercrab[Libiniaemarginata]),andfourfish(bluefish saltatrix],northernsearobin[Prionotuscarolinus],scup[Stenotomuschrysops],andAmericansandlance[Ammodytesamericanus]).Houdethought andscupprobably were obtainedfromlocaldumpsorasoffaldiscardedbyfishermen.NewJerseyFoodhabitsofHerringGullsalongtheNewJerseycoastvariedwithareaandseason(Tolonen1976).InDecember1970,naticidsnailspredominatedinthedietatSandyHookStateParkbutwereeatenuncommonlyelsewhere.InthePointPleasantarea,Spisulasolidissimawasimportant;MercenariamercenariawastheprimarypreyitematthesouthernendofLongBeachIsland.Fouryearslater,HerringGullsatSandyHookfedmostcommonlyonMercenaria,andSpissulawasstillthemostimportantfoodinthePointPleasantarea. replacedinimportanceatthesouthendofLongBeachIslandbyanotherpelecypod,thebayscallop(Aequipectenirradians).MarylandHerringGullsinlandatConowingoDamontheSusquehannaRiveronlyatefish.Identifiedpreyconsumedhereincludedyellowperch,Americaneels(AngUillarostrata),sunfish(Lepomissp.),andbullheadcatfish(Ictalurusspp.)(Tolonen1976).---NorthCarolinaTolonen(1976)foundthatthe wedge rangia(Rangiacuneata)wasparticularlyabundantamongthemolluscseatenbyHerringGullsinNorthCarolina.Hepointedoutthatthisclamandothermolluscsaredroppedinroadwaysinsuchabundancethatinthelate1960tsthestatespent$18,000-20,000annuallytokeepthehighwaysclearofthediscardedshells.InFebruary1971,Tolonen(1976)notedthatHerringGullsdroppedshellsonastretchofroadnearWhaleboneatarateequalto104molluscsperhourpermileofroad.FloridaKent(1981)reportedthatthefoodsdroppedandeatenbyHerring Point(FranklinCounty)inlatewinter1979consistedprimarilyofbivalves(77.1%byfrequency);decapodcrustaceans(13.9%)andechinoids(9.0%)werealsoeaten.Thefourmostimportantfoodswerethebayscallop,thepricklycockle(Trachycardiumegmontianum),aspidercrab(Libiniadubia),andaseaurchin(Lytechinusvariegatus)thatmadeup56.0,15.7.13.3,and9.0%ofthediet,respectively.Otherfoodseaten(noneofwhichmadeupmorethan2.4%ofthediet)werefourbivalves(thegreatheartcockle[Dinocardiumrobustum],Floridalucina[Lucinafloridana],sunrayvenus[Macrocallistanimbosa],andthesouthern campechensis])and212

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HerringGullthebluecrab.HerringGullspreferredthelargestavailablebivalves;theshelllengthofthesizeclassesofbayscallopandcockletakenmostfrequentlywere6-8em(2-3in)and3-6em(1-2in),respectively.Tolonen(1976)presentedextensivelistsoffoodeatenbyHerringGullsandGreatBlack-backedGullsalongtheAtlanticcoast.Thelists,however,donotdistinguishwhichspeciesatewhichfood.AllthefoodreportedfromFloridaprobablywaseatenbyHerringGulls.PreyitemsrecordedatMullet Key inlateMarch1970werethreegastropods(twowhelks[Busycyonspiratum,B.contrariumJandthecrownconch[Melongenacorona]),twopelecypods(thesouthernquahogandanunidentifiedrepresentativeofthefamilyArcidae),andaspidercrab(Libiniadubia).Tolonen(1976)alsorecordedHerringGullsfeedingonscrawledcowfish \LiiCtophrys quadricornis)atMullet Key, GulfBreezeandHighlandView,aswellasondeadbatfish(Ogcocephalussp.).IMPORTANTBIOLOGICALPARAMETERSEggLayingVariousstudiesindicatethategglayingbeginsinmid-orlateAprilorearlyMayandcontinuesuntillateMayorJune,oroccasionallyintoJuly.Muchofthelatelayingisapparentlycomposedofbirdsinitiatingsecondclutcheslaidafterthelossofthefirstone.StudiessummarizedbyErwin(1979a)indicatethatpeaklayingalongtheU.S.AtlanticcoastusuallyoccursfromlateApriltoearlyJune,virtuallythesameperiodasinmorenorthernportionsoftherange(Table27).MeanClutchSizeHerringGullsmaylay1-4eggs,butthemajorityusuallylay3(Table28).Mostofthestudieslistedbelowgivemeanclutchsizesbetween2.4and2.9.Variousfactorsaffectmeanclutchsize.MeanclutchsizeforHerringGullslayingearlyintheseasontendstobelargerthanforthoselayinglater(HaymesandBlokpoel1978a,HaycockandThrelfall1975,Table29).BurgerandLesser(1980)reportedthatHerringGullsnestingaloneinNewJerseyhadsignificantlylargermeanclutchsizes(mean=2.9)thandidthosenestingincolonies(mean=2.5).Pierotti(1982)showedthatHerringGullsnestingingrassymeadowsinNewfoundlandhadsmallerclutchsizesthanthosenestingonexposed,rocky,marineterracesoronturf-coveredmaritimeslopes.ClutchsizeofHerringGullsnestingclosertothepredatoryGreatBlackbackedGullwaslowerthanthosenestingfartheraway(Erwin1971,Table28).Thismaynotrepresentatruedifferenceinclutchsize,butmaybetheresultofpredationordisturbancebyGreatBlack-backedGulls.IncubationPeriodAtGullIsland,Newfoundland,themeanincubationperiod(+1.5days)for44firsteggswas29.4days;for48secondeggs,28.2days; and for28thirdeggs,27.1days(HaycockandThrelfall1975).AtGreatIsland,Newfoundland,Pierotti(1982)reportedthatin1977,firsteggsaveraged28-29daysandsecondandthirdeggsaveraged26days.Thefollowingyear,incubationperiodsforfirst,second,andthirdeggsaveraged29,27,and26days,respectively.Incubationperiodsat33nestsatKawinawLake,Manitoba,rangedfrom24to27days,averaging25.8(Vermeer1971a).Mean213

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HerringGullTable27.LayingperiodsreportedatsomeNorthAmericanHerringGullcolonies(a).ColonyLocalityLayingperiodandyearMANITOBAPeakSourceKawinawLakeearlyMay-earlyJune,197115-21MayONTARIOVermeer1973cBrothersIslandScotchBonnetIslandGraniteIslandGraniteIslandGullIslandGreatIslandmid-Apr.-lateMay,1973lateApr.-lateMay,1973Apr.-lateMay,1975earlyMay-lateMay,1976NEWFOUNDLANDlateApr.-lateMay,1971---------,1977-7822Apr.-3May----(b)11-15May6-10May10-14May20-24MayTeeple1977GilbertsonandHale1974aRyderandCarroll1978RyderandCarroll1978HaycockandThrelfall1975Pierotti1982(a)Thetermsearly,mid-,andlaterefertothefirst,mid-,andterminalthirdsofthemonth,respectively.Unlessotherwisestated,figuresareforthemainlayingperiodanddonotincludebirdsthatrenested.FiguresfromPierotti(1982)aretherangeofmediandatesoflayingofthefirsteggforthreenestingareasineachyear.(b)PeaklayingforfirstclutchesatScotchBonnetIslandin1975was21-30April;peaklayingin1975attwootherOntariocolonies(ChantryIslandandPortColborne)was23-25Apriland25April-5May,respectively(Gilmanetal.1977)incubationperiodsreportedfromtheOldWorld(Lockley1932,Paludan1951,Drent1970,MacRobertsandMacRoberts1972a,Parsons1972)aresimilar,rangingfrom26.4days(Lockley1932)to30.1days(Drent1970).HatchingSuccessDatafromanumberofstudies(Table30)indicatethatundernormalcircumstances,about75%oftheeggslaidinacolonyhatch.Thisamountstoabout2.3eggspernest,ontheaverage.Hatchingsuccessistypicallyhigherfornestsinitiatedearlyintheseason(MorrisandHaymes1977,BurgerandShisler1980b,Table31).Three-eggclutchestypicallyhatchahigherproportionofeggsthandotwo-eggclutches,andone-eggclutchesareusuallyleastsuccessful(Table32).Occasionally,two-eggclutchesaremoresuccessfulthanthree-eggclutches(KadlecandDrury1968a,MorrisandHaymes1977),andsomestudies(e.g.,Paynter1949,Vermeer1971a,RyderandCarroll214

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HerringGullTable28.MeanclutchsizesreportedfortheHerringGullinNorthAmerica(a).Mean NumberclutchofsizeclutchesLocalityandyearofobservationSourceMANITOBA2.932.672.63 2.86 2.82 2.532.432.642.842.992.982.582.882.91.2.842.852.972.752.912.45(b) 1.82 (b)2.96(b)2.24(b)2.39(b)2.862.68 2.882.76161 KawinawLake,1971(3)ONTARIO70PortColborne,1973(3-75%)108PortColborne,1974(3-64%)312PortColborne,1975(3-75%)60PortColborne,1976(3-77%)97ScotchBonnetIsland,1973(3-59%)53ScotchBonnetIsland,1975(3-62%)44BrothersIsland,1973(3)143ChantryIsland,1975(3-86%)100GraniteIsland,1975(3-96+%)100GraniteIsland,1976(3-96+%)50TorontoOuterHarbor,1977MICHIGAN287BellowsIsland,1965(3-77%)344PismireIsland,1965(3-87%)215GooseIsland,1972(3-86%)120GooseIsland,1973(3-86%)450SouthManitouIsland,1972650SouthManitouIsland,1973370SouthManitouIsland,1974474SouthManitouIsland,1975281SouthManitouIsland,1976466SouthManitouIsland,1977 421SouthManitouIsland,1978364SouthManitouIsland,1979823RogersCity,1975786RogersCity,1976868RogersCity,1977855RogersCity,1978WISCONSINVermeer1971aMorrisandHaymes 1977MorrisandHaymes1977MorrisandHaymes1977MorrisandHaymes1977GilbertsonandHale1974aGilmanet a1. 1977Teeple1977Gilmanet a1. 1977RyderandCarroll1978RyderandCarroll1978HaymesandBlokpoel1978aLudwigandTomoff1966LudwigandTomoff1966Chamberlin1975Chamberlin1975Shugart1977bShugart1977bShugart1977bSouthernetal.1980Southernetal.1980Southernetal.1980Southernetal.1980Southernetal.1980Patton1979Patton1979Patton1979Patton19792.49(c)100LittleSisterIsland,1964(3-61%)Keith1966a215

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HerringGullTable28.Continued.MeanclutchsizeNumberofclutchesLocalityandyearofobservationSourceNEWFOUNDLAND2.70(d)GullIsland,1970HaycockandThrelfall19752.73109GullIsland,1971HaycockandThrelfall19752.44-(e,n303GreatIsland,1976-1978Pierotti19822.652.27-(e,g)343GreatIsland,1976-1978Pierotti19822.672.16-(e,h)280GreatIsland,1976-1978Pierotti19822.51NEWBRUNSWICK2.47100KentIsland,1947(3-54%)Paynter1949MAINE2.62(b)47colonies,1944(3-65%)GrossinKadlecandDrury1968a2.59(b)9colonies,1965(3-60%)KadlecandDrury1968a2.6899 3colonies,1968Hunt1972a2.492944colonies,1969Hunt1972a2.514304colonies,1970Hunt1972aMASSACHUSETTS2.55(b)7colonies,1964(3-65%)KadlecandDrury1968a2.60(b)6colonies,1965(3-70%)KadlecandDrury1968a2.45567 EggRock,1962(3-58%)Keith1966a2.83249Coatue.1963(3-83%)KadlecandDrury1968a lUiODE ISLAND2.41275BlockIsland,1963(3-55%)KadlecandDrury1968a2.84258BlockIsland,1965(3-84%)KadlecandDrury1968a2.89266BlockIsland,1966(3-88%)KadlecandDrury1968a2.85370SandyPoint,1963(3-85%)KadlecandDrury1968a2.88(i)48SandyPoint,1969Erwin19712.69(j)48SandyPoint,1969Erwin1971216

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HerringGullTable28.Concluded.Mean NumberclutchofsizeclutchesLocalityandyearofobservationSourceNEWJERSEY2.65 2.76(k)124IslajoIsland,1975808colonies,1977Burger1977aBurgerandShisler1980b(a)Someofthesefiguresarederivedfromcountsofcontentsofnestsduringshorttermvisits.Suchreportsareoftenmisleadinganddonotgiveanentirelyadequateassessmentofclutchsizebecause(1)clutchesmaybeincompleteatthetimeofthevisit;(2)lossofeggsiswidespreadandsometimesoccursduringlaying(KadlecandDrury1968a);and(3)HerringGullsoftenbuildneststhatneverhold clutches. (Paynter1949,KadlecandDrury1968a).Generallyspeaking,themoredetailedthestudy,thecloserthemeanclutchsizeapproaches3(KadlecandDrury1968aandauthorscitedtherein).Insomeinstanceswehavecalculatedclutchsizefromdatagivenintheoriginalpapers.Whenpossible,welistedinparenthesesthemostfrequentlyobservedclutchsizeandthepercentageofneststhatcontainedthisnumberofeggs.(b)Emptynestsexcluded.(c)Meanclutchsizeisfromtheearliestnestcount;Keithbelievedthatmeanclutchsizeinthiscolonyatthemediandateofclutchinitiationwouldhavebeenbetween2.90and3.00.(d)Figureishighestaverageclutchsizeobtainedonanyonevisit.,(e)Nisthecombinedsampleforthreeyears.(f)Figuresarerangeofvaluesinrockyhabitat.(g)Figuresarerangeofvaluesonturf-coveredslopes.(h)Figuresarerangeofvaluesingrassymeadows.(i)MeanclutchsizeforbirdsnestingfarfromGreatBlack-backedGulls.(j)MeanclutchsizeforbirdsnestingnearGreatBlack-backedGulls.(k)Thenumberofclutchesdoesnotincludenestsinwhichnoeggswerelaid.1978)havefoundnorelationshipbetweentimeoflayingandhatchingsuccess.Disturbancebymanorotheranimalsalsoinfluenceshatchingsuccess.HerringGullslayingearlyatacolonyinRhodeIslanddidnothaveasignificantlyhigherhatchingsuccess(87%)thanthoselayinglateintheseason(78%)whentheynestedfarfromGreatBlack-backedGulls,buthatchingsuccessofearlynesters(77%)wassignificantlyhigherthanthatoflatenesters(65%)whenthetwospeciesnestedtogether(Erwin1971).Hunt(1972a)reportedthatcoloniesdisturbedbypicnickersinMainehadasignificantlylowerhatchingsuccessratethanthosethatremainedundisturbed.217

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HerringGullTable29.ComparisonofmeanclutchsizesforearlyandlateclutchesinNorthAmericanHerringGulls.LocalityYearNumber NumberEarlyofLateofclutchesclutchesclutchesclutchesSourceScotchBonnetIsland1973(c)2.57ScotchBonnetIsland1975(d)2.57ChantryIsland1975(d)2.91TorontoOuter Harbor-Y977(e)2.77KawinawLakeGooseIsland___ -'1c.=9-=-7-=-1.--:('!l_ 3.001973(b)2.891022.7482Vermeer1971a552.8451Chamberlin1975792.3318GilbertsonandHale1974a421.9111Gilmaneta1.19771262.2417Gilmanetal.1977222.4328 Haymes-and Blokpoel1978a(a)Earlyclutchesarethosebegunpriorto17May,themeandateofclutchinitiation;lateclutchesarethosebegunthereafter.(b)Nestswitheggshatchingbefore1Junewereconsideredearlynests.(c)Lateclutcheswerethosebegunafter22 Mayandwereconsideredbytheauthorstoberenestingattempts.(d)Lateclutcheswerethosethatwereconsideredsecondnestingattempts.(e)Lateclutcheswerethoseinitiatedafter7 May. AgeatFledgingAgeatfirststrongflightfor12youngHerringGullsatGullIsland,Newfoundland,rangedfrom42to48days(mean = 45.2days)(HaycockandThrelfall1975).Paynter(1949)estimatedafledgingtimeofabout43days.Theaverageageoffledgingfor6chicksatGray'sRock,Massachusetts,wasabout51days,withtheearliestfledgingatanageof35-44daysandtheoldestat56-61days(Kadleceta1.1969).Goethe(1956ainKadlecetal.1969)reportedarangeoffledgingagesforEuropeanbirdsof43-62days.FledgingSuccessInsuccessfulGreatLakescolonies,eachpairofHerringGullsproducedabout1.4-1.5fledglings,amountingtoabout46fledglingsper100eggslaid,or58-73fledglingsper100eggshatched(Gilmaneta1.1977).Itisdifficulttocomparestudiesbecauseofthedifferencesinterminologyandmethodsofcalculation,butTable33listsproductivityfiguresforanumberofstudiesofNorthAmericanHerringGullcolonies.ProductivityofyounginNorthAmericanHerringGullcoloniesvarieswidely.Particularlylowproductivityisoftenassocatiatedwithpesticidepollution.KadlecandDrury(1968a)concludedthatnormalproductivityinHerringGullcoloniesisapproximately0.8-1.4youngperpair.Earlyneststendtofledgemoreyoung(Pierotti1982).218

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HerringGullTable30.RatesofhatchingsuccessreportedfortheHerringGullinNorthAmerica.Vermeer1971aSourcesites)MorrisandHaymes1977sites)MorrisandHaymes1977sites)MorrisandHaymes1977MorrisandHaymes1977GilbertsonandHale1974aGilmanetal.1977Teeple1977Gilmanetal.1977RyderandCarroll1978RyderandCarroll1978HaymesandBlokpoel1978aONTARIOMANITOBALocalityandyearofobservationKawinawLake,1971PortColborne,1973(twoPortColborne,1974(twoPortColborne,1975(twoPortColborne,1976ScotchBonnetIsland,1973(a)ScotchBonnetIsland,1975Brother'sIsland,1973(b)ChantryIsland,1975GraniteIsland,1975GraniteIsland,1976TorontoOuterHarbor,1977PercentofeggslaidNumberthatofhatchedeggs69.933045.5,54.555,13241.0,53.183,14746.7,61.8105,17854.216616.3245199624.11167240679.629984.229865.1129MICHIGAN28.3828BellowsIsland,1965LudwigandTomoff196680.0+1,000PismireIsland,1965LudwigandTomoff196680.4215GooseIsland,1972Chamberlin197568.7611GooseIsland,1973Chamberlin197547.9119SouthManitouIsland,1972Shugart1977b68.7179SouthManitouIsland,1973Shugart1977b56.1148SouthManitouIsland,1974Shugart1977b32.31,160SouthManitouIsland,1975Southernetal.198010.0511SouthManitouIsland,1976Southernetal.198061.81,390SouthManitouIsland,1977Southernetal.198014.4945SouthManitouIsland,1976Southernetal.198033.3870SouthManitouIsland,1979Southernetal.1980WISCONSIN41LittleSisterIsland,1964Keith1966a219

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HerringGullTable30.Continued.PercentofeggslaidthathatchedNumberofeggsLocalityandyearofobservationNEWFOUNDLANDSource72.974.8-78,877865.1-80.489367.8-78.6672GullIsland,1971GreatIsland,1976-78(c)GreatIsland,1976-78(d)GreatIsland,1976-78(e)NEWBRUNSWICKHaycockandThrelfall1975Pierotti1982Pierotti1982Pierotti198271.38047.231.0 37.068.874.779.3 75.4678251.82472569929443044258 266121 4848328 Ken tIsland,1947KentIsland,1976MAINE3colonies,19684colonies,19694colonies,1970MASSACHUSETTSCoatue,1963RHODEISLANDBlockIsland,1965BlockIsland,1966SandyPoint,1963SandyPoint,1969(f)SandyPoint,1969(g)NEWJERSEYIslajoIsland.1975Paynter1949Gilmanetal.1978Hunt1972aHunt1972aHunt1972aKadlecandDrury1968aKadlecandDrury1968aKadlecandDrury1968aKadlecandDrury1968aErwin1971Erwin1971Burger1977a(a)FigurescalculatedfromTable1ofthecitedsourceandincludebothfirstandsecondnestingattempts.(b)DataarecalculatedfromTable3ofthecitedsourceandincludebothfirstandsecondclutches,butonlyclutchescontainingtwoandthreeeggs.(c)Figuresareforbirdsnestinginrockyhabitat.(d)Figuresareforbirdsnestingonturf-coveredslopes.220

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HerringGullTable30Concluded.(e)Figuresareforbirdsnestingingrassymeadows.(f)DataareforHerringGullsnestingnearGreatBlack-backedGulls.(g)DataareforHerringGullsnestingfarfromGreatBlack-backedGulls.Table31.ComparisonofhatchingsuccessforearlyandlatenestsofNorthAmericanHerringGulls.PercentPercentofeggsofeggshatched,hatched,earlyNlateNLocalityYearnests(eggs)nests(eggs)SourceBellowsIsland1965(a)28.48285534?LudwigandTomoff1966KawinawLake197170.432162.2151Vermeer1971aGooseIsland1973(b)78.615975.9145Chamberlin1975BrothersIsland1973(c)25.08421.932Teeple1977ScotchBonnetIsland197317.220311.942GilbertsonandHale1974a J:>cotch BonnetIsland1975(d)22108a21Gilmaneta1.1977ChantryIsland1975(d) 76-----m-25 39 Gilman et aL-f977 TorontoOuter1977(e)80.36151.5 68-Haymes-and BlokpoelHarbor1978aPortColborne(CF)1973(f)42.32648.330MorrisandHaymes1977PortColborne(CF)1974(f)45.96127.322MorrisandHaymes1977PortColborne(CF)1975(f)43.86441.251MorrisandHaymes1977PortColborne(L)1973(f)65.77041.962MorrisandHaymes1977PortColborne(L)1974(f)64.911412.133MorrisandHaymes1977PortColborne(L)1975(f)68.010054.523MorrisandHaymes1977PortColborne(L)1976(f)56.811149.155MorrisandHaymes1977(a)FiguresareestimatesbasedonintermittentvisitsandrepresentwhatLudwigandTomoffbelievedtobeinitialandsecondnestingattempts.(b)Earlynestswerethosethathadhatchedeggsby1June.(c)Earlyandlatenestsarethosebelievedtobefirstandsecondclutches.(d)Theauthorsconsideredlatenestssecondnestingattempts.(e)Latenestswerethoseinwhicheggswerefirstlaidafter7 May.(f)CF=CanadaFurnacecolony,L = Lighthousecolony.221

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HerringGullTable32.Comparisonofhatchingsuccessinthree-andtwo-eggclutchesinsomestudiesofNorthAmericanHerringGulls.HatchingSuccessLocalityYearThree-eggNTwo-eggNclutches(eggs)clutches(eggs)SourceKentIslandSandyPointCoatueBlockIslandBlockIslandGullIslandGooseIslandBrothersIslandBrothersIslandTorontoOuterHarborPortColborne(CF)PortColborne(CF)PortColborne(CF)PortColborne(L)PortColborne(L)PortColborne(L)PortColborne(L)19471963 1963 196519661971(a)19721973(b)1973(b)19771973(c)1974(c)1975(c)1973(c)1974(c)1975(c)1976(c)80.2%76.867.675.379.3 76.481.727.2 22.280.547.645.058.749.552.565.757.0162324102648 69917155266188742758463114144 13856.4%57.675.070.280.334.576.016.721.438.266.723.531.370.057.557.165.778Paynter194926KadlecandDrury1968a20KadlecandDrury1968a84KadlecandDrury1968a66KadlecandDrury1968aHaycockandThrelfall197550Chamberlin197518Teeple1977 14Teeple1977 34 HaymesandBlokpoel1978aMorrisandHaymes197718MorrisandHaymes197716MorrisandHaymes197730MorrisandHaymes197720MorrisandHaymes197714MorrisandHaymes197718MorrisandHaymes1977(a)Figurefortwoeggclutchessubsumeshatchingsuccessforoneeggclutchesaswell.(b)Thefirstsetoffiguresisforfirstnestingattempts;thesecondsetisforsecondnestingattempts.(c)CF=CanadaFurnacecolony.L=Lighthousecolony222

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HerringGullTable33.ProductivityinNorthAmericanHerringGullcolonies(a).Numberofyoungproducedperpairorpernest0.210.060.080.35-0.520.120.05-0.060.150.100.06-0.180.480.32-0.480.651.410.791.321.381.551.48(b)(b)(b)(b)(b)(d)(b)(b)(d)(c)(c,e)(c,O(b)(c)(b)(b)(b)(b)LocalityandyearofobservationONTARIOSnakeIsland,ELO,1972Presqu'ilePark,NLO,1972BlackAntIsland,SLR,1972BigChickIsland,WLE,1973ScotchBonnetIsland,NLO,1972ScotchBonnetIsland,NLO,1973ScotchBonnetIsland,NLO,1975BrothersIsland,ELO,1972BrothersIsland,ELO,1973PortColborne,ELE,1973PortColborne,ELE,1974PortColborne,ELE,1975PortColborne,ELE,1975PortColborne,ELE,1976GraniteIsland,NLS,1975GraniteIsland,NLS,1975GraniteIsland,NLS,1976ChantryIsland,ELH,1975MICHIGANSourceGilbertson1974aGilbertson1974aGilbertson1974aGilbertson1974aGilbertson1974aGilbertsonandHale1974aGilmaneta1.1977Gilbertson1974aTeeple1977MorrisandHaymes1977MorrisandHaymes1977MorrisandHaymes1977Gilmaneta1.1977MorrisandHaymes1977RyderandCarroll1978Gilmaneta1.1977RyderandCarroll1978Gilmaneta1.19770.31-0.421.22-1.301.960.650.510.390.010.001.380.000.00(b,d)(b,d)(c)(b)(b,O(b,O(b,O(b,O(b,f)(b,O(b,OBellowsIsland,GTB,1965PismireIsland,NLM,1965GooseIsland,NLH,1973SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,SouthManitouIsland,NLM,WISCONSIN1972 1973 1974 19751976197719781979LudwigandTomoff1966LudWigandTomoff1966Chamberlin1975Schugart1977bSchugart1977bSchugart1977bSouthernetal.1980Southernetal.1980Southerneta1.1980Southernetal.1980Southerneta1.19800.3-0.4(b)LittleSisterIsland,GB,1964Keith1966a223

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HerringGullTable33.Continued.Numberofyoungpro-ducedperpairorpernest0.9(c)1.32-1.84(c,g)1.58-1.87(c,h)1.03-1.28(c,i)1.06(c)0.80(c,j)0.34(c,j)0.28(c,j)1.09(c)1.02(c)0.93(c)1.42(c)1.35(c) 1. 79(k)2.44(1)1.43LocalityandyearofobservationNEWFOUNDLANDGullIsland,1971GreatIsland,1976-78GreatIsland,1976-78GreatIsland,1976-78NEWBRUNSWICKKentIsland,1947MAINEcolonies,1968colonies,1969colonies,1970MASSACHUSETTSCoatue,1963RHODEISLANDBlockIsland,1963BlockIsland,1965BlockIsland,1966SandyPoint,1963SandyPoint,1969SandyPoint,1969NEWJERSEYcolonies,1977SourceHaycockandThrelfall1975Pierotti1982Pierotti1982Pierotti1982Paynter1949Hunt1972aHunt1972aHunt1972aKadlecandDrury1968aKadlecandDrury1968aKadlecandDrury1968aKadlecandDrury1968aKadlecandDrury1968aErwin1971Erwin1971BurgerandShisler1980ba)Abbreviationsafterlocalitiesusedinthistableareasfollows:ELE,easternLakeErie;ELH,easternLakeHuron;ELO,easternLakeOntario;GB,GreenBay,northernLakeMichigan;GTB,GrandTraverseBay,northernLakeMichigan;NLO,northernLakeOntario;NLH,northernLakeHuron;NLM,northernLakeMichigan;NLS,northernLakeSuperior;WLE,westernLakeErie;SLR,St.LawrenceRiver.224

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HerringGullTable33.Concluded.b)Figuresrepresentthenumberofyoungproducedperpair.c)Figuresrepresentthenumberofyoungproducedpernest.d)Figuresrepresentestimatesofminimumandmaximumproduction,respectively.e)Figuresaretherangeofvaluesfortwocolonies.f)Figuresareforyearswhenproductionwasaffectedbyaresidentpopulationofredfoxes(Vulpesvulpes).g)Figuresarerangeofvaluesinrockyhabitat.h)Figuresarerangeofvaluesonturf-coveredslopes.i)Figuresarerangeofvaluesingrassymeadows.j)Huntconsideredanychickthatreachedaweightof500gashavingfledged.k)FiguresareforbirdsnestingnearGreatBlack-backedGulls,arerecalculatedfromTable1ofthecitedsource,andrepresentthenumberofyoungproducedpernest.Erwin(1971)listed1.76youngproducedperpair.1)DataareforbirdsnestingfarfromGreatBlack-backedGulls,arerecalculatedfromTable1ofthecitedsource,andrepresentthenumberofyoungproducedpernest.Erwin(1971)listed2.39youngproducedperpair.DisturbancebyhumansisoftenamajorfactorinthefailureofHerringGullcolonies(Burger1981a).However,Hunt(1972a)foundnosignificantdifferenceinproductivitybetweendisturbedandundisturbedcoloniesinMaine.Huntpointedoutthatcoloniesnearfoodsourcesproducedsignificantlymoreyoungthanmoredistantcolonies.Nestinghabitatmayalsorelatetofledgingsuccess.Pierotti(1982)reportedthatHerringGullsnestingonexposedrockyterracesandonturf-coveredslopesatGreatIsland,Newfoundland,consistentlyfledgedmoreyoungthandidthosenestingingrassymeadows.Hethoughtthatthelowersuccessingrassymeadows mayhavebeenduetopredationbytheGreatBlack-backedGullsthatpreferredthisareaasnestinghabitat.MortalityofEggsandYoungProbablythetwomajorcausesofnestingfailureinHerringGullsareenvironmentalcontaminationbypollutantsandfailuresrelatedtodisturbance,eitherbyhumansorbyintroducedpredators.NestingsuccesshasbeengreatlyreducedatanumberofcoloniesontheGreatLakesinrecentyears,largelybecausehatchingfailuresattributedtopesticides(Gilmanet al. 1977,Teeple1977).Withrecentdecreasesinorganochlorinecontamination,reproductivesuccessinmanyareashasreturnedtonormal(Weselohet al. 1979).Foxes(Vulpesvulpes)andothermammalianpredatorsalsotakeyounggulls,andsometimesarethereasonfewyoungfledge(Shugart1977b,Patton1979,Southernet al. 1980).Humandisturbanceofcoloniesandtheassociateddisruptionmaybe eitherbyresearchinvestigators(Burger1981a,Pierotti1982),byvandals,casualvisitorssuchascampers,boatersandpicnickers(ShugartinScharf1979),orevenbysupersonicaircraft(Burger1981d).EggsmaybeeatenbyHerringGulls(Burger1977a)andyoungbirdsmaybekilledandeatenbyneighboringgulls(Chamberlin1975).Youngoftenarekilled(Kadlecet al. 1969,Pierotti1982)whentheywanderfromtheirparents'territoryintoterritoriesofotherHerringGulls(HaycockandThrelfall1975,RyderandCarroll1978).OthersaretakenbyGreatBlack-backedGullsinsomecolo-225

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HerringGullnies(Paynter1949,Pierotti1982)andsome maybetakenbyotheravianpredators.ReportedorsuspectedavianpredatorsincludeCommonCrows(Corvusbrachyrhynchos)(Chamberlin1975),andGreatHornedOwls(Bubo virgIIiIaiiUs) (ShugartinScharf1979).--Exposuretoinclementweathermaycausethedeathofyoungbirds(Kadlecetale1969)andstormsmayresultinthelossofnestsfromflooding(Burger1977a,ShugartinScharf1979).Diseaseoccasionallyaccountsforsomedeaths,(Pierotti1982,ShugartinScharf1979),butapparentlyisseldomamajorfactorinnestingfailures.MostmortalityofHerringGullchickstakesplaceinthefirst10daysfollowinghatching(Kadlecetale1969,HaycockandThrelfall1975,RyderandCarroll1978).RenestingSomeHerringGullsrelayiftheylosetheirfirstclutches,buttheproportionthatdo,inrelationtovariouscircumstances,ispoorlyknown.Teeple(1977)believedthat13of34pairsrelaidonBrothersIslandnearKingston,Ontarioin1973.HaycockandThrelfall(1975)reportedthat7replacementclutchesatGullIsland,Newfoundland,wereinitiated12-32days(mean=12.6days,n=5)afterthelossofthefirstclutch.Parsons(1976a)gavetheintervalbetweenlossofthefirstclutchandrenestingas12-15days.Theintervalisslightlylongerforbirdsinitiallynestinglate,andforbirdswhoseclutchesaretakenafter15daysofincubation,andshortestforearlynesterslosingtheirclutchimmediatelyafterlaying.RenestingisfrequentintheLakeOntariocolonies,wherepesticidelevelsarehighandhatchingsuccessisverypoor(GilbertsonandHale1974a,Teeple1977).AgeatFirstBreedingChabrzykandCoulson(1976)foundthatmanyof1,151bandedHerringGullsfirstbredat5yearsofage.Somebredat4years,andnonebredearlier.Birdsbreedinginsubadultplumageweremorecommoningrowingcoloniesthaninstableorspace-limitedcolonies.MaximumNaturalLongevityAHerringGullbandedinNewBrunswickwasobtainedinMarYlandafterreachinga minimumageof27yearsand3months(Clappetale1982a).Apreviouslypublishedlongevityrecordof36years(Pettingill1967)wasanerror(JonkelandPettingill1974).WeightDataonweightsofNorthAmericanHerringGullsaregivenin SUSCEPTIBILITYTOOILPOLLUTIONHerringGullsseldomdiefromoiling,butoccasionalmortalityfromoilinghasbeenrecorded(Table35).Certainlytheirpopulationshavenotbeenaffectedbyoilingtotheextentofsomealcidpopulations(BourneandDevlin1969).Gull'swinteringoffshoremaybecontaminatedbyfloatingoilwhiletheyroostonthewateratnight.HerringGullswinteroffshoreingreatnumbersinmanyareas,includingthesoutheasternUnitedStates(Robertson226

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HerringGullTable34.Weight(ingrams)ofNorthAmericanHerringGulls(a).MeanNumber Weight Range weighed Sample andseasonAreaSource12321014-1618180ad.males,summer NewfoundlandThrelfallandJewer197813341222-15685ad.males,summerAlaskaUSNMspec.13911248-161418ad.males,summerAlaskaUSNM,WilliamsoncolI.1093-133632ad.malesNewfoundland Haycock andThrel-fall1975 999832-127478ad.females,summer NewfoundlandThrelfallandJewer1978937-11189ad.femalesNewfoundland Haycock andThrel-fall1975 11331050-128212ad.females,summerAlaskaUSNM,WilliamsoncolI.57-701215newlyhatchedyoung NewfoundlandPierotti1982347-484 56215dayoldyoung NewfoundlandPierotti1982 393 203-5831215dayoldyoungMassachusettsKadlecetal.1969 443 293-593 615dayoldyoungMassachusettsKadlecetal.1969776-98634230dayoldyoung NewfoundlandPierotti1982787553-1021 930dayoldyoungMassachusettsKadlecetal.1969 853 635-1071 6 30dayoldyoungMassachusettsKadlecetal.1969 982 640-1324 450dayoldyoungMassachusettsKadlecetal.1969 982 688-1276 450dayoldyoungMassachusettsKadlecetal.19699565-10524fresheggsNewfoundland Haycock andThrel-fall197588-9258-120404fresheggsNewfoundlandPierotti1982227

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HerringGullTable34.Concluded.a)Publishedweightsover100gramsareroundedtothenearestgram.Figuresforeggsand youngfromPierotti(1982)aretherangeofvaluesformeanstakeninthreehabitatsintwoyears.The numberofweightsgivenisthetotalnumberofbirdsuponwhichthesemeansarebased.Figuresforrangeforfresheggsareranges+2S.D.forthelargestandsmallestmeans,respectively.Figuresfor-rangefromKadlecetal.(1969)arethemean+1S.D.Thefirstsetoffiguresisforallchicksweighed.Thesecondis for allbelievedtohavefledged.and Mason1965).OiledHerring(andother)GullsarecommonlyseeninFloridaaroundsanitarylandfills(W.Hoffman,pers.comm.),butthesebirdsprobablybecomesmearedwithwastemotoroilwhiledigginginthelandfills.FilesintheBirdBandingLaboratory,Patuxent,Maryland,attributedthedeathof57ofthesegullstooiling,farmorethanforanyotherspeciesofLaruslisted.Mostofthebirdswerefoundinthenorthernportionsoftherange;onlyfivewerereportedfromthesoutheasternUnitedStates(fourfromTexas and onefromFlorida).HerringGullpopulationsarestableorincreasinginmostareasofNorthAmericaandintheOldWorld.NumberswinteringinthesoutheasternUnitedStatesareconsiderableandarelikelytoincreaseinthefuture.OildevelopmentinthesoutheasternUnitedStateswouldprobablynothaveasignficantimpactonoverallpopulationsofthisspecies.BIBLIOGRAPHY1983Baerends,G.P.and R.H.Drent(eds.).TheHerringGullanditsegg.PartII.Theresponsivenesstoegg-features.Behaviour82:'xiiiand 416pp.Bergman,G.1983.Populationdynamics,colonyformationandcompetitioninLarusargentatus,fuscusandmarinusinthearchipelagoofFinland.Ann.Zool.Fenn.19:143-164.Peakall,D.B.,D.S.Miller,andW.B.Kinter.1983.ToxicityofcrudeoilsandtheirfractionstonestlingHerringGulls--l.physiologicalandbiochemicaleffects.Mar.Environ. Kes. 8:63-71.Sibly,R.M.andR.H.McCleery.1983a.IncreaseofweightofHerringGullswhilefeeding.J.Anim.Ecol.52:35-50.1983b.ThedistributionbetweenfeedingsitesofHerringGullsbreed-----ingatWalneyIsland,U.K.J.Anim.Ecol.52:51-68.228

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HerringGullTable35.Numberofdeadbirdsand number andpercentageofdeadHerringGullsfoundaftermajoroilingincidents.AreaDatesNumberofoileddeadbirdsNumberofdeadHerringGullsPercentofHerringGullsSourceIslandBeach,NewJerseyJan.194592(a)2729.35Kramer and Kramer 1945PooleHarbour,Dorset,EnglandSoutheastKent,EnglandNortheastEnglandMedwayEstuary,Kent,EnglandTayEstuary,ScotlandJan.1961 433(b,c)wintersof1963-598(b)64to196"5-66Jan.1966805Sept.19662,748(b)Mar.-Apr.1,168(c)19681240 1516572.77Bourne1968a7.86Gibson 19661.86Parrack19676.00Bourne1968a0.60Greenwood and Keddie 1968NortheastBritainMartha'sVineyard,MassachusettsSanFranciscoBay,CaliforniaNorthernScotlandWaddensea, Den markFirthofClyde,Ayrshire,ScotlandJan.-Feb.1970Feb.1970Jan.1971 May-Jun. 1971 Dec. 1972Jan.197410,992(a,b)541(b)3,221(b,d)1,101(b)9,151(b)279(b)32412330.29 0.740.030.180.03l.88Greenwoodetale1971CSLP1971Smailetal.1972 Bourne1971aJoensenand Hansen 1977Lloydetale1974(a)Totalincludessomebirdsthatwerenotoiled.(b)Totalincludesonlythosebirdsidentifiedtospecies.(c)Thiscountisofbothdeadandaliveoiledbirds.(d)Thisfigurerepresentsbirdsbroughttocleaning/receivingstations.229

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HerringGullAdulrHerringGullinbreedingplumage.PhotographbyPatLynch.230

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HerringGull1982Becker,P.H.andM.Erdelen.1982.WindrichtungundVegetationsdeckungamnestderSilbermowe(Larusargentatus).[VegetationsurroundingHerringGulls'(Larusargentatus)nestsinrelationtowinddirection.]J.Ornithol.123:117130.[InGermanwithEnglishsummary.] Bergman,G.1982.Populationdynamics,colonyformationandcompetitioninLarusargentatus,fuscusandmarinusinthearchipelagoofFinland.Ann.Zool.Fenn.19:143-164.Blokpoel,H.and D.V.Weseloh.1982.StatusofcolonialwaterbirdsnestingonLittleGallooIsland,LakeOntario.Kingbird32:149-157.Coulson,J.C.,N.Duncan and C. Thomas.1982.ChangesinthebreedingbiologyoftheHerringGull(Larusargentatus)inducedbyreductioninthesizeanddensityofthecolony.J.Anim.Ecol.51:739-756.Coulson,J.CP.Monaghan,J.Butterfield,_N.Duncan, C-. S.Thomas andH.Wright.1982.Variationinthewing-tippatternoftheHerringGullinBritain.BirdStudy29:111-120.Gochfeld,M.andJ.Burger.1982b.BiologicalconcentrationofcadmiuminestuarinebirdsoftheNewYorkBight.ColonialWaterbirds5:116-123.Got mark ,F.1982.ColonialityinfiveLarusgulls:acomparativestudy.OrnisScand.13:211-224.Hanssen_,o.J.lations.1982.EvaluationofsomemethodsofcensusinglaridpopuOrnisScand.13:183-188.Holley,A.J.F.1982.Post-fledginginteractionsontheterritorybetweenparentsandyoungHerringGullsLarusargentatus.Ibis124:198-203.Lemmetyinen,R.,P.Rantamaki andA.Karlin.1982.LevelsofDDTand PCB'sindifferentstagesoflifecycleoftheArcticTernSternaparadisaeaandtheHerringGullLarusargentatus.Chemosphere11:1050-1068.Mineau,P.,G.E.J.Smith,R.MarkelandC.-S.Lam.1982. HerringGullsfromhatchingtofledging.J.FieldOrnithol.53:394-402.Morris,R.D.andM.J.Bidochka.1982.MateguardinginHerringGulls.ColonialWaterbirds5:124-130.Mudge,G.P.andP.N.Ferns.1982.Thefeedingecologyoffivespeciesofgulls(Aves:Larini)intheinnerBristolChannel.J.Zool.(Lond.)197:497-510.Parsons,K.C.1982.Nest-sitehabitatandhatchingsuccessofgulls.ColonialWaterbirds5:131-138.231

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Peakall,D.B.,D.J.Hallett,J.R. Bond, G.L.Foureman andD.S.Miller.1982.ToxicityofPrudhoeBaycrudeoilanditsaromaticfractionstonestlingHerringGulls.Environ.Res.27:206-215.Perry,P.1982.TheuseofgullnestsbyEiders.Brit.Birds75:360-365.Pierotti,R.1982.HabitatselectionanditseffectonreproductiveoutputintheHerringGullinNewfoundland.Ecology63:854-868.Southern,L.K.,S.R.Patton,andW.E.Southern.1982.NocturnalpredationonLarusgulls.ColonialWaterbirds5:169-172.Walker,D.G.1982.HerringGullkillingaBlack-headedGull.Brit.Birds75:580-581.Wesehloh,D.V.andP.Mineau.1982.Demography,productivity,andcontaminantlevelsinHerringGullsinLakeErie.(Abstractonly.)ColonialWaterbirds5:179.1981Amlaner,C.J.andD.J.McFarland.SleepintheHerringGull(Larusargentatus).Anim.Behav.29:551-556.Andrews, k. 1981.ThegullpopulationonMonomoyIsland,Massachusetts,USA.(Abstractonly.)ColonialWaterbirds4:194.Burger,J.1981a.cularlygulls.Effectofhumandisturbanceoncolonialspecies,partiColonialWaterbirds4:28-36.1981b.FeedingcompetitionbetweenLaughingGullsandHerringGullsatasanitarylandfill.Condor83:328-335.1981c.MovementsofjuvenileHerringGullshatchedatJamaicaBay-----Refuge,NewYork.J.FieldOrnithol.52:285-290.1981d.BehaviouralresponsesofHerringGullsLarusargentatusto-----aircraftnoise.Environ.Poll.24:177-184.1981e.-----conflict.OnbecomingindependentinHerringGulls:parent-youngAmer.Nat.117:444-456.Burger,J.andM.Gochfeld.1981a.Age-relateddifferencesinpiracybehaviouroffourspeciesofgulls,Larus.Behaviour77:242-267.1981b.UnequalsexratiosandtheirconsequencesinHerringGulls------(Larusargentatus).Behav.Ecol.Sociobiol.8:125-128.1981c.Discriminationofthethreatofdirectvs.tangentialapproachtothenestbyincubatingHerringLarusargentatusandGreatBlack-backedGullsLarusmarinus.J.CompoPhysiol.Psychol.95:676-684.232

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Herring GullCoulson,J.C.,N.Duncan,C.S.Thomas andP.Monaghan.1981.Anage-relateddifferenceinthebilldepthofHerringGullsLarusargentatus.Ibis123:499-502.DeWire, R.C.1981.BandinggullsatChimonIsland[Connecticut].Conn.Warbler1:17-18.Dijkeen,L.J.1981.HetverschilinkoptekiningbijnestjongenvanZilvermeeuw(Larusargentatus)enKleineMantelmeeuw(Larusfuscus).[ThedifferencesinheadmarkingsofjuvenalHerringGullLarusargentatusandLesserBlack-backedGullLarusfuscus.]Watervogels6:19-23.[InDutchwithEnglishsummary.]-----------Duncan,N.1981.TheAbbeysteadandMallowdalegullcolonybeforecontrol.BirdStudy28:133-138.Elliott,R.E.1981.OrbitalringcoloursofHerringGullsinBritain.BirdStudy28:66-68.Erwin,R.M.,J.GalliandJ.Burger.useinAtlanticcoastseabirds.1981.ColonysitedynamicsandhabitatAuk98:550-561.Ferns,P.N.andG.P.Mudge.1981.Accuracyofnestcountsata mixedcolonyofHerringandLesserBlack-backedGulls.BirdStudy28:244-246.Fox,G.A.,C.R.Cooper andJ.P.Ryder.Gullsbyusingexternalmeasurements.1981.PredictingthesexofHerringJ.FieldOrnithol.52:1-9.Grant, P, J.andR.A. Hume.1981.HerringGullwithcharacteristicsofMediterraneanraceinKent.Brit.Birds:350-351.Holley,A.J.F.1981.NaturallyarisingadoptionintheHerringGull.Anim.Behav.29:302-303.Kent,B.W.1981.PreydroppedbyHerringGulls(Larusargentatus)onsoftsediments.Auk98:350-354.King,B.1981.Inlandground-nestingbyHerringGulls.Brit.Birds74:264265.Mallalieu,M.and T.N.Holdge.1981.Yellow-leggedHerringGullsinNorthKent.Brit.Birds74:351-352.McGill-Harelstad,P.1981.TheeffectsofinterspecificcompetitiononreproductivesuccessofGreatBlack-backedandHerringGulls.(Abstractonly.)ColonialWaterbirds4:197.McGill-Harelstad,P.andT.L.Taigen.1981.Thermogeniccapacityofgullchicks.(Abstractonly.)Am.Zool.21:914.233

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HerringGullMineau,P.andD.V.C.Wesehloh.1981.Low-disturbancemonitoringofHerringGullreproductivesuccessontheGreatLakes.ColonialWaterbirds4:138-142.Morris,R.D.,M.C.Benkel,A.BiernackiandJ.M.Ross.1981.A newtransmitterassemblyforHerringGulls.J.FieldOrnithol.52:242-244.Nicholson,J.K.1981.Thecomparativedistributionofzinc,cadmium,andmercuryinselectedtissueoftheHerringGull(Larusargentatus).CompoBiochem.Physiol.68C:91-94.----V.1981.AninvestigationofcourtshipfeedinginHerringGullsLarusargentatus.Ibis123:218-223.Nilsen,J-E.lark.]1981.Gratrutarjagarsanglarka.VarFagelvarld40:66.[HerringGullschasingSky-Norstrom,R.J.,A.P.GilmanandD.J.Hallett.1981.TotalorganicallyboundchlorineandbromineinLakeOntarioHerringGulleggs,1977,byinstrumentalneutronactivityandchromatographicmethods.Sci.TotalEnviron.20:217-230.Oliver,P.J.1981.HerringGullwithyellowlegsnestinginLondon.Brit.Birds74:353.Portnoy,-J.W.andM.A. Soukup.1981.Gulluseofafreshwaterkettleholepond:quantificationandseasonalvariation.(Abstractonly.)ColonialWaterbirds4:200.Quinney,T.E.,B.N.MillerandK.R.S.Quinney.1981.GullsrobbingpreyfromGreatBlueHerons(Ardeaherodias).Can.Field-Nat.95:205-206.Redman,P.S.1981.YellOW-leggedHerringGullsinFranceandBritain.Brit.Birds74:349-350.Ryttman,H.and H.Tegelstrom.1981.LowdegreeofisozymevariationwithinandbetweenHerringGull(Larusargentatus),LesserBlack-backedGull(Larusfuscus)andtheirBritishandSwedishsubspecies.Hereditas94:143-148-.--Ryttman,H.,H.TegelstromandH.Jansson.1981.GenetiskaundersokningaravartsambandetmellangratrutLarusargentatusochsilltrutLarusfuscus.[GeneticstudiesoftherelationshipbetweenHerringGullLarus argentatus andLesserBlack-backedGullLarusfuscus.]Var 40:239-248.[InSwedishwithEnglishsummary-.-]----Snell,R.R.1981.HerringGullattacksandeatsadultmaleOldsquaw.WilsonBull.93:110-111.234

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HerringGullStanley,P.I.,T.Brough,M.R.Fletcher,N.HortonandJ.B. A.Rochard.1981.TheoriginsofHerringGullswinteringinlandinsouth-eastEngland.BirdStudy28:123-132.Treleaven,R.B.1981.PeregrinefeedingHerringGullchicktoyoung.Brit.Birds74:97.Triplet,P.1981.Notesaproposdel'actionpredatriceduGoelandargentesur1'Anguille.NosOiseaux36:169.[InFrench.]Vinicombe,K.1981.Yellow-leggedHerringGullsinBritain.Brit.Birds74:352-353.Weber,J. W. 1981.TheLarusgullsofthePacificNorthwest'sinterior,withtaxonomiccomments onseveralforms.(PartII--conclusion.).ContinentalBirdlife2:74-91.Witt,H.-H,J.Crespo,E.deJuana,andJ.Varela.1981.ComparativefeedingecologyofAudouin'sGull(Larusaudouinii)andtheHerringGull(L.argentatus)intheMediterranean.Ibis123:519-526.-1980Andrle,R.F.1980.L.argentatus.ThreemoreprobablehybridsofLarushyperboreusandWilsonBull.92:389-393.Braun,B.M.,P.A.HeinzandG.H.Heinz.onRed-breastedMerganserducklings.1980.HerringGullpredationWilsonBull.92:403.Burger,J.1980a.NestingadaptationofHerringGull(Larusargentatus)tosaltmarshesandstormtides. BioI. Behav.5: 1980b.Territorysizedifferencesinrelationtoreproductivestage-----andtypeofintruderinHerringGulls(Larusargentatus).Auk97:733-741.Burger,J.andF.Lesser.1980.NestsiteselectioninanexpandingpopulationofHerringGulls.J.FieldOrnithol.51:270-280.Burger,J.andJ.Shisler.1980b.TheprocessofcolonyformationamongHerringGullsLarusargentatusnestinginNewJersey.Ibis122:15-26.Burger,J.,M.Fitch,G.Shugartand W. Werther.1980.PiracyinLarusgullsatadumpinNewJersey.Proc.1979Conf.ColonialWaterbirdGroup3:87-98.Camberlein,G.1980.Methodesd'effraymentduGoelandargenteappliqueesalaprotectiondelamytiliculturedanslesCotesduNord.[BirdscaringawaymethodsagainstHerringGullsandtheirprotectiveeffectsonthecultureofmusselsintheCotesduNordregion.]Bull.Mens.Off.Nat1-ChasseNo.Spec. Sci. Tech.(Paris)pp.261-267.[InFrenchwithEnglishsummary.]235

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HerringGullCamberlein,G.andD.Flote.1980.DynamiqueetgestiondelapopulationdeGoelandargente(Larusargentatus)enBretagne.(Abstractonly.)OiseauRev.Fr. 273-275.Cambi,D.1980.AccertamentoeconfermadellanidificazionedelGabbianoreale(Larusargentatus)suILagodiGarda.Notapreliminare.[ConfirmationofHerringGulls(Larusargentatus)nestingon LakeGarda.Preliminarynote.]Riv.Ital.Ornitol.50:19-25.[InItalianwithEnglishandFrenchsummaries.]Coulson,J.C.,N.Duncan andP.Monoghan.1980.VariationinHerringGulls.Brit.Birds73:230-231.Dunn, E.H.andI.L.Brisbin,Jr.1980.Age-specificchangesinthemajorbodycomponentsandcaloricvaluesofHerringGullchicks.Condor82:398-401.Erwin,R.M.1980b.Breedinghabitatusebycoloniallynestingwaterbirdsintwomid-AtlanticU.S.regionsunderdifferentregimesofhumandisturbance.BioI.Conserv.18:39-51.Fitch,M.1980.Monogamy,polygamy,andfemale-femalepairsinHerringGulls.Proc.1979.Conf.ColonialWaterbirdGroup3:44-48.Goethe,F.1980.Lebens-undFortpflanzungsalterbeiHeringsmowenLarusf.fuscusinder undbeiSilbermowenL.a. argentatus:im Freiland.[Age andreproductiveageofLesserBlack-backedGullsLarusf.fuscusincaptivityandofHerringGullsL.a.argentatusinth-e--Fenn.57:40-41.[InGermanwith-FinnishandEnglishsummaries.]Goodermote,D.Superior.1980.HerringGullnestcountsonthenorthshoreofLake Loon52:15-17.Grant,P.J.1980.FieldidentificationofwestPalearcticgulls.Part3.Audouin's,Herring,LesserBlack-backed,GreatBlack-backedandGreatBlack-headedGulls.Brit.Birds73:113-158.Graves,J.A. and A.Whiten.1980.AdoptionofstrangechicksbyHerringGull,LarusargentatusL.Z.Tierpsychol.54:267-278.[InEnglishwithGermansummary.]Hamas,M.J.1980.HerringGullsnestingonBeaverIsland.Jack-PineWarbler58:93.Hosey,G.R. andF.Goodridge.1980.Establishmentofterritoriesintwospeciesofgullon WalneyIsland,Cumbria.BirdStudy27:73-80.Ingold,J.L.1980.PlayinHerringGulls.Inl.BirdBanding52:38.236

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HerringGullKilpi,M.,H.Puntii,andT.Toivonen.1980.NumbersofgullsnestingonthenortherncoastoftheGulfofFinland.OrnisFenn.57:153-160.[InEnglishwithFinnishsummary.]King,B.1980.Foot-paddlingbypairsofHerringGulls.Brit.Birds73:312.Kuschert,H.1980.Morphologisch-biometrischeUntersuchungenanSilbermowen(Larusargentatus)einerBinnenlandkoloniesSchlewig-Holstein.[MorphologicalandbiometricalresearchonHerringGulls(Larusargentatus)fromaninlandcolonyinSchleswig-Holstein.]Angew.Ornithol.4:190-194.[InGermanwithEnglishsummary.]Kuschert,H.and G. Vauk.1980.DanischeOstsee-Silbermoew(Larusargentatus)alsBrutvogelimBinnenlandSchleswig-Holsteins(PlonerSee).[DanishHerringGullLarusargentatusasabreedingbirdininlandSchleswig-Holstein(Ploner-See).]Vogelwarte30:147.Leach,M.C. M.,S.V.KearseyandC.Wells.1980.The1978-79surveyofHerringGull(Larusargentatus)coloniesontheYorkshireandClevelandCoast.Naturalist(Leeds)105:107-114.Lustick,S.,M.AdamandA.Hinko.1980.andtheradiativeheatloadinbirds.Interactionbetweenposture,color,Science208:1052-1053.Monaghan,P.1980.Dominance anddispersalbetweenfeedingsitesintheHerringGull(Larusargentatus).Anim.Behav.28:521-527.Morris,R.D.andJ.E.Black.1980.RadiotelemetryandHerringGullforagingpatterns.J.FieldOrnithol.51:110-118.Myrberget,S.andO.Olsvik.1980.GramakeungespistelarveravFrostmaIer.[HerringGulleatswintermothlarvae.]Fauna(Oslo)33:41.[InNorwegianwithEnglishsummary.]Norstrom,R.J.,D.J.Hallett,F.I.OnuskaandM.E.Comba.1980.Mirex,anditsdegradationproductsinGreatLakesHerringGulls.Environ.Sci.Technol.14:860-866.Patten,S. M.,Jr.1980.InterbreedingandevolutionintheLarusglaucescens-Larusargentatuscomplexonthesouthcoastof thesis,JohnsHopkinsUniv./Baltimore,MD.250pp.Plath,L.1980.BrutenderSilbermowe(Larusargentatus)aufeinemHallendachinRostock.[Breedingof Larus-argentatus inRostock.]Beitr.Vogelkd.26:358.[InGerman.]--237

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HerringGullPlotz,J.1980.ParasitosendurchNematodenbefallbeiSilbermowen(Larusargentatus)und Mantelmowen(Larusmarinus)vonderInselHelgoland.[ParasitosisthroughnematodeinfectioninHerringGulls(Larusargentatus)andGreaterBlack-backedGulls(Larusmarinus)fromHelgoland.]Angew.Ornithol.4:195-200.[InGermanwithEnglishsummary.]Pulliainen,E.andA.Marjakangas.1980.EggshellthicknessinelevenseaandshorebirdspeciesoftheBothnianBay.Ornis.Fenn.57:65-70.Richards,C. E.1980.HerringGullscatchingGreaterPipe-fishes.Brit.Birds73:476.Robertson,I.S.1980.ImmatureHerringGullspickingupicesliverS.Brit.Birds73:35.Ryttman,H.,H.Tegelstrom,andH.Jansson.1980.ImmunoelectrophoreticalcomparisonofserafromHerringGull(Larusargentatus)andLesserBlackbackedGull(Larusfuscus)(Aves).Hereditas92:113-116.Shugart,G. W. 1980.FrequencyanddistributionofpolygynyinGreatLakesHerringGullsin1978.Condor82:426-429.Sommani,E.1980.RipetutenidificazionideGabbianoReale(Larusargentatus)nellacittadiRoma.Riv.Ital.Ornitol.50: Italian.]Southern, W. E.1980a.ComparativedistributionandorientationAmericangulls.Pp.449-498inJ.Burger,B.C.OllaandH.(eds.).BehaviorofMarineAnimals.Vol.4:Marine NY.xviiand 515pp.ofNorthE.Winn PlenumPress1980b.Theimpactoffoxpredationongullbreedingsuccess,SleepingBearDunesNationalLakeshore.Pp.111-129inProc.2ndConf.Sci.ResearchNat.Parks,Vol.12:TerrestrialBiology:Zoology.Natl.ParkServ.,Washington,D.C.Southern,W.E.,S.R.Pattonand L.A.Hanners.1980.DifferentialresponseofRing-billedGullsandHerringGullstofoxpredation.Proc.1979Conf.ColonialWaterbirdGroup3:119-127.Sutherland,A.1980.ErythristicHerringGulleggsinCaithness.Scott.Birds11:84.Vauk,G.,E.Vauk-HentzeltandM.Stede.1980.Nachweise vonGeflugltuberkulose(Tuberkulosisavium)beifreilebendenSilbermowen(Larusargentatus).[Tworecordsoffowl-tuberculosis(Tuberkulosisavium)infreelivingHerringGulls(Larusargentatus).]Angew.Ornithol.4:185-189.[InGermanwithEnglishsummary.]Ward,L.D.andJ.Burger.1980.SurvivalofHerringGullanddomesticchickenembryosaftersimulatedflooding.Condor82:142-148.238

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HerringGullWestin,P.1980.Gratrutfangarko1trastiflykten.[HerringGullcatchingmigratingBlackbirdintheair.]VarFagelvarld39:44.[InSwedishwithEnglishsummary.]Whilde,A.1980.The numberofgullsbreedinginthewestofIreland.IrishNat.J.19:328.1979Bourne,W.R.P.1979.ProlongedparentalcareinHerringGullsnestingintownineasternScotland.BirdStudy26:196-197.Burger,J.Gulls.1979a.Competitionandpredation:HerringGullsversusLaughingCondor81:269-277.1979b.Colonysize:atestforbreedingsynchronyinHerringGull-----(Larusargentatus)colonies.Auk96:694-703.Burger,J.andJ.K.Shisler.1979.TheimmediateeffectsofditchingasaltmarshonnestingHerringGullsLarusargentatus.BioI.Conserv.15:85-103.Butler,R.G. andP.Lukasiewicz.1979.AfieldstudyoftheeffectofcrudeoilonHerringGullchickgrowth.Auk96:809-812.Carlsson,L.1979.Trutarsomnaringsparasitpahagar.[Gullskleptoparasitizingheron,Ardeacinerea.]VarFagelvarld38:50.[InSwedishwithEnglishsummary.-]----Duhautois,L.1979.MiseaupointsurlesincursionsdesGoelandsargentesLarusargentatussp.PL.enIle-de-France.Passer16:29-41.E10we,K.D.and S.Payne.1979.Aging youngHerringGullsfrommeasurementsofbodyparts.Bird-Banding50:49-55.Evans, R. M.1979.ResponsivenessofyoungHerringGullstostimulifromtheirownandotherspecies:effectsoftrainingwithfood.Can.J.Zool.57:1452-1457.Foxall, R. A.1979.PresumedhybridsoftheHerringGullandtheGreatBlack-backedGull.Am.Birds33:838.Gilman,A.P.,D. B.Peakall,D.J.Hallett,G. A. Foxand R. J.Norstrom.1979.HerringGulls(Larusargentatus)asmonitorsofcontaminationintheGreatLakes.Pp.280290inAnimalsasMonitorsofEnvironmentalPollutants.Proc.Symp.Environ.Pollut.,Natl.Acad.Sci.Henley,C.A.1979.FactorsinfluencingegglossandchickmortalityinLarusargentatusand fuscus.Ph.D.thesis,Dniv.Oxford/England.239

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HerringGullKallander,H.1979.Labbenpatrutenochtrutenpalabbenellerhundenpakattenochkattenpahundendel2.[SecondarykleptoparasitismbygullsonArcticSkuas.]Anser18:299.[InSwedishwithEnglishsummary.]Karmborg, T.1979.Trutardodarkoltrast.[HerringGull,Larusargentatus,andGreatBlack-backedGull,L.marinus,killingBlackbird.]VarFagelvarld38:50.[InSwedish with Englishsummary.]Kempf,C.,J.Lavergne,B.SittlerandF.Steimer.1979.TentativedenidificationduGoelandargenteLarusargentatussurIeRhinfrancais.Alauda47:37-38.Lebeurier,E.cation?1979.LeGoelandargenteest-ilfideleasonlieudenidifiOiseauRev.Fr.Ornithol.49:233-234.Lustick,S.,B.BattersbyandM.Kelty.1979.ileHerringGullenergeticsandbehavior.EffectsofisolationonjuvenEcology60:673-678.Mille,J.L.1979.AproposducomportementpredateurduGoelandargente.[RegardingthepredatorybehavioroftheHerringGull.]Alauda47:40.Miller,D.S.,D. B. PeakallandW.B.Kinter.1979.Grudeoilingestionimpairsosmo-regulatoryandnutrienttransportinHerringGulls.Fed.Am.Soc.Exp.Biol.6lstAnnu.Meet.,Chicago,1977.Fed.Proc.36:1008.Monaghan,P.1979.AspectsofthebreedingbiologyofHerringGullsLarusargentatusinurbancolonies.Ibis121:475-481.Monaghan,p.andN.Duncan.1979.HowoldisaHerringGull?Notnecessarilyasoldasitlooks.Brit.Birds72:100-103.Nunnally,S.,D.Nunnally,R. Needham andR.Lennon.1979.Nocturnalfeedingofgullsatalightedpier.Chat43:63.Patton,S.R.1979.ShortandlongtermeffectsofRedFoxnocturnalpredationonthenestingsuccessofcolonialbreedingHerringandRing-billedGulls.M.S.thesis,N.Ill.Univ./Dekalb,1L.viand107pp.Pierotti,R.1979.a mixedcolony.ThebehaviorofHerringandGreatBlack-backedGullsin(Abstractonly.)Pac.SeabirdGroupBull.6:46.Radford,D.J.1979.GulldeathsontheNorthUlstercoast,February1978.IrishNat.J.19:391-395.Sobey,D.G.andJ.B.Kenworthy.1979.TherelationbetweenHerringGullsandthevegetationoftheirbreedingcolonies.J.Ecol.67:469-496.240

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HerringGullLudwig,F.C.1976.Toyballooncausesgullmortality.Jack-PineWarbler54:179.Martelli,C.1976.GabbianirealiLarusargentatuspresentineu'exaeroportodiCentocelle(Roma).Riv.Ital.Ornitol.46:274-275.Moore,F.R.1976.HerringGulls.ThedynamicsofseasonaldistributionofGreatLakesBird-Banding47:141-159.Morris,R.D.andR.A.Hunter.1976.Monitoringincubationattentivenessofground-nestingcolonialseabirds.J.Wildl.Manage.40:354-357.Noseworthy,C.M.andJ.Lien.1976.OntogenyofnestinghabitatrecognitionandpreferenceinneonatalHerringGullchicks,LarusargentatusPontoppidan.Anim.Behav.24:637-651.Parsons,J.1976a.FactorsdeterminingthenumberandsizeofeggslaidbytheHerringGull.Condor 78:481-492.1976b.NestingdensityandbreedingsuccessintheHerringGullLarusargentatus.Ibis118:537-546.Parsons,J.,G.ChabrzykandN.Duncan.1976.post-fledgingsurvivalinHerringGulls.EffectsofhatchingdateonJ.Anim.Ecol.45:667-675.Patten,S. M.,Jr.1976.SympatryandinterbreedingofHerringandGlaucouswingedGullsinsouthernAlaska.(Abstractonly.)Pac.SeabirdGroupBull.3:25-26.Patten,S. M.,Jr.andL.R.Patten.1976.BreedingecologyoftheGulfofAlaskaHerringGullgroup.Pp.271-368inEnvironmentalAssessmentoftheAlaskanContinentalShelf, ofPrincipalInvestigators,Vol.2.NOAA,Environ.Res.Lab.,Boulder,CO.Schmidt-Nielsen,B.1976.IntracellularconcentrationsofthesaltglandoftheHerringGullLarusargentatus.Am.J.Physiol.230:514-521.Sobey,D.1976.TheeffectofHerringGullsonthevegetationoftheIsle'ofMay.Bot.Soc.Edinb.Trans.42:469-485.Spitzer,G.1976.ZurErnahrunggelbfussigerSilbermowen(Larusargentatusmichahellis)inderAdriaund imBinnenland.[Onthenutritionoftheyellow-leggedHerringGull(Larusargentatusmichahellis)intheAdriaticareaandintheinland.]Vogelwarte28:298306.[InGermanwithEnglishsummary.]Tolonen,K.E.1976.BehavioralecologyofLarusargentatusandLarusmarinus:agespecificdifferentialinfeedingefficiency,aprobablefactorintheevolutionofdelayedbreeding.Ph.D.thesis,YaleUniv./NewHaven,CT. 275pp.Williams,B.M.,D.W.RichardsandJ.Lewis.1976.SalmonellainfectionintheHerringGull(Larusargentatus).Vet.Rec.98:51.251

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HerringGullWinkel,W.1976.ExperimentelleFreiland-UntersuchungenzumBruttriebderSilbermowe,(Larusargentatus).UberdenEinflussvonFleckung,Farbe,Grosseform,Gewicht,undZahlderEier.[ExperimentalstudyofthebroodingdriveoftheHerringGull(Larusargentatus)inthewild.Ontheinfluenceofthemarking,colour,size,shape,weight,and numberoftheeggs.]Vogelwarte28:212-229.[InGermanwithEnglishsummary.] 1975 Anon.1975a.HerringGullssurviveafterinadvertentburialinanicestore.SeaSwallow24:63.1975b.HerringGullwingtagging.J.Gloucester.Nat.Soc.26:124.Armistead,H.T.1975.BreedingofGreaterBlack-backedGull,HerringGull,andGadwallatSmithIsland,Maryland.Md.Birdlife31:131-134.Banaja,A.A.,J.L.James andJ.Riley.1975.Anexperimentalinvestigationofadirectlife-cycleinReighardiasternae(Diesing1864),apentastomidparasiteoftheHerringGull(Larusargentatus).Parasitology71:493-503.Barth,E.K.1975a.Moult and taxonomyoftheHerringGullLarusargentatusandtheLesserBlack-backedGull fuscusinnorthwesternEurope.Ibis117:384-387.1975b.TaxonomyofLarusargentatusandLarusfuscusinnorthwestern-----Europe.OrnisScand.6:49-63.-----------Bock,F.1975.AngabenzurBrutbiologiederMittelmeersilbermowe(Larusargentatusmichahellis).Egretta18:65-66.Chamberlin,M.L.1975.northernLakeHuron.ivand75pp.BreedingecologyofHerringGullsonanislandinM.S.thesis,CentralMich.Univ.fMt.Pleasant,MI.Cuthbert,F.J.andW.E.Southern.1975.A methodformarkingyounggullsfor indiyidual recognition.Bird-Banding46:252-253.Davis,J.W.F.1975a.SpecializationinfeedinglocationbyHerringGulls.J.Anim.Ecol.44:795-804.1975b.Age,egg-sizeandbreedingsuccessintheHerringGullLarusargentatus.Ibis117:460-473.Evans, R. M.calls.1975.ResponsivenessofyoungHerringGullstoadult"mew"Auk92:140-143.252

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HerringGullFox,G.A.,A.P.Gilman,D.J.Hallett,R.J.Norstrom,F.I.Onuska andD.B.Peakall.1975.HerringGullproductivityandtoxicchemicalsintheGreatLakesin1975.ToxicChemicalsDiv.,Can.Wildl.ServoManuscriptRep.34.35pp.Franchimont,J.1975.Unimportantdortoirdemouettesrieuses(Larusridibundus)etdegoelandsargentes(Larusargentatus)surlaMeuse, alafrorrtierebelgo-nederlandaise.Aves12:286-287.Galusha,J.G.1975.SocialbehaviourofLarusargentatusandLarusfuscus.Ph.D.thesis,Univ.Oxford/Oxford,England.135pp.Groves,S.andG.Peabody.1975.DeadHerringGullisfoundinsidedeadGoosefish.Bird-Banding46:76.Harrington,B.A.1975.PelagicgullsinwinteroffsouthernCalifornia.Condor77:346-350.Harris,J.T.andS.W.Matteson.1975.Gullsandternsasindicatorsofman'simpactuponLakeSuperior.Univ.Wise.SeaGrantProgramTech.Rep.227.45pp.Hartwig,E.andM.Sohl.1975.ZurNahrungderSilbermowe(Larusargentatus)aufderNorseeinselSylt.I.ZusammensetzungderNahrung.[OnthefeedingecologyoftheHerringGull(Larusargentatus)ontheislandofSylt(NorthSea).I.Compositionoffood.]Zool.Anz.194:350-360.[InGermanwithEnglishsummary.]Haycock,K.A. and W.Threlfall.1975.ThebreedingbiologyoftheHerringGullinNewfoundland.Auk92:678-697.Helle,E.1975.Sammakotharmaalokinravintona.[FrogsasfoodofHerringGulls.]OrnisFenn.52:32-33.[InFinnishwithEnglishsummary.]Hesselwood,J.1975.Areportonthefishinglineenquiry.Naturalist(Leeds)929:70.Hoogerwerf,A.1975.denopVlieland.ZilvermeeuwenendegerineaanwasbijEidersenBergeenPiper13:117-124.[InDutchwithEnglishsummary.]Kadlec,J.A.1975.Recoveryratesandlossofaluminum,titanium,andincoloybandsonHerringGulls.Bird-Banding46:230-235.Kallander,H.1975.[PiracybyHerringGullsLarusargentatuson Whooper Swans cygnus.]Anser14:29-38.[InSwedishwithEnglishsummary.]King,B.1975.Gullbehaviourduringsea-mistintheBristolChannel.BristolOrnithol.8:111-112.253

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HerringGullKury,C.R.andM.Gochfe1d.1975.Humaninterferenceandgullpredationincormorantcolonies.Bio1.Conserv.8:23-34.Monaghan,P.1975.Therooftopgull.Wildlife(Lond.)17:170-172.Moore,F.R.1975.InfluenceofsolarandgeomagneticstimulionthemigratoryorientationofHerringGullchicks.Auk92:655-664.O'Meara,M.1975.Building-nestingHerringGullsinCountyWaterford.IrishNat.J.18:152-153.Owen,B.A.1975a.InlandbreedingbygullsinGloucester1974.J.GloucesterNat.Soc.26:35-36.1975b.InlandbreedingbygullsinGloucester1975.J.Gloucester ------Nat. Soc.26:107-108.1975c.InlandbreedingbygullsinGloucester.J.GloucesterNat. ------Soc. 26:249-250.Parnell,J.F.andR.F.Soots.nestinginNorthCarolina.1975b.HerringandGreatBlack-backedGullsAuk92:154-157.Parsons,J.1975a.SeasonalvariationinthebreedingsuccessoftheHerringGull:anexperimentalapproachtopre-fledgingsuccess.J.Anim.Eco1.44:553-573.1975b.AsynchronoushatchingandchickmortalityintheHerringGullLarusargentatus.Ibis117:517-520.Scharf,W.C.andG.Shugart.1975.Nestlingbanding:anindexofreproductiyesuccessinaHerringGullcolony.In1.BirdBandingNews47:125-129.Schwarz,D.andH.W.Nehls.1975.Si1bermowen(LarusargentatusL.)ohneSa1zdrusen(Glandulaesupraorbita1es)imFrei1andversuth.Beitr.Voge1kd. 21:305-311.Sobey,D.G.1975.TherelationshipbetweenHerringGullsandthevegetationoftheirnestingandroostingsites.Ph.D.thesis,Dniv.Aberdeen/Aberdeen,Scotland.Spaans,M.J.andA.L.Spaans.1975.Enke1egegevensoverdebroedbio1ogievandeZi1vermeeuwLarusargentatusopTersche11ing.[SomedataonthebreedingbiologyoftheHerringGullLarusargentatusontheDutchFrisianIslandofTersche11ing.]Limosa48:1-39.[InDutchwithEnglishsummary. ] 254

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HerringGullTuaev,D.G.,V.V.Vasil'evandF.A.Abushev.1975.[OnelucidationofthecausesofdeathofLarusargentatusintheCaspianSea.]Pp.237239inA.1.Cherepa.nov(ed.).[Transcontinentalconnectionsofmigratorybirdsandtheirroleinthedistributionofarboviruses.]Akad. Nauk SSSR.SibirskoeOtdelenie,biologicheskiiInstitut.Izdatel'stvo'Nauka',Novosibirsk.[InRussianwithEnglishsummary.] VandenSteen,J.1975.Deproblematiekantrentsubspecifiekedeterrninatievangeelpoot-zilvermeeuwen.Wielewaal41:359-364.1974Barth,E.K.1974a.SvartbakdruknetGramake.drownsaHerringGull.]Fauna27:95-96.summary. ][GreatBlack-backedGull[InNorwegianwithEnglish1974b.RaseravSildemakeog Gramake iNorge.[Subspeciesofthe -----Lesser Black-backedGullandtheHerringGullinNorway.]Fauna27:236237.[InNorwegianwithEnglishsummary.]Baudinette,R.V.andK.Schmidt-Nielsen.1974.EnergycostofglidingflightinHerringGulls.Nature(Lond.)248:83-84.Britton,P.L.1974.Broad-billedSandpipersandHerringGullswinteringontheNorthKenyacoast.Bull.EastAfr.Nat.Hist.Soc.1974:112-113.Citron,J.D.1974.GullsstealfishfromCommonMergansers.DelmarvaOrnithol.9:71-72.Drury,W.H.andJ.A.Kadlec.1974.ThecurrentstatusoftheHerringGullpopulationinthenortheasternUnitedStates.Bird-Banding45:297-306.Eggers,J.1974.Vorkommen undHerkunftderLechmowe(Larusridibundus)im HamburgerRaumimVergleichzurSturm-,Silber-,und Mantelmowe(Laruscanus, argentatus, marinus).Hamburg.AvifaunaBeitr.12:95-144.[InGermanwith summary.]Filonov,K.P.,V.I.LysenkoandV.D.Siokhin.1974.[PeculiaritiesofLimicolaeandLarinestingonislandsofMolochnyEstuary(theSeaofAzov).]Vestn.Zool.1974:52-58.[InRussianwithEnglishsummary.]Fimreite,N.,E.Brun,A.Frostlie,p.FrederichsenandN.Gundesen.1974.MercuryineggsofNorwegianseabirds.Astarte7:71-75.Firth,R.,Jr.1974.Experimentalcross-fosteringofHerringGullandGreatBlack-backedGullchicks.Auk91:139-144.Forsythe,D.M.1974a.Anecologicalstudyofgullpopulationstoreducethebird-aircraftstrikehazardatCharlestonAirForceBase.AirForceWeaponsLab.Tech.Rep. 73. 142pp.255

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HerringGullGauthreaux,S.A.,Jr.[ed.]1974.Proc.Conf.ontheBioI.AspectsoftheBird/AircraftCollisionProblem,5-7February1974. Dep.Zool.,ClemsonUniv./Clemson,SC.535pp.Gilbertson,M.1974a.PollutantsinbreedingHerringGullsinthelowerGreatLakes.Can.Field-Nat.88:273-280.Gilbertson,M.andR.Hale.1974a.CharacteristicsofthebreedingfailureofacolonyofHerringGullsinLakeOntario.Can.Field-Nat.88:356-358. ______ 1974b.Ontario.EarlyembryonicmortalityinaHerringGullcolonyinLake Can.Field-Nat.88:354-356.Gochfeld,M.1974b.Prevalenceofsubcutaneousemphysemainyoungterns,skimmersandgulls.J.Wildl.Dis.10:115-120.1974c.Gullpredationonmigrantpasserinebirds.Englehardtia6:12-13.Grant,D.R.1974.Localgullmovementsasahazardtoaircraft.BirdStudy21:169-179.Grubb,T.C.,Jr.1974.IndividualdistanceintheHerringGull.Auk91:637-639.Hibbert,F.W.1974.CootattackingHerringGull.Brit.Birds67:242.Jonkel,G.M.andO.S.Pettingill,Jr.1974.Retractionofalongevityrecordfora36-year-oldHerringGull.Auk91: 432.Jorgensen,O.H.andI.Kraul.1974.EggshellparametersandresiduesofPCBandDDEineggsfromDanishHerringGulls argentatus.OruisScand.5:173-179.Jouanin,C.1974.NotesurLarusargentatusatlantisauxilesSelvagens.Cyanopica1:19-24.[InFrench.]Kihlman,J.andL.Larsson.1974.OntheimportanceofrefusedumpsasafoodsourceforwinteringHerringGullsLarusargentatusPont.OrnisScand.5:63-70.Melville,D.1974.AnalysisofHerringGullpelletscollectedinCo.Antrim.SeabirdRep. 4:40-46.Michaelsen,J.1974.InnlandshekkingavGramake.[InlandbreedingofHerringGull.]Fauna27:95.[InNorwegianwithEnglishsummary.]Patten,S.,Jr.andA.R.Weisbrod.1974.SympatryandinterbreedingofHerringandGlaucous-wingedGullsinsoutheasternAlaska.Condor76:343-344.256

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HerringGullRee,V.1974.TidligkroppsmytinghosGramake.[EarlybodymoultintheHerringGull.]Fauna27:93-94.[InNorwegianwithEnglishsummary.]Reperant,J.andJ.-P.Raffin.argente(LarusargentatusfuscusgraellsiiBrehm).Nat.278:2335-2338.1974.LesprojectionsvisuellesIegoelandargentatusPontopp.)etIegoelandbrun(LarusC.R.Hebd.SeancesAcad.Sci.Ser.D.Sci.Ryder,J.P.1974.Organochlorineandmercuryresiduesingulls'eggsfromwesternOntario.Can.Field-Nat.88:349-352.Scanes,C.G.,P.Cheeseman,J.G.Phillips,andB.K.Follett.1974.SeasonalandagevariationofcirculatingimmunoreactiveluteinizinghormoneincaptiveHerringGulls,Larusargentatus.J.Zool.(Lond.)174:369-375.Scharf, W. C.andB.Buckingham.1974.CongenitalfootabnormalityinaHerringGull.Inl.BirdBandingNews46:30-32.Sturmhoefel,E.1974.Silvermowe-schmarotzbeiTurmfalken.Falke21:140.Threlfall,W., E.EveleighandJ.E.Maunder.1974.Seabirdmortalityinastorm.Auk91:846-849.Williams,T.C.,J.M.Williams,J.M.TealandJ. W. Kanwisher.1974.HomingflightsofHerringGullsunderlowvisibilityconditions.BirdBanding45: Witt,H.1974.ZurNahrungsokologiederMittelmeersilbermowe(Larusargentatusmichahellis)aneinemBrutplatzaufSardinien.[OnthefeedingecologyofaSardiniancolonyofMediterraneanHerringGulls(Larusargentatusmichahellis).]Vogelwelt95:148-150.[InGermanwithEnglishsummary.] 1973Andrle,R.F.1973.AsecondprobablehybridofLarusmarinusandL.argentatusontheNiagaraRiver.Can.Field-Nat.87:170-171.Angles,R.1973.HerringGull(Larusargentatus)takingadultStormPetrel(Hydrobatespelagicus).Brit.Birds66:495-496.Anon.1973.AnadventurousHerringGull.Brit.TrustOrnithol.News55:9.Bourget,A.A.1973.RelationofeidersandgullsnestinginmixedcoloniesinPenobscotBay,Maine.Auk90:809-820.257

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HerringGullChan,S. W. C.andJ.G.Phillips.1973a.Secretionofl7-deoxycorticosteroidsbyHerringGull(Larusargentatus)adrenals.Gen.CompoEndocrinol.20:274-282.1973b.Variationsinthevitroproductionofcorticosteroidsbythe-----HerringGull(Larusargentatus)adrenalglands.Gen.CompoEndocrinol.20:283-290.Delius,J.D.1973.Agonisticbehaviourofjuvenilegulls,aneuroethologicalstudy.Anim.Behav.21:236-246.Drury, W. H.1973-74.PopulationchangesinNewEnglandseabirds.BirdBanding44:267-313;45:1-15.Evans,R.M.1973.DifferentialresponsivenessofyoungRing-billedGullsandHerringGullstoadultvocalizationsoftheirownandotherspecies.Can.J.Zool.51:759-770.Fitzgerald,G.R.andJ.C.Coulson.1973.ThedistributionandfeedingecologyofgullsonthetidalreachesoftheriversTyneandWear. Vasculum58:29-47.Forsythe,D.M.June1972.1973.GullpopulationsatCharleston,S.C.,June1971toChat37:57-62.Franzmann,N.-E.1973.GulbenetSolvrnage(Larusargentatusomissus)ynglendepaChristianso.Feltornithologen15:209.Godfrey, W. E.marinus.1973.Morepresumedhybridgulls:LarusargentatusX L. Can.Field-Nat.87:171-172.Goethe,F.andE.Schuz.1973.Zum"Typus"vonLarusargentatusheugliniBreeimStaatlichenMuseumfurNaturkundezuStuttgart.Stuttg.Beitr.Naturkd.Ser.A.(BioI.)261:1-8.[InGermanwithEnglishsummary.]Hedgren,S.andL.Larsson.1973.'Labbliknande'trutar.VarFagelvarld32:288.[InSwedishwithEnglishsummary.]Hunt,G.L.,Jr.andM. W. Hunt.1973.HabitatpartitioningbyforaginggullsinMaineandnorthwesternEurope.Auk90:827-839.Isenmann,P.1973a.Dispersionhivernaledugoelandargente(Larusargentatus)surIelittoralatlantiquedel'estuairedelaLoireaupaysBasque.ArVran5:101-108.1973b.BiometrischeUntersuchungenanderGelbfussigenSilber------mowe(Larusargentatusmichahellis)ausderCamargue.[BiometricalstudyofthepopulationoftheYellow-leggedHerringGull(Larusargentatusmichahellis)intheCamargue.]Vogelwarte27:16-24.[InGermanwithEnglishsummary.] 258

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HerringGullIsenmann,P.1973c.DonneessurlesdeplacementserratiquesdeGoelandsargentesapiedsjaunes(Larusargentatusmichahellis)nesenMediterranee.OiseauRev.Fr.Ornithol.43:187-195.[InFrenchwithEnglishsummary.]1973d.Distributionhivernaleen1972-1973duGoelandargenteapiedsjaunes(Larusargentatusmichahellis)surIelittoralatlantiqueduMorbihanauPaysBasque.OiseauRev.Fr.Ornithol.43:260-262.Jorgensen,O.H.1973.SomeresultsofHerringGullringinginDenmark19581969.Dan.Ornithol.Foren.Tidsskr.67:53-63.Macdonald,D.1973.HerringGullcolonyinpinewood.Brit.Birds66:228.Macdonald,S.M.andC.F.Mason.1973.Predationofmigrantbirdsbygulls.Brit.Birds66:361-363.MacRoberts,M.H.1973.ExtramaritalcourtinginLesserBlack-backedandHerringGulls.Z.Tierpsychol.32:62-74.McLannahan,H.M.C.1973.SomeaspectsoftheontogenyofcliffnestingbehaviourintheKittiwake(Rissatridactyla)andtheHerringGull(Larusargentatus).Behaviour44:36-88.Moore,F.R.1973.HerringGullpostbreedingdispersalandrelatedchickorientationbehavior.M.S.thesis,N.IllinoisUniv./Dekalb,IL.80pp.Noseworthy,C.,S.StokerandJ.Lien.1973.HabitatpreferencesinHerringGullchicks.Auk90:193-194.Nystrom,M.1973.Extinction,disinhibitionandspontaneousrecoveryofthepeckingresponseinyoungHerringGulls.Behaviour45:271-281.Podmore,A.1973.HerringGullpiracy.BirdLife1973:36.Sanger,G.A.1973.PelagicrecordsofGlaucous-wingedandHerringGullsinthenorthPacificOcean.Auk90:384-393.VanZurk,H.1973.michahellis).Legoelandargentemediterraneen(LarusargentatusRivieraSci.8:55-56.Vermeer,K.1973b.FoodhabitsandbreedingrangeofHerringGullsintheCanadaprairieprovinces.Condor75:478-480.1973c.Comparisonofegg-layingchronologyofHerringandRing-billed-----GullsatKawinawLake,Manitoba.Can.Field-Nat.87:306-308.Williams,T.C.andJ.M.Neal.1973.Theflightofblindfoldedbirds.Bird-Banding44:102-109.259

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HerringGull1972Anderson,R.B.something?1972.Gulls,gullsand moregulls-aretheytellingusMaineFishGame14:8-10.Andrle,R.F. 1972.AnotherprobablehybridofLarusmarinusand L.argentatus.Auk89:669-671.Blus,L.J.,C.D.Gish,A.A.BelisleandR.M.Prouty.relationshipofDDEresiduestoeggshellthinning.376-377.1972.LogarithmicNature(Lond.)235:Brackbill,H.1972.Fowlthatdon'tbefoul.Md.Conserv.48:4-7.Cooke, F.andR.K.Ross.1972.DiurnalandseasonalactivitiesofapostbreedingpopulationofgullsinsoutheasternOntario.WilsonBull.84:164-172.Dexter,R.W.chusetts.1972.RecoveriesofHerringGullsbandedatCape Ann,Massa(Abstractonly.)Proc.Internatl.Ornithol.Congr.15:639.Dittberner,H. 1972. DieGrossmowen(Larusmarinus, 1. fuscusand 1. argentatus)alsGastvogelmarkischerGewasser.Beitr.Vogelkd.18:141-155.Drury,W.H.andI.C.T.Nisbet.1972. ThebiologyofHerringGullsinNewEngland.EcologyofMigratoryBirds:A Symposium.Res.Rep.2.278pp.importanceofmovementsinthePp.173-212inPopulationU.S.FishWildl.ServoWildl.Ensor,D.M.andJ.G.Phillips.1972a.Theeffectofageandenvironmentonextrarenalsaltsecretioninjuvenilegulls(LarusargentatusandL.fuscus).J.Zool.(Lond.)168:119-126.1972b. Theeffectofdehydrationonsaltandwaterbalanceingulls-----(LarusargentatusandL.fuscus).J.Zool.(Lond.)168:127-137.Forsythe,D.M.Carolina.1972.Ring-billedandHerringGullrecoveriesfromSouthBird-Banding43:276-281.Frankum,R.1972.HerringGullsinUganda.Bull.EastAfr.Nat.Hist.Soc.1972:81-82.Haarmann,K.1972.Das BrutvorkommenderSilbermowe(Larusargentatus) an derNiederelbe.Vogelkd.Ber.Nieders.4:79-81.Hunt,G.L.,Jr.1972a.Influenceoffooddistributionand humandisturbanceonthereproductivesuccessofHerringGulls.Ecology53:1051-1061.1972b.HerringGullpopulationdynamics:thesignificanceofman'swasteproducts.(Abstractonly.)Proc.Internatl.Ornithol.Congr.15:652.260

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HerringGullKarlsson,K.-A.,B.E.SamuelssonandG.O.Stern.1972.Identificationofaxylose-containingcerebrosideinthesaltglandoftheHerringGull.J.LipidRes.13:169-175.Kumari,E.1972.DistributionchangesandlivingconditionsoftheHerringGull(Larusargentatus)intheEastBalticarea.(Abstractonly.)Proc.Internatl.Ornithol.Congr.15:663.Lyster,I.H.J.1972.MolekillsHerringGull.Scott.Birds7:207-208.MacRoberts,B.R.andM.H.MacRoberts.1972a.TherelationshipbetweenlayingdateandincubationinHerringandLesserBlack-backedGulls.Ibis114:93-97.1972b.SocialstimulationofreproductioninHerringandLesser-----Black-backedGulls.Ibis114:495-506.Mill,P.J.1972.AnultrastructuralstudyofthebodywallmusclesofthepentastomidReighardiasternaeDies.1864,parasiticoftheHerringGullLarusargentatusPontopp.Z.Zellforsch.Mikrosk.Anat.130:12-17.Mollering,K.1972.QuantitativeUntersuchungenzurBrutbiologiesderSilbermowe(Larusargentatus)aufderVogelinselMellum. Abh.Landesmus.Naturkd.Munster34:79-87.[InGerman.]Nisbet,I.C.T.andW.H.Drury.1972.Post-fledgingsurvivalinHerringGullsinrelationtobrood-sizeanddateofhatching.Bird-Banding43:161-172.Nystrom,M.1972.Onthequantificationofpeckingresponsesinyounggulls(Larusargentatus).Z.Tierpsychol.30:36-44.Parsons,J.Gull.1972.Eggsize,layingdateandincubationperiodintheHerringIbis114:536-541.Seymour,N.R.1972.Successofthreegullspeciesfeedingon swarmingantsinAntigonishCounty,NovaScotia.Can.Field-Nat.86:391-392.Spaans,A.L.1972.Humaninfluenceonthefood-supplyoftheHerringGull(Larusargentatus)inHolland.(Abstractonly.)Proc.Internatl.Ornithol.Congr.15:688.Stenman,0.,R.KomuandA.Ermala.1972.Kloraloosiharmaa-jamerilokkimyrkkyna.[FinnishexperienceofpoisoningHerringGullsandGreatBlack-backedGullswithchloralose.]SuomenRiista24:107-116.[InFinnishwithEnglishsummary.] Thomas,G.J.reserves.1972.AreviewofgulldamageandmanagementmethodsatnatureBioI.Conserv.4:117-127.261

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HerringGullTuaev,D.G.,V.I.Vasilev,andF.A.Abushev.1972.[OntheoccurrenceofmassdestructionoftheHerringGullinanestingcolonyonGlinyanIslandintheAzerbaidjanSSR.]Ornitologiya10:394-395.[InRussian.]Voipio,P.1972.SilbermowenderLarusargentatuscachinnans--GruppealsBeseidlerdesbaItischenRaumes.[LarusargentatuscachinnansasacolonizeroftheBalticarea.]Ann.Zool.Fenn.9:131136.[InGerman.]Winkler,R.1972.LapneumatisationducranechezLarusargentatusmichahellis.Alauda40:272-277.[InFrenchwithEnglishsummary.]1971Bednorz,J.1971.Mewapospolita(Laruscanus),mewasrebrzysta(Larusargentatus)irybitwawielkodzioba(Hydroprognecaspia)gniezdzasienapolskimwybrzezu.[CommonGull(Laruscanus),HerringGull(Larusargentatus),andCaspianTern (Hydroprogne-caspia) nestingonPolishsea----coast.]NotatkiOrnitol.12:67-71.[InPolishwithEnglishsummary.]Bjerk,J. E. andG.Holt.1971.ResiduesofDDEandPCBineggsfromHerringGull(Larusargentatus)andCommonGull(Larus inNorway.ActaVet.Scand.12:429441. M.1971.UberdenEinflussvonPhenobarbitalaufdasBrutuerhaltenderSilbermowe(LarusargentatusargentatusPontopp.).[EffectsofphenobarbitolonbroodcarebehaviouroftheHerringGull(LarusargentatusargentatusPontopp.).]Z.Tierpsychol.28:487-493.[InGermanwithEnglishsummary.]Brackbill,H.1971.HerringandRing-billedGullspairedorcourtinginMarylandinJanuaryandFebruary.Auk88:438-440.Cuss,J.and H.M.Blair.1971.ErythrismineggsofHerringGullinScotland.Scott.Birds6:282.Delius,J.D.1971a.ForagingbehaviourpatternsofHerringGullselicitedbyelectricalforebrainstimulation.Experientia27:1287-1289.1971b.Neuralsubstratesofvocalizationsingullsandpigeons.-----Exp.Brain.Res.12:64-80.Drury,W.H.andI.C.T.Nisbet.1971.TheimportanceofmovementsinthebiologyofHerringGullsinNewEngland.Symp.PopulationEcologyofMigratoryBirds,PatuxentWildl.Res.Cent.,October1969.Erwin,R.M.1971.Thebreedingsuccessoftwosympatricgulls,theHerringGullandtheGreatBlack-backedGull.WilsonBull.83:152-158.Goldman,P.1971.HerringGullpredationoncommonwatersnakeinLakeErie.WilsonBull.83:196-197.262

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Ivanovs,M.1971.Lake,1970.HerringGullHafft,J.H.1971.HerringGullattacksEaredGrebe.Condor73:253.Harris,M.P.1971.EcologicaladaptationsofmoultinsomeBritishgulls.BirdStudy18:113-118.NestingofRing-billedandHerringGullsatMilleLacs Loon43:58-59.Jehl,J.R.1971.AhybridGlaucousXHerringGullfromSanDiego.Calif.Birds2:27-32.Kadlec,J.A.1971.Gullcolonies.EffectsofintroducingfoxesandraccoonsonHerringJ.Wildl.Manage.35:625-636.Kask, E.1971.[Asea-gull(Larusargentatusomissus)nestinatreetop.]EestiLoodus14:352.[InEstonian.]Loos-Frank,B.1971.ZurTrematodenfaunadelSilbermowe(Larusargentatus)andersudlichenNordsee.Vogelwarte26:202-212.[InGermanwithEnglishsummary.]Nisbet,I.C.T.andW.H.Drury.1971.Strategyofmanagementofanaturalpopulation:theHerringGullinNewEngland.Pp.441-454inWorldConf.BirdHazardstoAircraft,Sept.1969.Natl.Res.Counc.,Kingston,Ontario.542pp.Parsons,J.1971a.CannibalisminHerringGulls.Brit.Birds64:528-537.1971b.ThebreedingbiologyoftheHerringGull,Larusargentatus.-----Ph.D.thesis,Univ.Durham/Durham,England.Pearson,J.C. and G.Prevot.1971.Galactosomumtimondavidisp.n.(Trematoda:Heterophyidae)fromLarusargentatus,withanoteonthemetacercaria.J.Parasitol.57:1227-1230.Prevot,G.1971a.Contributional'etudedesMicrophallidaeTravassos,1920(Trematoda).CycleevolutifdeMicrophalluspachygrapsiDeblock&Prevot,1969.ParasiteduGoelandapiedsjaunes(Larusargentatus).Ann.Parasitol.Hum.Compo46:453-461.[InFrenchwithEnglishsummary.]1971b.Cycleevolutifd'AporchismassiliensisTimon-David,1955-----(DigeneaEchonostomatidae),parasitedugoelandLarusargentatusmichahellisNaumann.Bull.Soc.Zool.Fr.96:197-208.[InFrenchwithEnglishsummary.]Spaans,A.L.1971.OnthefeedingecologyoftheHerringGullLarusargentatusPont.inthenorthernpartoftheNetherlands.Ardea59:73-188.Thompson,G.1971.Thephotopicspectralsensitivityofgullsmeasuredbyelectroretinographicandpupillometricmethods.VisionRes.11:719-731.263

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HerringGullTinbergen,N.tracked.1971.Clevergullsanddumbethologists--or:thetrackersVogelwarte26:232-238.Vermeer,K.1971a.SomebreedingaspectsofHerringGullsatKawinawLake,Manitoba.BlueJay29:207-208.1971b.AsurveyofmercuryresiduesinaquaticbirdseggsintheCan-----adianprairieprovinces.Trans.N.Am.Wildl.Nat.Res.Conf.36:138-152.Winkel,W.1971.UberdenBruttreibderSilbermowe(Larusargentatus)inBeziehungzurFarbederEier.[OnthebreedingdriveoftheHerringGull(Larusargentatus)inrelationtothecoloroftheeggs.]Vogelwarte26:249-254.[InGermanwithEnglishsummary.]1970Anderson,D.W.1970.Chlorinatedhydrocarbons:theirdynamicsandeggshelleffectsonHerringGullsandotherspecies.Ph.D.thesis,Univ.Wisconsin/Madison,WI.168pp.Andersson,A.1970.Foodhabitsandpredationofaninland-breedingpopulationoftheHerringGullLarusargentatusinsouthernSweden.OrnisScand.1:75-81.Baerends,G.P.,R.H.Drent,P.GlasandH.Groenewold.1970.AnethologicalanalysisofincubationbehaviourintheHerringGull.Pp.135-235inG.P.BaerendsandR. H.Drent(eds.).TheHerringGullanditsegg.BehaviourSuppl.17.ixand 312pp.Corkhill,P.1970.Effectofegg-puncturingonlimitingchickproductioninHerringGulls.BirdStudy17:340-342.Davis,J.W.F.1970.EggssizesandhatchingweightsintheHerringandLesserBlack-backedGulls.Rep. SkokholmBirdObs.1970:24-31.Delius,J.D. andG.Thompson.1970.BrightnessdependenceofcolourpreferencesinHerringGullchicks.Z.Tierpsychol.27:842-849.Drenckhahn,D.1970.UntersuchungenanregioolfactoriaundnervusolfactoriusderSilbermowe(Larusargentatus).Z.Zellforsch.Mikorsk.Anat.106:119-124.Drent,R.H.1970.FunctionalaspectsofincubationintheHerringGull.Pp.1-132inG.P.Baerendsand R.H.Drent(eds.).TheHerringGullanditsegg.BehaviourSuppl.17.ixand312pp.Drent,R.H.,K.PostumaandT.Joustra.1970.TheeffectofeggtemperatureonincubationbehaviourintheHerringGull.Pp.235-261inG.P.BaerendsandR. H.Drent(eds.).TheHerringGullandits BehaviourSuppl.17.ixand 312pp.264

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HerringGullGobeil,R.E.1970.ArterialsystemoftheHerringGull(Larusargentatus).J.Zool.(Lond.)160:337-354.Hakansson,C.H. and B.Malcus.1970.SecretivepressureexertedbythestimulatednasalsaltglandinLarusargentatus(HerringGull).ActaPhysiol.Scand.78:249-254.Harris,M.P.1970a.RatesandcausesofincreasesofsomeBritishgullpopulations.BirdStudy17:325-335.1970b.AbnormalmigrationandhybridizationofLarusargentatusand-----L.fuscusafterinterspecicfosteringexperiments.Ibis112:488-498.Hatch,J.J.1970.PredationandpiracybygullsataterneryinMaine.Auk87:244-254.Holley,A.J.F.1970.SomeaspectsofrearingtheyounginLarusgullsatStertIsland--aninterimreport.BristolOrnithol.3:100-117.Ingolfsson,A.1970b.HybridizationofGlaucousGullsLarushyperboreusandHerringGulls argentatusinIceland.Ibis112: Komnick,H.and E.Kniprath.1970.MorphometrischeUntersuchungenanderSalzdrusevonSilbermowen.Cytobiologie1:228-247.[InGermanwithEnglishsummary.]Kumerloeve,H.1970.WeitereNachweisederSilbermowe,LarusargentatusalsKleinasiatischerBinnenland-Brutvogel.Vogelwarte25:267268.Lina,P.H. C.1970.ObservationsontheinfectionofyoungHerringandLesserBlack-backedGullsbyectoparasites.Ardea58:258-261.Lippens,L.1970.LapresenceauZwin, a Knokke(littoralBeIge),d'unhybrideprobabledeGoelandbourgmeste(Larushyperboreus)etduGoelandargente(Larusargentatus).Aves7:105-106.Lohmer,K.and G. Vauk.1970.EinweitererBeitragzurErnahrungHelgolanderSilbermowen(Larusargentatus).Vogelwarte25:242-245.[InGermanwithEnglish Mikkola,H.1970.IdentificationofalbinoHerringGullsLarusargentatus.OrnisFenn.47:172-176.Parsons,J.1970.Relationshipbetweeneggsizeandpost-hatchingchickmortalityintheHerringGull(Larusargentatus).Nature(Lond.)228:1221-1222.265

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HerringGullRyden,O.1970.Onthebehaviourinanalarmprovoking,constantstimulussituationofa"naturalexperiment".ObservationsonapairofHerringGulls(Larusargentatus)andCommonTerns(Sternahirundo)breedingonthesameislet.J.Ornithol.111:48-53.------Southern,W.E.1970a.EnroutebehaviorofhomingHerringGullsasdeterminedbyradiotracking.WilsonBull.82:189-200.1970b.-----andwind.Orientationingulls:effectofdistance,directionofrelease,LivingBird9:75-95.Stromar,L.1970.Prilogpoznavanjubiologijegalebovaklaukavaca,LarusargentatusPontopp.,icigraobicnih,SternahirundoL.,tenjihovihmedusobnihodnosananekimKornatskim otoctma. [Thebiologyof,theHerringGull,LarusargentatusPontopp.,andCommonTern,SternahirundoL.andinterrelationshipson someKornatIslands.]Larus21-22:87-93.[EnglishtranslationfromSerbo-Croatian.]Stunkard,H.W.1970.SpeciesofRenicola(Trematoda)inthekidneysofthegull,Larusargentatus.WoodsHoleMar.BioI.Lab.Bull.139:438-439.Thompson,G.1970.Colourvisioninbirds,withspecialreferencetoHerringandLesserBlack-backedGull(LarusargentatusandLarusfuscus).Ph.D.thesis,OxfordUniv./Oxford,England.-----------Vermeer,K.1970.BreedingrecordsofHerringGullsinAlbertaandCaliforniaGullsinManitoba.Can.Field-Nat.84:182.Weaver,D.K.1970.ParentalbehavioroftheHerringGull(Larusargentatussmithsonianus)anditseffectonreproductivesuccess.Ph.D.thesis,Univ.Mich./AnnArbor,MI. 87pp.Weaver,D.K.andJ.A.Kadlec.1970.A methodfortrappingbreedingadultgulls.Bird-Banding41:28-31.1969Able,J.H.1969.ElectronmicroscopicdemonstrationofadenosinetriphosphohydrolaseactivityinHerringGullsaltglands.J.Histochem.Cytochem.17:570-584.Beer,J.V.1969.ObservationsonthedispersalofgullsmarkedonSteepHolmandtheDenny.SteepHolmGullRes.Stn.Rep.1968:4-9.Bongiorno,S.F.andJ.Swinebroad.1969.IncreaseinHerringGullcolonyinCape May,NewJersey.WilsonBull.81:99-100.Chinery,D.J.1969.GreatSkuaentangledwithHerringGull.Brit.Birds62:116-117.266

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HerringGullCleeves,T. R. 1969.HerringGullcatchingandeatingbat.Brit.Birds62:333.Dyck,J.1969.Traekkendedvaegfalk(Falcocolumbarius)druknetafMager.[MigratingMerlin(Falcocolumbarius)drownedbygulls.]Dan.Ornithol.Foren.Tidsskr.63:130-131.[InDanishwithEnglishsummary.]Felton,C.1969.HerringGullenteringwindowforfood.Brit.Birds62:76.Friend,M.andD.O.Trainer.1969.AspergillosisincaptiveHerringGulls.Bull.Wildl.Dis.Assoc.5:271-275.Goertz,M.andF.Goethe.1969. Ringfunde Nordsee-Silbermowen(Larusargentatus).Teil1:BeringungenaufderInselMemmert.Auspicium3:305-317.Guillou,J.J.1969.DifferencesecologiquesentreLarusa.argentatusetLarusfuscusgraellsienhivernagedansdesmileux'naturels'.Alauda37:338-345.Hakansson,C.H.andB.Malcus.1969.SecretiveresponseoftheelectricallystimulatednasalsaltglandsinLarusargentatus(HerringGull).ActaPhysiol.Scand.76:385-392.Harms,W.1969. Eine weitereAnsiedlungderSturmmowe(Laruscanus)undBruteinerSilbermowe(Larusargentatus)inHamburg.Ornithol.Mitt.21:165.Harris,M.P.1969.EffectoflayingdateonchickproductioninOystercatchersandHerringGulls.Brit.Birds62:70-75.Harris,M.P.andP.Hope-Jones.1969.SexualdifferencesinmeasurementsofHerringandLesserBlack-backedGulls.Brit.Birds62:129-133.Ingolfsson,A.1969a.Behaviourofgullsrobbingeiders.BirdStudy16:45-52.1969b.Sexualdimorphismoflargegulls(Larusspp.).Auk86:732--737.Kadlec,J.A.andW.H.Drury,Jr.1969.LossofbandsfromadultHerringGulls.Bird-Banding40:216-221.Kadlec,J.A.,W.H.Drury,Jr.,andD.K.Onion.1969.Growth andmortalityofHerringGullchicks.Bird-Banding40:222-233.King,B.1969.HerringGullsfeedingonalgae.Brit.Birds62:333.Kumerloeve,H.1969.DieSilbermowe,Larusargentatus,alsBinnenlandBrutvogelimostlichenKleinasien.Vogelwarte25:47-49.267

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HerringGullLohmer,K.andG.Vauk.1969.NahrungsokologischeUntersuchungenanubersommerndenSilbermowen(Larusargentatus)aufHelgolandimAugust/September1967.Bonn.Zool.Beitr.20:110-114.[InGerman.]Ludwig,J.P.andC.E.Ludwig.1969.TheeffectsofstarvationoninsecticidecontaminatedHerringGullsremovedfroma LakeMichigancolony.Proc.Conf.GreatLakesRes.1969:53-60.Parsons,J.1969.ChickmortalityinHerringGulls.IbisIll:443-444.Rea,S.C.1969.Plasticdevicecausesgullmortality.WilsonBull.81:105-106.Southern,W.E.1969.Gullorientation.SkyconditionsinrelationtoRing-billedandHerringTrans.Ill.StateAcad.Sci.62:343-349.Vauk,G.andK.Lohmer.1969.EinweitererBeitragzurErnahrungderSilbermowe(Larusargentatus)inderDeutschenBucht.Veroeff.Inst.Meeresforsch.Bremerhaven12:157-160.[InGermanwithEnglish summary.] Wallace,D.I.M. 19n9. HerringGullsatTarkwa,Lagos:aspeciesnewtoNigeria.Bull.NigerianOrnithol.Soc.6:59-60.Walters,M.P.atnight.1969.HerringandLesserBlack-backedGullsfollowingboatsBrit.Birds62:387.1968Andersson,A.1968a.Gratrutensnaringsekologiochinverkanpakustfagelfaunanmedsarskildhansyntillejdern.[Ph.D.]thesis,Univ.Lund/Lund,Sweden.103pp.1968b.GratrutarnapaHallandsVaderoochderasgrannar.Skanes------Nat.56:100-109.Barth,E.K.1968a.EggdimensionsandlayingdatesofLarusmarinus,L.argentatus, fuscus,and NyttMag. Zool:-rs: 5-34.1968b.ThecircumpolarsystematicsofLarusargentatusandLarus-----fuscuswithspecialreferencetotheNorwegianpopulations.NyttMag.Zool.15(Suppl.1):1-50.Bourne,W.R.P.1968c.Thewing-tippatternsofgulls.Brit.Birds61:138141.Campbell,R.w.1968.StatusofbreedingHerringGullsatBridgeLake,BritishColumbiafrom1933to1963.Can.Field-Nat.82:217-219.Cowan,P.J.1968.FeedingrelationshipbetweenmulletandHerringGulls.Brit.Birds61:31.268

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HerringGullDrost,R.1968.DressurvonSilbermowenLarusargentatusaufakustischeSignale.Vogelwarte24:185-187.[InGerman.]Drury,W.H.,Jr.andW.J.Smith.1968.HerringGullsandintrusionbyyoung.DefenseoffeedingareasbyadultEvolution22:193-201.Focke,E.1968.ZumVerhaltenjungerSilbermowen(Larusargentatus)beiBedrohungdurchaduleArtgenossen.Vogelwarte24:262-266.[In Geroudet,P.1968.L'expansionduGoelandargenteLarusargentatusmichahellisdansIebassinduRhoneetenSuisse.NosOiseaux29:313-335.[InFrench.]Kadlec,J.A. andW.H.Drury.1968a.StructureoftheNewEnglandHerringGullpopulation.Ecology49:644-676.1968b.Aerialestimationofthesizeofgullbreedingcolonies.J.------Wildl.Manage.32:287-293.Karpovich,V.N.andI.P.Tatarinkova.1968.[PopulationdynamicsofGreatBlack-backedandHerringGullsattheislandsontheMurmancoastandtheirinfluenceontheefficiencyofthereproductionoftheCommonEider.]Pp.85-104inE.Kumari(ed.).[TheCommonEider(Somateriam.mollissimaL.) U.S.S.R.]ValgusTallin.[InRussianwithEnglishsummary.]Rabinowitch,V.E.1968.Theroleofexperienceinthedevelopmentoffoodpreferencesingullchicks.Anim.Behav.16:425-428.Rogers,A.E.F.1968.HerringGulltakingsmallpasserineatsea.Brit.Birds61:467.Raux,G.andW.Thonen.1968.LanidificationduGoelandargenteLarusargentatusmichahellisauFanel.NosOiseaux29:335-338.Schreiber,R.W.1968.SeasonalpopulationfluctuationsofHerringGullsincentralMaine.Bird-Banding39:81-106.Southern,W.E.1968.DispersalpatternsofsubadultHerringGullsfromRogersCity,Michigan.Jack-PineWarbler46:2-6.Threlfall,W.1968a.ThefoodofthreespeciesofgullsinNewfoundland.Can.Field-Nat.82:176-180.1968b.StudiesofthehelminthparasitesoftheAmericanHerringGull-----(LarusargentatusPont.)inNewfoundland.Can.J.Zool.46:1119-1126.1968c.ThefoodofHerringGullsinAngleseyandCaernarvonshire. ------Nat. Wales.11:67-73.269

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HerringGullThrelfall,W.1968d.AHerringGullchick(Larusargentatus)withanabnormalbill.Auk85:506-508.vanImpe,J.1968.EenpaarZilvermeeuwen(Larusargentatus)broedendopeenknotwilg.[HerringGullsbreedinginaheadedwillow.]Limosa41:62-63.{InDutchwithEnglishsummary.]vanTets,G.F.1968.Seasonalfluctuationsinthemortalityratesofthreenorthernandthreesouthern-hemispheregulls.CSIROWildl.Res.13:1-9.Voipio,P.1968.ZurVerbreitungderargentatusundcachinnans-Mowen.[ThedistributionofthegroupsofargentatusandcachinnansHerringGUlls.]OrnisFenn.45:73-83.[InGermanwithEnglishandFinnishsummaries.]1967Abel,J.H.1967.CellularchangesintheadenohypophysisoftheHerringGullduringosmoticstress.(Abstractonly.)Anat.Rec.157:201.Barth,E.K.19.67.StandardbodymeasurementsinLarusargentatus,L.fuscus, and marinus.NyttMag.Zool.14:7-83.Blum,V.1967.80SilbermowenimSeewinkel.Egretta10:31.Brosset,A.1967.LeGoelandargente,Larusargentatusmichahellisdestructeurdesapropreponte.Alauda35:73-74.Brown,R.G.B.1967b.SpeciesisolationbetweentheHerringGullLarusargentatusandLesserBlack-backedGullL.fuscus.Ibis109:310-317.1967c.BreedingsuccessandpopulationgrowthinacolonyofHerring-----andLesserBlack-backedGulls,Larusargentatusand fuscus.Ibis109:502-515.Drent,R.H.1967.FunctionalaspectsofincubationintheHerringGull(LarusargentatusPont.).Ph.D.thesis,Univ.Groningen/Netherlands.Griffiths,J.inland. 19U7. HerringandLesserBlack-backedGullsnestingonroofsBrit.Birds60:426.Ingolfsson,A.1967.Thefeedingecologyoffivespeciesoflargegulls(Larus)inIceland.Ph.D.thesis,Univ.Mich./AnnArbor,MI.xiiand186pp.MacMillan,A.T.1967.The"Carrick"gullandothers--Icelandoralbino?Scott.Birds4:493-502.Mclaughlin,F.L.1967ms.BehaviorstudyoftheinterrelationshipsofgullsandeidersinmixedbreedingcoloniesonPenobscotBay,Maine.Unpubl.filerep.,Maine Coop.Wildl.Res.Unit,Orono,ME.270

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HerringGullMoisan,G.andJ.-M.Poulin.1967.Deplacementssaisonniersettauxdemortalitedugoelandargente(Larusargentatus)deslIesRazades,Quebec.Nat.Can.(Que.)94:421-43-5-.---Pettingill,O.S.,Jr.1967.A36-year-oldHerringGull.Auk84:123.Schreiber,R.W.1967.RoostingbehavioroftheHerringGullincentralMaine.WilsonBull.79:421-431.Southern,W.E.1967.TheroleofenvironmentalfactorsinRing-billedandHerringGullorientation.Ph.D.thesis,CornellUniv./lthaca,NY.Stromar,L.1967.FouryearsofbandingHerringGulls(LarusargentatusPontopp.)ontheisletsofMrkanandBobara.Larus [EnglishtranslationoftheSerbo-Croatian.]Threlfall,W.1967a.StudiesonthehelminthparasitesoftheHerringGullLarusargentatusPontopp.,innorthernCaernarvonshireandAnglesey.Parasitology57:431-453.1967b.DiseasesandpathologicalconditionsoftheHerringGull -----(Larus argentatusPontopp.)excludinghelminthinfestations.Bull.Wildl.Dis.Assoc.3:62-67.Viellard,J.1967.GuignettesetGoelandsargentesdanslesAlpes.OiseauRev.Fr.Ornithol.37:148-149.Weaver,D.K.1967.TheHerringGull,withemphasisonmortalityoftheeggsandyoung.M.S.thesis,Univ.Mich./AnnArbor,MI. Wolk,R.G.andJ.Bull.1967.DifferentialnestingschedulesofHerringGullson LongIsland,NewYork.Kingbird17:5-6.1966Abel,J.H.1966.AphysiologicalandcytochemicalstudyofcompensorygrowthinthenasalglandsoftheHerringGull,Larusargentatus.Ph.D.thesis,BrownUniv./Providence,RI.Barth,E.K.1966.MantlecolourasataxonomicfeatureinLarusargentatusandLarusfuscus.NyttMag.Zool.13:56-82.Bernis,F.1966.PenetraciondeLarusargentatushastalaprovinciadeToledo.Ardeola12:239.Blosch,M.1966.DieAktivitatderSalzdrusen.EineUntersuchunganfreilebendenundgefangenenSilbermowen,Larusargentatus.Vogelwarte23:225231.[InGerman.]271

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HerringGullBusnel, R.-G. andJ.Giban.1966. Reactions d'unpopulationdeLarusargentatusatlantisdel'iledeMadere adessignauxacoustiquesdedetressed'oiseauxeuropeens.Bocagiana11:1-9.[InFrenchwithEnglishand Germansummaries.]Delius,J.D.1966a.PentobarbitalanaesthesiaintheHerringandLesserBlack-backedGull.J.SmallAnim.Pract.7:605-609.1966b.Displacementactivitiesandarousal.Nature(Lond.)214:------1259-1260.Harriman,A.E. andM.R.Kare.1966.ToleranceforhypertonicsalinesolutionsinHerringGulls,StarlingsandPurpleGrackles.Physiol.Zool.39:117-122.Keith,J.A.1966a. Reproduction inapopulationofHerringGulls(Larusargentatus)contamintedbyDDT.J.Appl.Ecol.3(Suppl.): 1966b. Reproduction inapopulationofHerringGulls,Larusargent-----atus,contaminatedbyDDT.M.S.thesis,Univ. King,B.1966.HerringGullstryingtosnatchfishfromGreatNorthernDivers.Brit.Birds59:247.Ludwig,J.P.1966.Lakes1960-1965.HerringandRing-billedGullpopulationsoftheGreatGreatLakesRes.Div.Univ.Mich.Publ.15:80-89.Ludwig,J.P.and C.S.Tomoff.1966. Reproductive successandinsecticideresiduesinLakeMichiganHerringGulls.Jack-PineWarbler44:77-85.Miller,J.C.1966a.HerringGullsnestingatStoneHarbor,NewJersey.Cassinia49:31.1966b.TheGreatBlack-backedGullnestinginNewJerseyandaddi-----tionalnotesonnestingHerringGulls.Cassinia49:31.Paynter, R. A.1966.A newattempttoconstructlifetablesforKentIslandHerringGulls.Bull.Mus.CompoZool.133:491-528.Schoennagel,E.1966.Lachmowen(Larusridibundus)undSilbermowen(Larusargentatus)fangenDreistachligeStichlinge(Gasterosteus Ornithol.Mitt.18:144.Segre,A., R. Noble andJ.P.Hailman.1966.Afive-eggHerringGullnest.Bird-Banding37:290.Smith,N.G.1966.Adaptationstocliff-nestinginsomeArcticgulls(Larus).Ibis108:68-83.272

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HerringGullTemme,M.1966.UberdasVerhaltensrepertoirederMantelmowe(LarusmarinusL.)imWinterhalbjahrsowieeinigeBemerkungenuberdasVerhaltenderSilbermowe(LarusargentatusPontopp.)imgleichenZeitraum.J.Ornithol.107: German.]Threlfall,W.1966.tatusPontopp.).ThehelminthparasitesoftheHerringGull(LarusargenTech.Commun.Commonw.Bur.Helminthol. 1965Abel,J.H.andR.A.Ellis.1965.Aphysiologicalandhistochemicalstudyofregenerationinthenasal(salt)glandsofHerringGulls.Bull.Mt.DesertlsI.BioI.Lab.5:1.Bergman,G.1965.Trutarnaskonkurrensforhallandenfodobehovochrelationertillandraskargardsfagler.[CompetitionandfeedingecologyofgullsinasouthFinnishskerrygard.]Zool.Revy 27:58-77.[InSwedishwithGerman summary.]Brisbin,I.L.,Jr.1965.AquantitativeanalysisofecologicalgrowthefficiencyintheHerringGull.M.S.thesis,Univ.Georgia/Athens,GA.Gobb,J.L. S.1965.AbnormalnestingbehaviourofeidersandHerringGulls.J.Scott.Ornithol.Club3:252-253.Drury,W.H.1965a.ResultsofastudyofHerringGullpopulationsandmovementsinsoutheasternNew England. Pp.207-219inR.G.BushnelandJ.Giban(eds.).Golloque:Ieproblemedes oiseaux surlesaerodromes.Inst.Natl.Rech. Agron.Paris.326pp.1965b.Gullsvs.terns:clashofcoastalnesters.Mass. Audubon 64:------207-211.Harris,M.P.1965.ThefoodofsomeLarusgulls.Ibis107:43-53.Harris,M.P.andW.J.Plumb. 1965.ExperimentsontheabilityofHerringGullsLarusargentatusandLesserBlack-backedGullsL.fuscustoraiselargerthannormalbroods.Ibis107:250-257.-------Rogers,C.H.1965.HerringGullnestingcolonyinBarnegetBay.Cassinia48:38.Schreiber, k. W.1965.SeasonalpopulationfluctuationsandroostingbehavioroftheHerringGullincentralMaine.M.S.thesis,Univ.Maine/Orono, Thomas,L.P.and S. B. Thomas.1965.HerringGullsdivingforstarfish.Q.J.Fla.Acad.Sci.28:195-196.273

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HerringGullVass,S. E.1965.GullbreedingrecordsfromPrinceEdwardIsland.Can.Field-Nat.79:152-154.Williams,I.C.andM.P.Harris.1965.TheinfectionofthegullsLarusargentatusPont.,L.fuscusL.,and L.marinusL.withcestodaonthecoastofWales.Parasitology55:237-256.1964Andersson,S.1964.EttgratrutmaterialfromVasterviksskargard.[AmaterialofHerringGullfromtheArchipelagoofVastervik.]VarFagelvarld23:337-347.[InSwedishwithEnglishsummary.]Baerends,G.P.1964.Lareconnaissancedel'oeufparIeGoelandargente.Bull.Soc.Sci.Bretagne37:193-208.Barth,E.K.1964.Larusfuscus.119-12-1-.--VariationinthemantilecolorofLarusargentatusandPreliminaryreport.K.Nor.Vidensk.Selsk.Forh.37:Bonting,S.L.,L.L.Caravaggio,M.R. Canady,andN.M.Hawkins.1964.Studiesonthesodium-potassium-activatedadenosinetriphosphate.XI.ThesaltglandoftheHerringGull.Arch.Biochem.Biophys.106:49-56.Bowes, andP.W.Schwalbe.1964.GreatBlueHeronchasingaHerringGull.Cassinia47:37-38.Brackbill,H.1964.Pseudo-sleepingby aHerringGull.Condor 66:309.Goethe,F.1964.Mittelmeer-Silbermowe(LarusargentatusmichahellisNaum.)inCuxhavengefangen.Bietr.Naturkd.Nieders.17:28-31.Gross,A.o.1964.AlbinismintheHerringGull.Auk81:551-552.Harris,M.P.1964a.AspectsofthebreedingbiologyofthegullsLarus gentatus, fuscus,and L.marinus.Ibis106:432-456.1964b.Theincidenceofsomespeciesoftrematodeainthreespecies-----ofLarusgullsinWales.Ibis106:532-536.1964c.RecoveriesofringedHerringGulls.BirdStudy11:183-191.Hudec,K.1964.Durchzug undUberwinternderSilber-und Sturmmowe(Larusargentatus,Laruscanus)anderDonau.Zool.Listy13:363.[InCzech withGerman summary:-] Hunt,C.L.,Jr.andW.J.Smith.1964.Theswallowingoffishby youngHerringGullsLarusargentatus.Ibis106:457-461.274

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HerringGullJarback,A.1964.Gratrut(Larusargentatus)fangandegrasparv(Passerdomesticus)iflykten.[HerringGull(Larusargentatus)catchesHouseSparrow(Passerdomesticus)in flight.]--vir Fagelvarld23:324.[InSwedishwithEnglishsummary.]Kadlec,J.A.1964ms. Theeffect of amplifiedsoundson movementsofHerringGulls.Unpubl.prog.rep.U.S.FishWi1d1. Servo Komnick,H.1964.ElektronenmikroskopischeUntersuchungenzuefunktionellenMorphologiedesIonen-transportesinderSalzdrusevonLarusargentatus.IVTeil:FunktionelleMorphologiederEpithelze1lendesSammelkanals.Protoplasma58:96-127.Mathiasson,S.1964.Gratrutochgratruts-bekampning.Zool.Revy 26:41-51.[InSwedishwithEnglishsummary.]Nickell,W.P.1964a.FatalentanglementsofHerringGulls(Larusargentatus)andCommonTerns(Sternahirundo).Auk81:555-556.1964b.TwoalbinoHerringGullsfoundatRogersCity,Michigan.Auk------81:560.Pomeroy,M.K.andM.D. B. Burt.1964.CestodesoftheHerringGull,LarusargentatusPontoppidan,1763,fromNewBrunswick.Can.J.Zool.42:959973.Pricam,R.1964.PremierenidificationduGoelandargente(Larusargentatus)surlesrivesdulacLeman.OiseauRev.Fr.Ornithol.34:151-153.Rawson,D.1964.SequencesinthematurationofthegenitaliainTetrabochriuserostris(Loennberg,1889)fromtheintestineofLarusargentatusar gentatus Pontoppidan.Parasitology54:453-465.Richards,G.A.1964.FulmarincubatingeggsofHerringGullwithitsown.Brit. Birds 57:31.Southern,W.E. andG.D.Schnell.1964.HerringGullsfeedingonsealamprey.Jack-PineWarbler42:231.Steinbacher,G.1964.ZumNahrungsbedarfderSilbermowe(Larusargentatus).Vogelwelt85:23-24.1963Ames,J.E.1963.HerringGullnestingontheNorthCarolinacoast.Chat27:29.Blondel,J.1963.LeproblemeducontroledeseffectifsduGoelandargente(LarusargentatusmichahellisNaumann)enCamargue.TerreVie110:301 French.]275

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HerringGullDrury,W.H.,Jr.1963a.easternNewEngland.Wildl.Servo 68pp.HerringGullpopulationsand movementsinsouthMass.AudubonSoc.cyclostyledrep.toU.S.FishEmlen,J.T.1963.DeterminantsofcliffedgeandescaperesponsesinHerringGullchicksinnature.Behaviour22:1-15.Goethe,F.1963.VerhaltensunterschiedeswischeneuropaischenFormanderSilbermowengruppe(Larusargentatus-cachinnans-fuscus).J.Ornithol.104:129-141.[InGermanwithEnglishsummary.]Hailman,J.P.Carolina.1963.HerringGullextendsbreedingrangesouthtoNorthAuk80:375-376.Komnick, H,1963.ElektronenmikroskopischeUntersuchungenzurfunktionellenMorphologiedesIonen-transportesinderSalzdrusevonLarusargentatus.ITeil:BauundFeinstrukturderSalzdruse.Protoplasma56:274-314.Lemmetyinen, R.1963.LokkienesiintymisestajaravinnostaGullkronanselankoillisosassa.[OccurrenceandfeedinghabitsofgullsinthenortheasternpartoftheareaofGullkrona.]SuomenRiista16:69-82.[InFinnishwithEnglishsummary.]Libby,W.L.andW.T.vanVelzen.1963.Theeffectofamplifiedsoundson movementsofHerringGulls.U.S.Dep.Int.Bur.SportFishWildl.,PatuxentWildl.Res.Cent.,SectionAnimalDepredationsCtrl.Studies.Prog.Rep. Aug.1,1962-July31,1963. WorkUnitF-34.3.20pp.Ludwig,J.P.1963.ReturnofHerringGullstonatalcolony.Bird-Banding34:68-72.Minoranskii,V.A.1963.[NestingoftheHerringGull(Larusargentatus)inLakeManych-Gudilo.]N.D.V.S.BioI.Nauki3: Russian.]Paynter,R.A.,Jr.1963.NorthAmericanHerringGullsnestingon abuilding.WilsonBull.75:88.Pemberton,R.T.1963.HelminthparasitesofthreespeciesofBritishgulls,LarusargentatusPont.,L.fuscusL. and L.ridibundusL.J.Helminthol.37:57-88.----Williamson,F.S.L. and L.J.Peyton.1963.InterbreedingofGlaucous-wingedandHerringGullsintheCookInletregion,Alaska.Condor 65:24-28.1962Clement,R.C.andR.E.Woodruff.1962.AhistoryofthenestinggullsandternsofRhodeIsland,1889-1961.NarragansettNat.5:108-111,125-128.276

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HerringGullDennis,J.V.andW.Pepper.1962.More onthetravelsofHerringGulls.EBBANews25:139-144.Gerdes,K.1962.RichtungstendenzenvomBrutplatzverfrachterLichmowen(Larusridibundus)unterAusschlussvisuellerGelande-und Himmelsmarken. Z.Wiss.Zool.166:352-410.[InGerman.]Harris,M.P.1962a.DifficultiesintheageingoftheHerringGullandLesserBlack-backed'Gull.BirdStudy9:100-103.1962b.HerringGullwithabnormalbill.Brit.Birds55:236.1962c.-----islands.TheLarusgullsbreedingon Skomer, SkokholmandGrassholmNat.Wales8:56-58.Ludwig,C.C.1962.HerringGulls--1961recoveries.Inl.BirdBandingNews 34: 55-56.Ludwig,J.P.1962b.AsurveyofthegullandternpopulationsofLakesHuron,Michigan,andSuperior.Jack-PineWarbler40:104-119.Riley,J. W. andD.L.Miller.1962.UnusualdeathofHerringGull.Brit.Birds55:590-591.Smith,W.J.1962.Driftinggulls.Nat.Hist.71:54-59.Thesleff,S.andK.Schmidt-Nielsen.1962.ofthesaltglandoftheHerringGull.Anelectro-physiologicalstudyAm.J.Physiol.202:597-600.Vauk,G.1962.DasSilbermowenproblemaufHelgoland.Ber.Internatl.Rat.VogelschutzHelgol.2:1-6.Voous,K.H.1962.AnotherpresumedhybridofLesserBlack-backedGullandHerringGullintheNetherlands.Ardea50:171-172.1961Baerends,G.P.1961.Perceptic-perikelenbijdeZilver-meeuw.Versl.GewoneVergad.Akad. Amst.70:133-138.Drost,R.,E.FrockeandG.Freytag.1961.EntwicklungundAufbaueinerPopulationderSilbermowe,Larusargentatusargentatus.J.Ornithol.102:404-429.[InGermanwithEnglishsummary.]Ehlert,W.1961.WeitereUntersuchungenuberdieNahrungsweltderSilbermowe(Larusargentatus)aufMellum.Vogelwarte21:48-50.[InGerman.]277

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HerringGullGillespie,M.1961.TravelsofHerringGulls.EBBANews24:88.Goethe,F.1961.Zur TaxonomiederSilbermowe(Larusargentatus)imsudlichendeutschenNordseegebiet.Vogelwarte21:1-24.[InGermanwithEnglishsummary.]Isenmann,P.and B.Schmidt.1961. argentatusXLarusmarinus.Observationsaproposd'unhybrideLarusOiseauRev.Fr.Ornithol.31: Lesperance,T.1961.HerringGullscatchingflyingants.Kingbird11:150.Ludwig,C.C.andF.E.LudWig.1961.HerringGullsandCaspianTerns--1960recoveries.Inl.BirdBandingNews33:28-29.Macpherson,A.H.1961.ObservationsonCanadianArcticLarusgulls,andonthetaxonomyofL.thayeriBrooks.Arct.Inst.N.Am.Tech.Pap.7:1-40.Nero,R.W.1961.Drylandgullcololy.BlueJay19:166-168.Rooke,K.B.1961.AprobablehybridGreatBlack-backedGullXHerringGullinDevon.Brit.Birds54:161-163.Spitzenberger,F.1961.ZurErnahrungeineristrischenSilbermowen-Kolonie(Larusargentatusmichahellis).Vogelwarte21:50-52.[InGerman.]Stevens,C.J.1961.Unusualnest-sitesofHerringGullsinCornwall,includingatreeandvariousbuildings.Brit.Birds54:244-245.Tinbergen,N.1961.TheHerringGull'sWorld.(Revisededition.)BasicBooks,NewYork.Voous,K.H.1961.Micro-geographicalvariationinNetherlandsHerringGulls,Larusargentatus.Ardea49:69-72.Wilcox,L.1961.HerringGullscatchflyingants.Kingbird11:199.1960Baerends,G.P.,K.H.Postuma andT.Joustra.1960.brutendenSilbermoweaufdieTemperaturdesEies.13:586-588.[InGerman.]DieReaktionderArch.Neerl.Zool.Bergman,G.1960.UberneueFuttergewohnheitenderMowenandenKustenFinnlands.OrnisFenn.37:11-28.[InGermanwithFinnishsummary.] Brown,P.B.1960.FulmarincubatingeggsofHerringGull.Brit.Birds53:127.278

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HerringGullDrury,W.H.,Jr.1960.BreedingactivitiesofLong-tailedJaeger,HerringGullandArcticTernonBylotIsland,NorthwestTerritories,Canada.Bird-Banding31:63-79.Goethe,F.1960.Silbermowe.FelsbrutertumundweiterebeachienswerteTendenzenbeiderProc.Internatl.Ornithol.Congr.12:252-258.[InGerman.]Grahn,Y.1960.Hackningavstrandskata(Haematopusostralegus)ochgratrut(Larusargentatus)isjonSalen,KronobergsIan.[BreedingofOystercatcher(Haematopusostralegus)andHerringGull(Larusargentatus)atLakeSalen,Smaland.]VarFagelvarld19: [InSwedjshwithEnglishsummary.]Hansen,L.1960.Krydsningmellum Solvmage(LarusargentatusPont.)ogSildemage(LarusfuscusL.)fundetynglendesammenmedenSolvmage.[AhybridbetweenHerringGull(LarusargentatusPont.)andLesserBlackbackedGull(LarusfuscusL.)breedingwithaHerringGull.]Dan.Ornithol.Foren.Tidsskr.54:85-87.[InDanishwithEnglishsummary.)Jehl,J.R.,Jr.1960.AprobablehybridofLarusargentatusand L.marinus.Auk77:343-345.Johansen,H.1960.DieVogelfaunaWestsiberiens.TeilIII.(NonPasseres).9.Fortsetzung:Alcidae,Laridae.J.Ornithol.101:316-339.[InGerman. ]Mylne,C.K.1960.TechniquesofHerringGullsandJackdawspreyingonPuffins.Brit.Birds53:86-88.Stegmann,B.1960.ZurSystematikdesRassenkreisesLarusargentatus.J.Ornithol.101:498-499.[InGerman.]1959Baerends,G.P.1959.Theethologicalconcept"releasingmechanism"illustratedbyastudyofthestimulielicitingegg-retrievingintheHerringGull.Anat.Rec.128:518-159.Borg,K.1959.Nabblangdochkroppsvikt hos gratrut(Larusargentatus),havstrut marinus)ochsilltrut(L.fuscus).[BilllengthsandbodyweightsforHerringGulls(Larus argentatUST7 GreatBlack-backedGulls(L.marinus)andLesserBlack-backedGulls(L.fuscus).]VarFagelvarld 18: 311-317.[InSwedishwithEnglishsummary.-]-----Elliott,J.J.1959.Waysofthe"SeaGulls".Kingbird9:116-120,142-148.Focke,E.1959.ZurErnahrungderSilbermowe(Larusargentatus).Vogelwarte20:86-88.[InGerman.]Hobbs,J.N.1959.Gullsattackingmigrantthrushes.Brit.Birds52:313.279

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HerringGullHofslund,P.B.Minnesota.1959.FallmigrationofHerringGullsfromKnifelsland,Bird-Banding30:104-114.Smith,W.J.1959.MovementsofMichiganHerringGulls.Bird-Banding30:69-104.Spaans,A.L.1959.HetZilvermeeuwenproblemofVlieland.LevendeNat.62:25-29Voous,K.H.1959.GeographicalvariationoftheHerringGull,Larusargentatus,inEuropeandNorthAmerica.Ardea47:176-187.1958 Amadon,D.1958.TheencroachmentofHerringGulls(Larusargentatus)on moredesirablesea-fowlintheeasternUnitedStates.Internatl.Comm.BirdPreserv.Bull.7:114.Drost,R.1958.UberdieAnsiedlungvonjunginsBinnenlandverfrachtetenSilbermowen(Larusargentatus).Vogelwarte19:169-173.[InGerman.]Duchrow, H.1958.StosstauchenundBeuteraubenbeiMowen.Vogelwelt79:183184.Fange,R.,K.Schmidt-NielsenandH.Osaki.1958.ThesaltglandoftheHerringGull(Larusargentatus).BioI.Bull.(WoodsHole)115:162-171.Frings,H.,M.Frings,J.Jumber,R. G.Bushnel,J.Gibon andP.Gramet.1958.ReactionsofAmericanandFrenchspeciesofCorvus andLarustorecordedcommunicationsignalstestedreciprocally.Ecology39:125-131.Goethe,F.1958.AnhaufungenunversehrterMuschelndurchSilbermowen.[MusselsandLarusargentatus.]Nat.Yolk88:181-187.[InGerman.]Kohl,I.1958.ringGulls.ContributionstosystematicstudiesoftheBlackSea'sHerAquila65:127-143.Kruijt,J.P.1958.SpecklingoftheHerringGullegginrelationtobroodingbehavior.Arch.Neerl.Zool.12:565-567.MorzerBruyns,M.F.1958.Gullswhicharea menacetootherspecies.TheHerringGullproblemintheNetherlands.Internatl.Comm.BirdPreserv.Bull.7:103-107.280

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HerringGullOlsson,V.1958.Dispersal,migration,longevityanddeathcausesofStrixaluco,Buteobuteo,Ardeacinerea,andLarusargentatus.Astudybasedon ringedinFenno-Scandia.ActaVertebr.1:91-189.Salmon,H.M.1958.LesserBlack-backedandHerringGullsnestingon afactory-roofinlandinGlamorgan.Brit.Birds51:399-401.1957Andersen,F.S.1957.Eggsizeandtheagecompositionofbirdpopulation.Vidensk.Medd. Dan.Naturhist.Foren.119:1-24.Andersson,R.1957.Gratrutar(Larusargentatus)ochhornugglor(Asiootus)medovanligajaktmetoder.[HerringGulls(Larusargentatus)andLongearedOwls(Asiootus)withunusualmethodsofhunting.]VarFagelvarld16: 307-308.--yIn SwedishwithEnglishsummary.]Borg,K.1957.Omnabblangdenhosgratruten(Larusa.argentatusPont.).[OnthebilllengthoftheHerringGull(Larus argentatusPont.).]VarFagelvarld16:36-43.[InSwedishwithEnglishsummary.]Dutemeyer,C.1957.SilbermowenbrutenaufHausdacherninBremerhaven.Ornithol.Mitt.9:135.Ehlert,W.1957.ZurErnahrungderSilbermowe(LarusargentatusPontopp.)inderVorbrutzeit.[ThefoodoftheHerringGullintheprebreedingseason.]Ornithol.Mitt.9:201-203.[InGerman.]Hickling,R.A.o.1957.Thesocialbehaviorofgullswinteringinland.BirdStudy4:181-192.Kumerloeve,H.1957.SilbermowenbrutenaufHausdachern.Ornithol.Mitt.9:112.Mills,D.H.1957.lobsterlarvae.HerringGullsandCommonTernsaspossiblepredatorsofJ.Fish.Res.BoardCan.14:729-730.Poulding,R. H.1957.Tuberculosisingulls:apreliminaryinvestigation.Bull.Br.Ornithol.Club77:144-149.Uspenskii,S.M.1957.Sezonnoerazmeshchenieimigratsiiskandinavskoiserebristolchaiki(LarusargentatusomissusPleske)po dannymkol'tsevaniyav SSSR.[SeasonaldistributionandmigrationsoftheScandinavianHerringGull(LarusargentatusomissusPleske)accordingtobandingdataintheUSSR.]TrudyByuroKol'tsevaniya9.Moskva. 1956Davis.T.A.w.1956.Gullsfeedingongrain.Brit.Birds49:400-404.281

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HerringGullEmlen,J.T.,Jr.1956.JuvenilemortalityinaRing-billedGullcolony.WilsonBull.68:232-238.Goethe,F.95pp.1956a.DieSilbermowe.[InGerman.]WittenbergLutherstadt:A.ZiemsenVerlag._____1956b.Fuchs,Vulpesvulpes(Linne,1758),reibtSchlafgesellschaftvonetwasechzigjugenlichenSilbermowen(LarusargentatusPontopp.)auf.Saugetierkd.Mitt.4:58-60.[InGerman.]-----1956c.FremdeSilbermowenformenandeutschenKusten.Dochsibirische-----Silbermowe(LarusargentatusbirulaePleske)beiHelgoland.Vogelwarte18:152-154.1956d.Eine nberannte SilbermowevonHelgoland("LarusargentatusdrostiKleinschmidt").Vogelwarte18:154-156.1956e.Eismeer-SilbermowenalsWintervogelinEckernfordeundauf-----Wanderoog.Vogelwarte18:156.Jungfer,W.1956.aufHelgoland.Schwarzmeer-Silbermowe(LarusargentatusponticusStegm.)Vogelwarte18:156-157.Moynihan,M. NotesonthebehaviorofsomeNorthAmericangulls.I.Aerialhostilebehavior.Behaviour10:126-178.1956b.CaliforniaGullsandHerringGullsbreedinginthesamecolony.------Auk73:453-454.Pricam,R.1956a.LeGoelandargenteravisseurdecanetons.NosOiseaux23:218-219.[InFrench.]1956b.UnGoelandargentemediterraneensurlaLoire.Alauda24:73.[InFrench.]Tinbergen,N.1956.Onthefunctionsofterritoryingulls.Ibis98:401-411.1955Drost,R.1955.NeueBeitragezurSoziologiederSilbermoweLarus argentatus.Proc.Internatl.Ornithol.Congr.11:564-569.[InGerman.]Frings,H.,M.Frings,B.CoxandL.Peissner.1955a.Auditoryandvisualmechanismsinfood-findingbehavioroftheHerringGull.WilsonBull. 67: 155-170.1955b.RecordedcallsofHerringGulls(Larusargentatus)asrepel-----lantsandattractants.Science121:340-341-.----Gladkov,N.A.andV.S.Zaletaev.1955.0rybokhozyaistvennomznacheniiserebristoichaikinaKaspiikommore.[TheimportanceoftheHerringGulltothefishingindustryoftheCaspianSea.]Vopr.Ikhtiol.4.[InRussian.]282

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HerringGuUGoethe,F.1955a.BeobachtungenbeiderAufzuchtJungerSilbermowen.[ObservationsonthedevelopmentofbehaviorinyoungLarusargentatus.]Z.Tierpsychol.12:402-433.[InGermanwithEnglishsummary.]1955b.VerglichendeBeobachtungenzumVerhaltenderSilbermowe(Larus-----argentatus)undderHeringsmowe(Larusfuscus).Proc.Internatl.Ornithol.Congr.11:577-582.[In German.]------------Gross,A.O.1955.ChangesofcertainseabirdpopulationsalongtheNewEnglandcoastofNorthAmerica.Proc.Internatl.Ornithol.Congr.11:446-449.Haan, H.J.H.de.1955.DezilvermeeuwLarusargentatusargentatusBrehm.broedendindeOspelerPeelMaandbl.44:112-114.Huntington,C. E.1955.AgediscriminationinabreedingcolonyoftheHerringGullLarusargentatus.Proc.Internatl.Ornithol.Congr.11:467-469.Tinbergen, N. 1955.Zilvermeeuwennieuws.LevendeNat.58:21-30.[InDutch.]Took,G.E.1955.NestingofHerringGullsonroof-topsinDover.Brit.Birds48:88-89.Voipio,P.1955.ThepopulationoftheYellow-leggedHerring-Gullon alargelakeinFinland.Proc.Internatl.Ornithol.Congr.11:537-539.1954Baerends,G.P.1954.EggrecognitionintheHerringGull.Proc.Internatl.Congr.Psychol.14:93-94.Bodenstein,G.1954.Sommerbeobachtungen vonLarusargentatusanteliusundLarushyperboreusinderSudlichten Ostsee:--vDgelwarte 17:44.Jungfer,W.1954.DieEntslehungeinesWarstnestesderSilbermowe.Nat.Yolk84:432-433.King,B.1954.UnusualbehaviorofHerringGull.Brit.Birds47:26.Maschlanka,H.1954.ProportionsmessungenanSilbermowen.Zool.Jahrb.Abt.AUg.Zoo1.Physio!.Tiere65:209-218.[InGerman.]Meijering,M.P.1954.ZurFragederVariationeninderErnahrungderSilbermowe,LarusargentatusPont.Ardea42:163-175.[InGerman.]Poulding,R.H.1954a.LossofringsbymarkedHerringGulls.BirdStudy1:37-40.1954b.ParasitismofaHerringGullbytheduckleech.Brit.Birds ------47: 306-307.1954c.SomeresultsofmarkinggullsonSteepholm.Proc.Bristol ------Nat. Soc.29:49-56.283

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HerringGullSchmidt,G.1954.ZumRaubenderSchmarotzer-Raubmowe.Vogelwelt75:147151.[InGerman.]Stokoe,R.1954.AblackHerringGullinCumberland.Brit.Birds47:132-133.Taber,W.1954.HerringGullsandthehurricane.Bull.Maine AudubonSoc.10:62-63.Voipio,P.1954a.UberdiegelbfussigenSilbermowenNordwesteuropas.ActaSoc.FaunaFloraFenn.71:1-56.[InGerman.]1954b.BeobachtungenuberdieSilbermowe,LarusargentatusPont.,amGross-SaimaaundihreBeziehungenzudenanderenMowenvogeln.DrnisFenn.31:57-80.[InGerman.] Young, C.ber.1954.ImmatureHerringGullsolicitingfoodfromadultinDecemBrit.Birds47:25.1953Bauer,K.1953.DieMittelmeer-Silbermowe(Larusargentatusmichahellis)imOesterreich.Vogelkd.Nachrichtenbl.Oesterr.3:1-2.Cross,A.1953b.InlandnestingofHerringGull.Scott.Nat.65:195.Eggeling,W.J.1953.HerringGullnestingon arooftop.Scott.Nat.65:195.Goethe,F.1953a.SozialeHierarchieinAufzuchtschwarmderSilbermowen.Z.Tierpsychol.10:44-50.[InGerman.]1953b.ExperimentelleBrutbeendigungundanderebrutethologische-----BeobachtungenbeiSilbermowen(Larusa.argentatusPontopp.).J.Ornithol.94:160-174.-------Paludan,K.1953.NogleresultaterafKobenhavnsZoologiskeMuseumsringmaerkningafLarusargentatus.Vidensk.Medd. Dan.Naturhist.Foren.115: 181-204.--yyn DanishwithEnglishsummary.]Rapp,W.F.,Jr.1953.Ring-billedGulls.FieldseparationoftheHerring,CaliforniaandNebr.BirdRev. 21:32-33.Tinbergen,N.1953.TheHerringGull'sWorld.AStudyoftheSocialBehaviorofBirds.Collins,London.NewNat.Monogr.xviand255pp.1952Boase,H.1952.Notesonthecourtshipdisplayofgulls.Brit.Birds45:320-323.284

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HerringGullDawn,W.1952.HerringGullregurgitatesuponBlack-backedGull'sdemand.Linn.Newsl.6(6/7):2-3.Drost,R.and E.Schuz.1952.DasVerha1tendermannlichenundweiblichenSilbermowen(Larusa.argentatusPont.)ausserhalbderBrutzeit.Vogelwarte16:108-116.Geroudet,P.1952.LeGoe1andargentesurIecourssuperieurduRhone.Alauda20:171-173.Kroneisl,R.1952.ContributiontothestudyofthetaxonomyofthegullsoftheAdriaticcoast.LarusargentatusPontoppidan--HerringGull.Larus4-5:105-118.[Englishtranslation.]Nisbet,I.1952.HerringGulldroppingandcatchingobjectinbill.Brit.Birds45:74.Pimlott,D.H.1952.TheeconomicstatusoftheHerringGullsoftheGrand MananArchipelago,NewBrunswick,1949.Can.Wildl.ServoWildl.Manage.Bull.(Ser.2)5:1-76.Steiniger,F.1952.BildervomTauchenderSilbermowe.Vogelwelt73:157159.[InGerman.]Tinbergen,N.1952.OnthesignificanceofterritoryintheHerringGull.Ibis94:158-159.1951 Bergman,G.1951.UntersichiedevonSilvermowe(Larusa.argentatus)und Heringsmoewe(Larusf.fuscus)inLebensweiseund Stimme.Vogelwarte16:17-18.[InGerman.-]---Bronoel,J.K.1951.HerringGullcensusonKnifeIsland.Flicker23:58.Drost,R.lauf.1951.Beobachtungenaneinerk1einerSilbermowen-PopulationJahresVoge1warte16:44-48.Fritsch,R.H.1951.BeobachtungenaneinerneugebildetenBrutkoloniederSi1bermowe(Larusa.argentatusPontopp.).Z.Tierpsychol.8:252-273.[InGerman.]----Gross,A.O.1951.TheHerringGull-cormorantcontrolproject.Proc.Internatl.Ornithol.Congr.10:532-536.Paludan,K.1951.ContributionstothebreedingbiologyofLarusargentatusandLarusfuscus.Vidensk.Medd. Dan.Naturhist.Foren.114:1-128.285

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HerringGullPeters,H.M.andR.Muller.1951.DiejungeSilbermowe(Larusargentatus)als"Platzhocker".Vogelwarte16:62-69.[InGerman.]Vleugel,D.A.1951.AcaseofHerringGullslearningbyexperiencetofeedaftertheexplosionofmines.Brit.Birds44:180.1950Adams, R.G.1950.Herring-Gull"paddling"ongrass.Brit.Birds43:163.Harberd,R.T.1950.Herring-Gullseatingcharcoal.Brit.Birds43:381.Harberd,R.T.andK.J.Witts.1950.BathingbehaviorofHerringGull.Brit.Birds43:381-382.Hofslund,P.B.1950.HerringGullnestsonKnifeIsland.Flicker22:128.Meinertzhagen,R.1950.OnaraceoftheLarusargentatus-fuscusgroupnewtotheBritishIsles.Bull.Br.Ornithol.Club70:17.------Nordberg,S.1950.ResearchesinthebirdfaunaofthemarinezoneintheAlandArchipelago.ActaZool.Fenn.63:1-59.Peters,H.M.and H.Lange.1950.UberdasVerhaltendorSilbermowe(Larusa.argentatus)zuihremGelege.(ZurFragedesKannibalismus.)Z-.---Tierpsychol.7:121-130.Simmons,K.E.L.1950."Up-ending"ofHerringGull.Brit.Birds43:163.Spencer,K.G.andA.Welch.1950.First-yearHerringGullsolicitingfoodinMay.Brit.Birds43:135.Stevens,C.J.1950.GullinCornwall.InlandbreedingandsubterraneannestingofHerringBrit.Birds43:94-95.Tinbergen,N.andA.C.Perdeck.1950.OnthestimulussituationreleasingthebeggingresponseinthenewlyhatchedHerringGullchick(Larus gentatusB.)[sic].Behaviour3:1-39.Yeager,L.E.1950.BaldEagleattackscrippledgull.WilsonBull.62:210.1949Hazelwood,A.1949.AgileflightmanoeuvreofHerringGull.Brit.Birds42:159.Ingram,C. andG.T.Kay.1949.GreatSkua'smethodofkillinglargebirds.Brit.Birds42:223-224.286

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HerringGullKrug,H.H.1949.Recoveryof19-year-oldHerringGull.Bird-Banding20:103.Medcof,J.C.1949."Puddling"--amethodoffeedingbyHerringGulls.Auk66:204-205.Parks,G.H.1949.DivingHerringGulls.Bird-Banding20:149.Paynter,R.A.,Jr.1949.Clutch-sizeandtheeggandchickmortalityofKentIslandHerringGulls.Ecology30:146-166.Poulding,R.H.1949.HerringGullwithyellowlegsinGloucestershire.Brit.Birds42:335.Rooke,K.B.1949.HerringGullwithyellowlegsinDorset.Brit.Birds42:29-30.Walker,A.B.1949.HerringGull"paddling"ongrassfield.Brit.Birds42:222-223.1948Bronoel,J.K.1948.HerringGullnestsonKnifeIsland,LakeSuperior.Flicker20:110-111.Buckalew,J.H.1948.NestingoftheHerringGullinVirginia.WoodThrush4:22.Hunt,O.D.1948.Unusualnesting-siteofHerringGull.Brit.Birds41:125-126.Isakov,Yu.A.1948.0kolonial'nomgnezdovaniichaekkhokhotuniinaostroveTaraba.[OnthecolonialbreedingofHerringGullsonTarabaIsland.]inSbornik"Chaikakhokhotun'yaiflamingonaKaspiiskommore",Moskva. TIn Russian.]Manville,R.H.1948.The HuronIslandrookery.Jack-PineWarbler26:152-155.Otterlind,G.1948a.Tillgratrutens(Larusa.argentatusPont.)naringsochspridningsekologi.[Onfood habies anddistributionoftheHerringGull(Larusa.argentatusPont.).]K.Fysiogr.Sallsk.LundForh.18:36-56.[InSwedishwithEnglishsummary.]1948b.Tillgratrutens(Larusa.argentatusPont.)naringsochsprid-----ningsekologi.[Onfoodhabitsand-distributionoftheHerringGull(Larusa.argentatusPont.).]FaunaFlora43:116-137.[InSwedishwithEnglishsummary. ] 287

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HerringGullWhite,M.andC.J.Stevens.1948."Smoke-bathing"ofStarlingandofHerringandBlack-headedGulls.Brit.Birds41:244.Wilmahn, B.1948.Ringmerkningsresultaturavsaeing,LarusargentatusPont.StavengerMus. Arbok57:40-46.[InSwedish.)1947Boswall,J.H.1947.Herring-Gullsnestingonbuildings.Brit.Birds40:255-256.Eaton,R.J.1947.AnoldHerringGull.Bird-Banding18:79.Elliott,H.F.I.andR.E.Moreau.1947.StartofincubationbyHerringGull.Brit.Birds40:286.Jacobson,M.A.1947.AnimpededHerringGull.Auk64:619.Marshall,H.1947.LongevityoftheAmericanHerringGull.Auk64:188-198.McMullen,T.E.1947.NestingofHerringGullinNewJersey.Auk64:321.Meiklejohn,M.F.M.1947.CourtshipofHerringGullonwater.Brit.Birds40:285-286.Paynter,R.A.,Jr.1947a.ThefateofbandedKentIslandHerringGulls.Bird-Banding18:156-170.1947b.Preliminaryreportontheeffectsofeggingontheclutch-----sizeofKentIslandHerringGulls.Annu. Rep. BowdoinSci.Stn.8:11-12.Salomonsen,F.1947.MaagekoloniernepaaHirsholmene.[Thegull-coloniesonHirsholmene.)Dan.Ornithol.Foren.Tidsskr.41:174-186.[InDanishwithEnglishsummary.)Stresemann,E.andN.W.Timofeeff-Ressovsky.1947.ArtentstehungingeographischenFormenkreisen.1.DerFormenkreisLarusargentatus-cachinnansfuscus.BioI.Zentralbl.66:57-76.[In German:rWoodward, B.1947.NotesontheloonsandHerringGullsofthewestMooseheadLakeregion,Maine.Bull.Maine AudubonSoc.3:41-44.1946Bunker,A.1946.Gullstakingfishfrommergansers.Can.Field-Nat.60:115.Crowell,E.M.andS.Crowell.1946.GullsattheWeepecketIslands.ThedisplacementofternsbyHerringBird-Banding17:1-10.288

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HerringGullLindroth,S.1946.Ettbidragtillfraganamgratrutens(LarusargentatusPontopp.)stallningsammarinreliktivarainsjoar.[AcontributiontothequestionoftheHerringGull(LarusargentatusPontopp.)asamarineremnantinourlakes.]VarFagelvarld5:97-114.[InSwedishwithEnglishsummary.]Poor,H.H.1946a.Plumageandsoft-partvariationsintheHerringGull.Auk63:135-151.1946b.BreedingoftheHerringGullinConnecticut.Proc.Linn.Soc.------N.y.54-57:43-44.Voous,K.1946.Suruncasd'hybridationnaturelleentreLarusfuscusetLarusargentatusdanslesPays-Bas.Alauda14: 2l-32.--rrn French.]1945Beaudette,F.R.1945.Aspergillosisandparasitisminagull.Bird-Banding16:99-101.Brown,R.H.1945.HerringGullfeedingindependentyoung.Brit.Birds38:217-218.Ellison,N.F.andW.Wilson.1945.Herring-GullnestinginCheshire.Brit.Birds38:376.Johnston,F.J.1945.Herring-GullsrobbingLapwings.Brit.Birds38:278.Jones,E.1945.HerringGullsnestingatBeaverBay.Flicker17:35.Lovell,H.B.ations.1945.ReactionofAmericanMerganserstoHerringGulldepredWilsonBull.57:202.Poor,H.H.1945.Primariesofanine-year-oldHerringGull.Bird-Banding16:102-103.1944Gross,A.O.1944.HerringGullrecoveries.BowdoinCall.Bull.9.Brunswick,ME.Haartman,L.V.1944.MasfragenEnkritikovenkritiker.Fin.Nat.IV.Leach,E.P.shire.1944._ScandinavianHerring-GullsinScotlandandLincolnBrit.Birds37:159.Lloyd,B.1944.HerringGullsfeedingindependentyoung.Brit.Birds38:39-40.Mason,A.G.1944.HerringGulls'behavior.IrishNat.J.8:1-8.289

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HerringGullPoor,H.H.1944.Color-bandedadultHerringGulls.Bird-Banding15:112114.Rollin,N.1944.Wintermeasurementsofgulls'testes.Ibis86:550-551.White,W.W.1944.Herring-Gullsnestingonahouseroof.Brit.Birds38:120.1943Dexter, R. W.1943.ShellheapsoftheHerringGull.Nautilus57:69.Griffin,D. R. 1943.HomingexperimentswithHerringGullsandCommonTerns.Bird-Banding14:7-33.Hennessey,T. R.1943.BandedHerringGullNo.40-656624.Can.Field-Nat.57:97.Poor,H.H.gion.1943.Color-bandedimmatureHerringGullsintheNewYorkreBird-Banding14:101-115.Ross,W. R. 1943.HormonaldeterminationofadultcharactersandsexualbehaviourinHerringGulls.J.Exp.Zool.94:181-203.Trautman,M.B.1943.HerringGullattackonnormalgolden-eye.WilsonBull.55:192.vanMarIe,J.G.andK.H.Voous,Jr.1943.argentatusenLarusfuscusinNederland.1942GeographischevormenvanLarusLimosa16:146-149.[InDutch.]Behle,W.1942.RecordsoftheHerringGull,SanderlingandLarkBuntinginUtah.Condor 44:230-231.Dathe,H.1942.UberdieRassenzugehorrigkeitderSilbermoweanderKustederDarsshalbinselinHerbst.Ornithol.Monatsber.SO:127.Davis,D. E.1942.NumberofeggslaidbyHerringGulls.Auk59:549-554.Rand, A. L.1942.Laruskumlienianditsallies.Can.Field-Nat.56:123126.Spear,I.M.1942.HerringGullcontrolexperiments.BowdoinSci.Stn.Bull.8:15-31.1941Aldous,C.M.1941.Reportofawholesaledie-offofyoungHerringGulls,HogbackIsland,MooseheadLake,Maine.Bird-Banding12:30-32.290

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Bodenstein,G.andE.Schuz.Ired.FurDeutschland.HerringGull1941.ErstnachweisvonLarusargentatusanteliusOrnithol.Monatsber.49:49-56.Brandt,M.1941.UeberdasBrutvorkommenderSilbermowe(Larusargentatusomissus)unddesSterntauchers(Colymbusstellatus)imOstbaltikum.J.Ornithol.89:257-267.[InGerman.] Brown,M.1941.Gullseatfruitofthecabbagepalmetto.Auk58:579.Hoff,C.C.1941. A caseofcorrelationbetweeninfectionofsnailhostswithCryptocotylelinguaandthehabitsofgulls.J.Parasitol.27:539.Latham, R.1941.HerringGullsasvegetableeaterson LongIsland,N.Y.Oologist58:12.Lippens,L.1941.MoeursdeGoelandargente.Gerfaut31:137.[InFrench.]Larusa.argentatusPontopp.Slijper,H.J.meeuwen.1941.TweewaarnemingeninNederlandvangeelpotigeZilverArdea30:270-271.[InDutch.]VanOordt,G.J.1941.EenZilvermeeuw,LarusargentatusPont.,broedendinhetpolderland.Ardea30:269-270.[InDutch.]Wetmore,A.1941.HerringGullinGuatemala.Auk58:257.1940Bergman,G.,E.Fabricius,andL.vonHaartman.1940.Enpreliminarundersokningoversilltrutens,Larusf.fuscusL.,ochgratrutens,Larusa.argentatusPontopp., Naringsbiolog:, sarskilturjakvardssynpunk-t-.-Ornis Fenn.17:33-41.Berthel,R.M.1940.HerringGullseatgrasshoppers.Flicker12:35.Cooke,M.T.1940.WinterrangeoftheHerringGull.Auk57:250-251.Davis,W.A. andL.S.McClung.1940.AspergillosisinwildHerringGulls.J.Bacteriol.40:321-323.Drost,R.andL.Schilling.1940.UeberderLebensraumdeutscherSilbermowen,Larusa.argentatusPontopp.aufGrund vonBeringungsergebnissen.Vogelzug 11: 1-22.[InGerman.]Gross,A.O.1940.ThemigrationofKentIslandHerringGulls.Bird-Banding11:129-155.Harriot,S.C. andJ.J.Hickey.1940.AChristmascensusofbandedHerringGulls.Proc.Linn.Soc.N.Y.50-51:33-34.291

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HerringGullHelmuth,W.T.1940.TheHerringGullon LongIsland.Univ.StateN.Y.Bull.Schools26:256-257,259. Koch,J.C.1940.GeelpootigeZilvermeeuwen,Larusargentatussubspec.,inNederland.Ardea29:227-230.[InDutch.]Lowe,W.P.1940.HerringGullandsnake.Ibis(14thSer.)4:728.Mayaud,N.1940.ConsiderationssurlesaffinitesetlasystematiquedeLarusfuscusetLarusargentatus.Alauda12:80-98.McKeever, C.K.1940.ThebreedingoftheHerringGull(Larusargentatussmithsonianus)onLongIslandin1939.Proc.Linn.Soc.N.Y.50-51:32-33.Throne,A.L.1940.AfeedinghabitoftheHerringGull.WilsonBull.52:34.Woodcock,A.H.1940.ObservationsonHerringGullsoaring.Auk57:219-224.1939Deusing,M.1939.TheHerringGullsofHatIsland,Wisconsin.WilsonBull.51:170-175.Harrington,R.W.,Jr.1939.ParasitesoftheHerringGullLarusargentatussmithsonianus.BowdoinColI.Bull.6:14-18.Kuhlemann,P.1939.BeobachtungenaneinerdurchFlusseeschwalben(Sternah.hirundoL.)ausvertauschtemEierbrutetenundaufgezogenenSilbermowe(Larus argentatusPontopp.).Z.Tierpsychol.3:75-84.[InGerman.] Latham, R.1939.HerringGullsonploughedlands.Oologist56:77-79.McClanahan, R.C.1939.ProtectingblueberriesfromdamagebyHerringGulls.U.S.Dep.Int.BioI.Surv.Wildl.Leafl.141.4pp.Mendall,H.L.1939.FoodhabitsoftheHerringGullinrelationtofreshwatergamefishesinMaine.WilsonBull.51:223-226.Meylan,O.1939.NotesurIeGoelandargenteLarusfuscus(argentatus)michahellisNaumann.Arch.SuisseOrnithol.1:456-465.Richter,R.1939.WeitereBeobachtungenaneinergemischtenKolonievonLarusfuscusgraellsiBrehm undLarusargentatusPontopp.J.Ornithol.87: [InGerman.]--Salomonsen,F.1939.Oologicalstudiesingulls.I.Egg-producingpowersofLarusargentatusPont.Dan.Ornithol.Foren.Tidsskr.33:113-133.292

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'!erringGull1938Cruickshank,A.D.1938.HerringGullsfollowingtheplow.Auk55:672.Drost,R.1938.GeschlechsunterschiedeamSchnabelderSilbermowe,Larusa.argentatusPont.Ornithol.Monatsber.46:129-131.[InGerman.-]-----Geyr vonSchweppenburg,H.1938.ZurSystematikderfuscus-argentatus-Mowen.J.Ornithol.86:345-365.[InGerman.]Harris,A.T.1938.NestingoftheHerringGullandCommonTern.Oologist55:102-104.Hickey,J.J.and R. P.Allen.1938.Firstsight-recoveriesofmarkedHerringGulls.Bird-Banding9:51-54.Kozlova,E.V.1938.FieldobservationsonthebreedingoftheHerring-Gull(Larusargentatusponticus)ontheCaspianSea.Ibis(14thSer.)2:245-254.Lyon,W.I.1938.AlbinisticHerringGulls.Bird-Banding9:102.Richter, R. 1938.BeobachtungenaneinergemischtenKolonievon Silbermowe(LarusargentatusPont.)und Heringsmowe(LarusfuscusgraellsiiBrehm).J.Ornithol.86:366-373.Shelley,L.O.1938.SomedetailedhistoryofHerringGull37-652822.BirdBanding9:54-55.1937Bodenstein,G.1937.VonderWanderungenderSeemowen(Larushyperboreus,L.marinus,L.argentatus,L.fuscus,und L.canus)inOstlichenOstsee nach den Berlngungsergebnissen:--schr. Phys.-Okon.Ges.Konigsberg69:223-234.Booy,H.L.andN. linbergen. 1937.NieuwefeitenoverdesociologievandeZilvermeeuwen.LevendeNat.41:325-334.Broekhuysen,G.J.,Jr.1937.GedragingenvangeslachtsrijpeennognietgeslachsrijpeZilver-enGroteMantelmeeuwen(LarusargentatusPont.etLarusmarinusL.)buitendebroedtijd.Ardea26:159172.[InDutchwithEnglishsummary.] Dewar,J.M.1937.HerringGullsfeedingonstarfish.Scott.Nat.1937:32.Goethe,F.1937.BeobachtungenundUntersuchungenzurBiologiederSilbermowe(Larusa.argentatusPontopp.)aufderVogelinselMemmertsand.J.Ornithol. 85: 1119.[InGerman.] 293

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HerringGullHolmes,P.F.1937.Gullstakingantsoffsurfaceoflake.Brit.Birds30:261.Lockley,R.M.1937.Black-backedandHerring-Gullsand Ravensfeedingonants.Brit.Birds30:325-326.Lowe,W.P.1937.Cangullsdive?Ibis(14thSer.)1:175.McLeod,J.A.1937.Twonewschistosomidtrematodesfromwater-birds.J.Parasitol.23:456-466.Steinbacher,G.1937.DasWiederkennendesGelegesbeiderSilbermowe(Larus argentatusPont.).Beitr.Fortpfl.Vogel13:23-25.Very,N.T.1937.BehaviorofyoungHerringGulls.Bird-Banding8:77-78.1936Campbell,J.W.1936.SpottedFlycatcher,GreatTitandgullstakingmoths.Brit.Birds30:172.Ivanenko,I.D.1936.Biolohiyaisil'sko-hospodarskeznachen'yachaikyrehotukhy.[ThebiologyandagriculturalimportanceoftheHerringGull.]Zb. Nauk. Pro Azov.Zapov.1936.[InUkrainian.]Mitchell,M.1936.HerringGullsfeedingonlarvaeofmoths.Brit.Birds30:92.Ringleben,H.1936.UberdieNahrungderaltenundjungenSilbermowen.Beitr.Fortpfl.Vogel12:165.[InGerman.]Tinbergen,N.1936.ZurSoziologiederSilbermowe,Larus argentatusPont.Beitr.Fortpfl.Vogel12:89-96.[InGerman.]1935Broekhuysen,G.J.,Jr.1935.GedragingenvannognietgeslachsrijpedochreedszelfstandigeZilver-enGroteMantelmeeuwen(LarusargentatusPont.etLarusmarinusL.).Ardea24:239-250.[InDutchwithGermansummary.]Drost,R.1935.ausgefarbt.Silbermowe,Larusa.argentatusPont.,im5JahrnochnichtOrnithol.Monatsber.43:116.[InGerman.]Grant,C.H.B. and C.W.Mackworth-Praed.1935.[OntheracesoftheHerringGullvisitingeasternAfricainthenon-breedingseason.]Bull.Br.Ornithol.Club56:33-34.Makkink, G.F.1935.BritishLesserBlack-backedGullXHerringGullinTexel.Ardea24:208-209.294

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HerringGullMeinertzhagen,R.1935.TheracesofLarusargentatusandLarusfuscuswithspecialreferencetoHerrB.Stegmann'srecentpaper Ibis(13thSer.)5:762-773.Mendall,H.L.1935.TherelationshipofcertainseabirdstothefishingindustryofthestateofMaine.Bull.Dep.SeaShore.Fish.Maine.28pp.Munro,J.A.1935.NestingcoloniesoftheHerringGullinBritishColumbia.Condor37:214-215.Pearson,T.G.1935.AHerringGullofgreatage.Bird-Lore37:412-413.Ritchie,J.1935.GreatageofHerringGull.Scott.Nat.1935:69-70.Schuz,E.alt.1935.Silbermowen(Larusa.argentatus)fast25und26JahreVogelzug6:134-135.-------1934Eaton,R.J.1934a.ThemigratorymovementsofcertaincoloniesofHerringGullsineasternNorthAmerica.PartII.Bird-Banding5:1-19.1934b.ThemigratorymovementsofcertaincoloniesofHerring-----GullsineasternNorthAmerica.PartIII.Bird-Banding5:70-84.Gabrielson,I.N.1934.TheHerringGullontheColumbiaRiver.Murrelet15:25.Hinchman,R.M.1934.A fewHerringGullrecoveries.Bird-Banding5:189-190.Hytonen,U.1934.Harmaalokin(Larus argentatusPontopp.)pesimisestasoillame.OrnisFenn.11:61-75.[InFinnishwithGermansummary.]Junge,G.C.A.1934.AdifferenceintimebetweentheegglayingofLarusfuscusL.andLarusargentatusPont.intheShetlands.Ardea23:169-171.Saunders,A.A.1934.Acurioushabitofgulls.Auk51:234.Shelley,L.O.1934a.HerringGullrecoveryfromPanama.Bird-Banding5:189.1934b.GullnotesfromtheNewHampshireandMainecoasts.Auk51:83.vanDobben,W.H.1934.Bijdragetothetmeeuwenvraagstuk.Org.ClubNed.Vogelkd.7:63-78.295

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HerringGull1933Allen,R.P.1933.BreedingrangeofHerringGullextended.Auk50:433-434.Eaton,R.J.1933.ThemigratorymovementsofcertaincoloniesofHerringGulls(LarusargentatussmithsonianusCoues)ineasternNorthAmerica.Bird-Banding4:165-176.Lonnberg, E. 1933.SomeremarksonthesystematicstatusoftheYe11owleggedHerring-Gulls.Ibis(13thSer.)3:47-50.Tomkins,I.R.1933. Ring-billed andHerringGullsattheSavannahRivermouthinJulyandAugust.Auk50:103.Schuz, E. 1933.VondenWanderungenderEismeer-undOstsee-Si1bermowen(Larus argentatus).OrnisFenn.10:17-19.1932Austin,O.L.1932d.TheoccurrenceofbotulisminaHerringGull.BirdBanding3:178.Duse,A.1932.Lacoloniadigabbianirea1idel1agodiGarda.Riv.Ita1.Ornito1.2:1-4.Lockley,R.M.1932.Incubation-periodsofLesserandGreatBlack-backedandHerring-Gulls.Brit.Birds25:310-313.Tinbergen,N.1932.WaarnemingenaanZi1vermeeuenindebroed-ko1onieteWassenaar.LevendeNat.37:248-252.1931Marcot,C.1931.Ausujetdesoeufsdu Goe1andargenteapiedsjaunesde1aMediterranee,Larusargentatusmichahe11esiiBruch.etdu Goe1andd'Audouin,Larus audOUinii Payr.OiseauRev.Fr.Ornitho1.1:391-392.Schuz,E.1931.22Jahren.AlteRingvoge1:Si1bermowe(Larus argentatus)von21und Voge1zug2:176-177.Steinbacher,G.1931.BeitragezurBrutbio1ogievon Si1bermoweandBrandseeschwa1be.J.Ornitho1.79:349-353.[InGerman.] van Dobben,W.H.1931.Einze1heitenuberFarbeundBrutbio1ogiederaufTersche11ingnistendenHeringsmowen(Larusfuscussubspec.)Ardea 20:143-147.[InGerman.] 296

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HerringGull1930Hortling,I.1930. En gulfotadgratrutLarusargentatusomissusSushkin?hackandeinS.Finland.Ornis 67.[InSwedishwithGermansummary.]Oldham,C.1930.Theshell-smashinghabitsofgulls.Ibis(12thSer.)6:239-243.vanSomeren,V. D. HerringGulls.1930.CuriouschangesofdietinBlack-headedand Nat.1930:132.von Hedemann,K.1930.Die NahrungderSilbermowe.Ornithol.Monatsschr.55:160.[InGerman.]1929Ruthke,P.1929.VonderSilbermowe.GefiederteWelt52:612-621.[InGerman. ]Sprunt,A.,Jr.1929.TheHerringGull(Larusargentatus)intheNorthCarolinamountains.Auk46:100.Tinbergen,N.1929.AbreedingpairofHerringGull(Larusa.argentatusPont.)XLesserBlack-backedGull(Larusfuscussubspec.).Ardea18:1.1928Brouwer,G.A.1928.Ardea17:72-73.VoorkomenvandwergeierenbijLarusargentatusPont.[InDutch.]Dietrich,F.1928. Die AbanderungenderSilvermoweneier.Beitr.Fortpfl.Vogel4:157-160.Lincoln,F.C.1928a.ThemigrationofyoungNorthAmericanHerringGulls.Auk45:49-59.1928b.HerringGulls:acorrection.Auk45:366.Portie1je,A.F.J.1928.ZurEthologiePsycho1ogiederSi1bermoweLarusargentatusPont.Ardea17:112-149.[InGerman.]Taning,A.V.Island.1928.SolvrnagenLarusargentatusargentatusPontopp.nyforDan.Ornitho1.Foren.Tidsskr.22:91-96.Tweeda1e,V.1928.Herring-Gullnestingonoccupiedhouse.Brit.Birds22:147.297

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HerringGull1927Rapine,J.1927.DifferenciationdesjeunesoiseauxdesespecesLarusargentatusargentatus(Pontopp.)etLarusfuscusaffinis Reinhardt. Rev.Fr.Ornithol.19:96-97.Shipman,C.M.1927.NestingoftheHerringGullandsomeotherbirdsonLakeErieIslands.Auk44:242-243.Skovgaard,P.1927.Maerkende HavmaagerII.[MarkingexperimentswithLarusargentatus.)Dan.Fugle8:108-120.1926Lawson, R. 1926.HerringGullsnestinginSalemBay.Bull.EssexCo.Ornithol.Club1926:52.Lonnberg, E. 1926.Dengulfotadegratrutens,LaruscachinnansPall.,systematiskastallning.FaunaFlora21:218-222.[InSwedishwithEnglishsummary.)1925Mackay,G.H.1925.BreedingoftheHerringGull(Larusargentatus)inMassachusetts.Auk42:517-518.1924Thomson, A.L.1924.ThemigrationsoftheHerringGullandLesserBlackbackedGull.Brit.Birds18:34-44.1923Bean,R.A.1923.AblackHerringGull.Yearb.MilwaukeePublicMus.3:178-181.Moreau,R.E.1923.Herring-Gulleatingitsownchick.Brit.Birds16:221-222.Strong,R.M.1923.FurtherobservationsonthehabitsandbehavioroftheHerringGull.Auk40:609-621.Wolfe,T.R.1923.TheHerringGullsofLakeChamplain.Auk40:621-626.1922Colthrup,C.W.1922.HerringGullshawkingforwingedants.Brit.Birds16:139.298

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HerringGull1921Lonnberg,E.1921. Den gulfotadegratruten,LarusargentatuscachinnansPallas,enforSverigenyfagel.FaunaFlora16:126-130.[InSwedish.]1920Dwight,J.1920.TheplumagesofgullsinrelationtoageasillustratedbytheHerringGull(Larusargentatus)andotherspecies.Auk37:262-268.1918Palmgren,R.1918.LarusargentatusBrunn[male]Xmarinus(L.)[female].He1singforsMedd.Soc.FaunaFloraFenn.44:125-127.1917Leege,o.1917.Die NahrungdieSi1bermowenander ostfrieschen Kuste.Ornitho1.Monatsschr.42:2-24.[InGerman.]1915Nichols,J.T.1915.EstimatedaverageageoftheHerringGull.Condor17:181-183.Strong,R.M.1915.argentatusPont.Strong,R.M.1914.argentatusPont.OnthehabitsandbehavioroftheHerringGull,LarusSmithson.Rep.1914:479-509.1914OnthehabitsandbehavioroftheHerringGull,LarusAuk31:22-49,178-199.1913 Cummings, S.G.1913.HerringGullsdiving.Brit.Birds7:201-202.1912Strong,R.M.1912.Gull.ScienceObservationsonthebreedingbehavioroftheHerring35:936.1906Ward,H.L.1906a.NotesontheHerringGullandtheCaspianTern(LarusargentatusandSternacaspia).Bull.Wis.Nat.Hist.Soc.4:113-134.1906b.WhydoHerringGullskilltheiryoung?Science24:593-594.299

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Herring GullDutcher,W.andW.L.Bailey.1903.Acontributiontothelifehistoryofthe Herring Gull intheUnitedStates.Auk20: 417-431. Rodger,A.M.1903. Hening Gull (I.arus argentatus)capturingabat.Scott.Nat.Hist.Soc.1903: 51. Mackay,G. H.1893.Larusargentatussmithsonianus.Auk10:76.Mackay,G. H.1892.HabitsoftheAmerican Herring Gull(Larusargentatus inNewEngland.Auk9:221-228.AdultHerringGullinwinterplumage.PhotographbyRogerB.Clapp.300

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GREATBLACK-BACKED GULL(Larusmarinus)IDA:Svartbag,DU:Mantelmeeuw,FI:Merilokki, FR: Goelandmarin, GE: Mantelmowe, IC:Svartbakur,IT:Mugnaiaccio,NW:Svartbak,PO:Mewasiodlata,SP:Gavion,SW:Havstrut] GENERAL DISTRIBUTIONNorthAmericaInsoutheasternCanada,GreatBlack-backedGullsbreedalongthecoastofLabrador,insouthcoastalQuebec,inNewfoundland,andintheMaritimeProvinces.Intheinterior,theybreedrarelyandlocallyontheGreatLakesinsouthernOntario(Godfrey1966).IntheUnitedStates,GreatBlack-backedGullsbreedfromthecoastofMainesouthtoNorthCarolina(AOU1957,ParnellandSoots1975,Map7).NorthAmericanbirdswinterregularlyinthebreedingrangesouthtotheGreatLakesandalongtheAtlanticcoasttoNorthCarolina(AOU1957).Thisspecieshasoccurredmoreandmore commonlyinrecentyearsasawinterresidentalongthecoastinstatessouthofNorthCarolinaandhasbeenrecordedasfarwestasTexas.AlthoughsteadilyrecordedingreaternumberseachyearsouthofNorthCarolina,theyarestilluncommonwintervisitorsinthatarea.GreatBlack-backedGullsarelesslikelytowanderfarfromtheirbreedingandwinteringrangesthanaremanyotherNorthAmericangulls,buttheyhavewanderedasfarnorthasBaffinIslandand Hudson Bay(Godfrey1966),andasfareastandsoutheastasBermuda,Aruba(intheNetherlandsAntilles),PuertoRico,Cuba,St.Bartholomew,andBarbados(Vaurie1965,Bond1971,Voous1977,Raffaele1981).WorldDistributionIntheOldWorld,GreatBlack-backedGullsbreedonlyinwesternEurasiabetweenabout420and770N (Voous1960,BOU1971).TheybreedinGreenland,Iceland,theFaeroes,GreatBritain,northwestFrance,northernDenmark,ontheAtlanticandBalticcoastsofNorway, Sweden, andFinland,andinpartofEstonia;theyalsobreedontheMurmanskcoastofRussia,offshoreinSpitsbergen,andonBearIsland(Vaurie1965,BOU1971,Crampetal.1974).Europeanpopulationsarebothsedentaryandmigratory,movingsouthasfarastheMediterraneanandtheBlackandCaspianSeasandwanderingcasuallytoMadeira,theCanaries,andtheAzores(Vaurie1965).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESNorthCarolinaFirstrecordedfromNorthCarolinaon5February1901(Pearsonetal.1942),theGreatBlack-backedGullhasgraduallybecome a commontoabundantwinterresidentalongthecoast(Map8).Theyaremostabun-301

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BIRDNAME: __-r'-'1.---.-"'.....(.8 0 "'-'-\,AlJ...II--'r__.loc--\-0, \ 88" -------,\---\ MO 1L \"-t-_--1-,-t-t JACICSON GULFOFMEXICO """", ).".),F-.J -I sA HOUSTONPORTMTH EXTDALLAS-----BreedingRangeMapforSoutheastern Dnned States 320-77, MALNO.YEAR NumbeA inboxesdenotemaximum estimates of breeding birdsatcoloniesinrecent years. first figure indicates maximum number of birds. Second figure indicatesyearinwhichestimatewasobtained.< .O__ -I------J---r--of/-0/.",/w oNMap7

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30 .,'DfItYT'ORTUGAS GREATBLACKBACKEDGULLBIRDNAME'.. N.---..-------------(--, f \ i < I $fit I I l,... I .J.:I.\ I tit I 1IONTlIO_., "JACKSON,
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GreatBlack-backedGulldantnorthofCapeHatteras(Map8).Thefirstknowninstanceofbreedingwithinthestatewasreported24-25June1972,whenParnellandSoots(1975)foundfouradultsandfourflightlessjuvenilesinacolonyofHerringGullsatOregonInlet.Fouradultswerepresentatthesamesitethefollowingsummer, and anestwiththreeeggswasfound1June1973.Threepairswerefoundnestingonadredge-spoilislandin1976(Portnoyetal.1981)andtenpairsofGreatBlack-backedGullsnestedatfivesitesin1977(ParnellandSoots1979ms).Nineofthenestsfoundin1977wereinRoanoke Sound onfourdredge-spoilislands(Map7)wheretheGreatBlack-backedGullswerealwaysfoundnestinginassociationwithnestingHerringGulls(ParnellandSoots1979ms).SouthCarolinaBurton(1970)consideredthesegullstoberarewintervisitorstoSouthCarolina,butsuggestedthattheywouldbeseenmorefrequentlyinthefuture.Through12December1968therewereonly13sightingsof16birds;mostoftheobservationsweremadeinJanuary(3)and December(6)(SpruntandChamberlain1949,Burton1970).GreatBlack-backedGullsareuncommoninwinterinSouthCarolina,butrecentChristmasCountsindicatethattheyaregraduallybecoming morecommoninnorthernareasofthestate(Map8).Forsythe(1973)recordeda maximumcountof20birdsatCharlestoninFebruaryinaperiodextendingfromJune1971toJune1972.GeorgiaAsrecentlyas1958GreatBlack-backedGullswereknownfromGeorgiaonlyonthebasisofasinglespecimentakeninChatham Countyprobablyabout1910(Burleigh1958).Dentonetal.(1977)considereditanuncommonvisitoralongthecoastfromDecembertoMarch.Anunusuallylargerecentconcentrationconsistedof20to25immaturebirdsseenonandoffshoreJekyllIslandon11February1979 (LeGrand1979b).GreatBlack-backedGullshavebecome morecommoninGeorgiainrecentyears.Florida-AtlanticCoastInFlorida,asinotherstatesalongthesoutheasternseaboard,theGreatBlack-backedGullhasbecome morecommonrecently.Howell(1932)knewitonlyfromtwo19th-centuryrecordsfromthenortheasterncoastofFlorida;oneofthemhadbeenmade by Audubon.Sprunt(1954)knewofanothersixrecords,allfromtheAtlanticcoastandonefromasfarsouthasLower Matecumbe Key.Sprunt(1963)laternotedthattherewererecordsfrommostyearssince1954.Atpresent,smallnumbersofthisspeciesarecommonlyseensouthtoCapeCanaveral,butthisgullisuncommonfromtheresouthtotheKeys(Kale1979msa).Therecentincreaseinthenumberofwinteringbirdsisdocumentedbyhighcountsmadealongthenortheastcoast.AsmanyasfivewereseennearCocoainthewinterof1955-56(Stevenson1956b),butonlythreeyearslater,duringthewinterof1958-59,atleast13werepresentatCanaveral(Stevenson1959a).In1960-61atleast28GreatBlack-backedGullswinteredatCanaveral;thiswasthehighestcountforFloridayetmade(Stevenson1970).Theaveragenumberseenon someofrecentChristmasCounts(1973-77)(Map8)suggeststhatthetotalpopulationwinteringontheAtlanticcoastisstillincreasing.Mostrecordsarefrom7 Novemberto27March; a fewothers(9October;304

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GreatBlack-backedGull18,25April)probablyrepresentearly-arrivingorlate-departingwinterresidents.TherearenowaboutadozenrecordsforMaythroughAugustofsummeringGreatBlack-backedGulls.Almostallrecordsarefromthecoast,butweknowoftworeportsofbirdsseenabout20 mi(32km)inland.Onewasseen25April1963 ontheSt.John'sRivernearJacksonville(Stevenson1963b);theotherwasseenonthesameriveratMandarin,9February1979(Stevenson1979a).Florida-GulfCoastGreatBlack-backedGullsareuncommontorareontheFloridaGulfcoast,butscatteredindividualsareseenwithincreasingfrequencyeachwinter.ThisspecieswasfirstrecordedfromtheGulfcoastin1948 when abandedbirdwasrecoveredatPanamaCityon17February(Imhof1974);weknowofapproximately22recordssincethen.Twotothreeindividualswereseenin1959,atleastsevenwereseeninthe1960's,andabout12-14individualswerereportedduringthe1970's.Thisspecieswaspresentinatleastsevenofthepastninewinters.Allbutsevenoftherecordsknowntousfallbetween13Octoberand29March.Fourothers(20August-28September)mayhavebeenofbirdsthatsum meredontheGulfcoast.Threelatespringrecords(15April-22May)mayhavebeenofverylatespringmigrantsorofbirdsthatremainedinFlorida;weknowofnoreportsforJuneorJuly.Alabama TheGreatBlack-backedGullisanoccasionalwintervisitoralongtheAlabamacoast.Imhof(1976b)listed7recordsfromcoastalareas;weknowofanadditionaltworecords,bothfromwellinland.1956 28 Dec. 195928Dec. 196321Nov. 1963 28Dec.1971 24Apr.1972 30Dec.1974 5 Nov. 197411Nov. 1977Jan.11 12imm.seenatBlakelyIslandimm.seenatCochraneCausewayad.seenatCochraneCausewayseenon DauphinIslandseenonW DauphinIslandseenatFortMorganad.seenatCochraneCauseway imm.seeninlandatWheelerNWRseeninlandnearDecaturImhof 1962b Imhof 1962b Imhof 1976b Imhof 1976b Imhof1971,1976b Imhof 1976b Imhof 1976bHamilton1975Hamilton1977TexasOberholser(1974)listedtheoccurrenceofthisgullinTexasashypotheticalonthebasisoftwosightrecords.Therehavebeenatleastfivesightrecordssincethen,allmadeduringthe1970'swhenthespecieswasextendingitsrangeinothersoutheasternstates.305

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GreatBlack-backedGull1949 21-27 Feb. imm.seenatRockportOberholser1974 1953 23Feb.imm.seenatRockportOberholser1974 1973ca.6 Mar.seenatLagunaAtascosaNWRWebster1973c 197311Nov.ad.seenatLagunaAtascosaNWRWebster1974a 197424Mar.-imm.seen,photogr.atBolivarFlatsWebster1974b,23July,[Thebirdseenini975wasbelieved1974d,1975b 1975 18 Mar.tobethesameoneseenin1974.]1976 4Jan.2seenatFreeportjettiesWebster1976b1978-winterseenatLagunaAtascosaNWRWebster1979b79SYNOPSISOFPRESENTDISTRIBUTIONANDABUNDANCEBreedingTheGreatBlack-backedGullbreedsintheNewWorldfromLabradortoNorthCarolina,becomingprogressivelylesscommonfromnorthtosouth.IntheOldWorld,thespeciesbreedsinGreenlandandIceland,andfromtheFaeroeIslandsandnorthwesternFrancenorthandeastthroughScandinaviatotheMurmanskcoastoftheU.S.S.R.ThebreedingdistributionintheNewWorldisconsiderablymoresoutherlythanintheOldWorld;itextendsfromabout620toabout340N (Voous1960).Thesizeoftheworldpopulationisunknownbutsomerecentinformationisavailableforpartsoftherange.ErwinandKorschgen(1979)indicatedthatabout17,405pairsnestedfromMainetoVirginiain1977.ThisfigurerepresentsallbutaninsignificantproportionofthetotalnumberbreedingwithintheUnitedStates.Incontrast,about22,000pairsbredinGreatBritainandIrelandin1969-70(Crampetal.1974).Anestimated410pairsnestedonthenorthcoastoftheGulfofFinlandin1980(Kilpietal.1980).MostoftheGreatBlack-backedGullsintheUnitedStatesbreedinMaine andMassachusetts,whichheld56.6% and 26.8%,respectively,ofthetotalreportedbyErwinandKorschgen(1979).ThefourlargestcolonieswerethoseatGardiner'sIsland,NewYork(1,200pairs);MonomoyIsland,Massachusetts(900);andSmuttynose(960)and Duck(800)Islands,Maine(Erwin1979a).Togetherthesecoloniesheldapproximately22%ofthebreedingGreatBlack-backedGullsofthenortheasternUnitedStatesin1977.WinterandMigrationExceptforMississippiandLouisiana,theGreatBlackbackedGullhasnowbeenrecordedinallthesoutheasternUnitedStates(Table36),whereitoccursasacommon(inNorthCarolina)toscarceorrare(northernGulfcoast)wintervisitor.AsintheOldWorld,populationsarebothsedentaryandmigratory;mostbirdsmoveonlyshortdistances.306

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GreatBlack-backedGullTable36.DatesofoccurrenceforGreatBlack-backedGullsinthecoastalsoutheasternUnitedStates(a).StateApproximatenumberofoccurrencesDatesofoccurrenceNorthCarolinamanythroughoutyear,winterpeakSouthCarolinauncommon,regular12 Nov. 12MayGeorgiauncommon Dec. Mar.Florida50+7 Nov. -27Mar.(9Oct.;18,25Apr.)-AtlanticcoastFlorida2313Oct.29 Mar.(20Aug. 28Sep.)-Gulfcoast(15 Apr. 22May)Alabama 9 5 Nov. 24Apr.MississippiunrecordedLouisianaunrecordedTexas811Nov. 31 Mar. (summer 1975)(a)Exceptionalorunseasonalrecordsaregiveninparentheses.IntheOldWorld,mostbirdsbreedinginGreatBritainwinterthere;afewmovetonorthwestFranceandIberia.GreatBlack-backedGullsfromNorway,northwesternRussia,andIcelandalsowintertosomeextentinGreatBritain(Crampetal.1974).ThewinteringareasofFenno-ScandinavianGreatBlackbackedGullsaresimilartothoseofHerringGulls.MostBalticbirdswinterinthesouthwesternBalticSea,andDanishandwestSwedishbirdswinterintheSkagerrak,Kattegat,andtheGreatBelt;NorwegianbirdsandthosefromtheMurmanskregionofRussiawinterintheeasternandwesternportionsoftheNorthSea,respectively(Bianki1967).HABITATNestingGreatBlack-backedGullsarepredominantlyresidentsofcoastalareas(Crampetal.1974)andnestmostlyonsmallislands.NorthAmericanbirdshavebeenrecordedonlow,rockyislandsincoastalareasandonislandsinfreshwaterlakes.ThosenestingonislandsinLakeGeorge,NovaScotia,usedavarietyofsites;somenestedonrockypeninsulas,othersinvegetationbackfromtheshore,andyetothersnestedamongbouldersorstumpsnearthewater(Bent1921).ThoseatLittleDuckIsland,Maine,nestedintwolargemeadowswithlarge,exposedgraniteoutcroppingsandvegetatedmostlywithgrasses,stingingnettle(Urticadioica),angelica(Angelicalucida),andrasp-berry(Rubusidaeus)(ButlerandTrivelpiece1981).---BirdsinGreatBritainusuallynestintheinteriorofthe"topsofstacks,smallislands,andholms"(Crampetal.1974).Verbeek(1979)notedthatGreatBlack-backedGullsatWalney,England,nestedneartheedgesofpondscreatedbydredging.Of53nests,51werewithin12.2m(40.0ft)of307

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GreatBlack-backedGulltheponds (meandistance=4.9m[16.1ft]);thetwoexceptionswere39.3and45.7m(129and 150ft)away. Theypreferredtonestonslightlyelevatedgravelpilesandridges(62%, n=50);therestofthenestswereinflatgrassyorsandyareas.AsimilarpreferenceforelevatednestingsituationswasfoundinWales(Harris1964a).Inbothareas,most(87%)ofthesegullsnestonislands,butsomenestonmoors,alonglakesandestuaries,andoncliffsinavarietyofsituations.Lesscommonlyusednestinghabitatsincludeshinglebanksandsaltmarsh(Davis1958).NestingonsaltmarshisalsouncommonintheeasternUnitedStates.Burger(1978)reportedsalt-marshnestinginNewJerseywherethesegullsusuallynestedongrassunderbushes(Ivafrutescens)amidthegreatestdensityofHerringGullnests. --Nestinghabitatsusedelsewherearesimilar(Dement'evand Gladkov1951,Gudmunsson1954,Godfrey1966),althoughthedegreetowhichonehabitatoranotherisusedvariessomewhatfromareatoarea.GreatBlack-backedGullsnestsolitarilyorindiffusecolonies(Harris1964a)buttendtobemoresolitaryintheirnestinghabitsthanmanyotherlarids.ColoniesalongthenortheasternseaboardoftheUnitedStatesaresmall.Anaverageof73.7breedingpairswasfoundin147coloniescensusedfromsouthernMainetoVirginiain1977(ErwinandKorschgen1979).Althougheightnortheasterncoloniescontained1,000ormorebreedingbirds,81.7%of350eastcoastcoloniescontained50pairsorless.FeedingGreatBlack-backedGullsuseawidevarietyofmostlycoastalfeedinghabitats.Theseincludebeaches,openocean,andestuaries.Thegullsoftencongregatewhereoffalorotherrefusecanbereadilyobtained(e.g.,garbagedumps,slaughterhouses,fishingpiers,sewers,fishingtrawlers).Theyalsofeedintidalpoolsandinshallowpartsoflakesandrivers(Gudmundsson1954).Hunt and Hunt(1973)comparedhabitatusedbyforaginggullsinMaineandnorthwesternEuropeandfoundthatsubstratesusedforfeedingbyGreatBlackbackedGullsweresimilarinbothareas.MoreofthesegullswereseenfeedingondumpsinbothMaine(45.2%,n=735)andnorthwestEurope(32.9%,n=137)thaninanyotherhabitat.Areasinwhichwastesweredischargedfromsewersandfish-processingplantswerealsofrequentlyused(25.9% and 32.1%,respectively).TheHuntsthoughtthattheintertidalzonewasunimportantasfeedinghabitatforthisspeciesasfewindividualsfedthere.However,Ingolffsson's(1976)analysisoftheoriginoffoodseatenbythesegullsinwesternIcelandsuggestedthatGreatBlack-backedGullstakemuchfoodfromthesurfaceoftheseaaswellasfromtheintertidalzoneandfromrefusedumps.BirdsbreedingatWalneyIsland,England,feedfrequentlyintheintertidalzone,butalsofeed(bypredationandkleptoparasitism)withinthecolony,andonagarbagedump(Verbeek1979).NonbreedingandOffshoreNonbreedingGreatBlack-backedGullsaremostlyfoundalongshorelines;theyarefoundinshoremoreoftenandarelesspelagicthanmostotherspeciesofLarus(Dement'evand Gladkov1951,Watson1966).Theyhavebeenseenfollowingfishingvesselsasfaroffshoreas12or15mi 308

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GreatBlack-backedGull(19-24km)inEngland(Davis1958),and LeeandBooth(1979)reportedthatGreatBlack-backedGulls(mostlyimmatures)occurredregularly20mi(32km)ormoreoffNorthCarolina.Rowlett(1980)foundonlyimmaturesoffshoreduringthesummerintheChesapeakeBightnorthofNorthCarolina.ObservationsmadeoffthecoastsofNovaScotia(Brown1967a),Newfoundland (Brown1968,NettleshipandTull1970),andGreenland(Brown 1968)alsoindicatethatGreatBlack-backedGullsremainclosetothecoasts,butmaymoveoffshoretofeed.Thesegullsareonlyoccasionallyreportedwelloutatsea.Sage(1968)sawthreebirdsabout70mi(110km)fromthewestcoastofIrelandandanotherindividual550mi(885km)westofSaintKilda;RankinandDuffy(1948)foundthemcommontoabout100 mi(160km)offCapeSable,NovaScotia,andat100150 mi(160-240km)offthecoastofNorway.FOODANDFEEDINGBEHAVIORGreatBlack-backedGullsfeedopportunistically.TheywereconsideredtheleastspecializedfeedersoffivespeciesofLarusstudiedinIceland(Ingolfsson1969a)andobtaintheirfoodby awidevarietyoftechniques.Wheninmarinesituationstheysurface-seize,sittingonthewaterandimmersingheadandnecktoseizebothliveanddeadfood.GreatBlack-backedGullsalsosurfaceplungebyhoveringoverthewateranddroppingtoseizefoodatorlessthanafootbelowthesurface(Ingolfsson1969a).Suchsurface-plungingusuallyisperformedfromheightsofonlya fewfeet,butSeymour(1974)witnessedthesegullsdivingoneels(Anguillarostrata)fromheightsofSOto100ft(15-30m);thegullspartiallyfoldedtheirwings,dropped5to10ft(1.5-3m)abovethesurface,broketheirdivewiththeirwings,andplungedintothewaterwithoutentirelysubmerging.GreatBlack-backedGullsalsokleptoparasitizeotherbirds.Kock(1974)sawthesegullsstealingfishfromCommonMurres(Uriaaalge)atHelgolandintheNorthSea.TheyalsostealmusselsfromCommonEiders(Somateriamollissima)(Roberts1934,Ingolfsson1969a)andTuftedDucks(Aythyafuligula)(HarrissonandHurrell1933),butapparentlydosolessfrequentlythansomeotherlaridssuch HerringGulls(Ingolfsson1969a,Prys-Jones1973).Occasionally,kleptoparasitismistheprimarywayinwhichfoodisobtained.GreatBlack-backedGullsatWalneyIsland,England,obtainmostoftheirfoodfromothergulls,mostlyHerringGulls,andtoalesserextentLesserBlackbackedGulls(Larusfuscus)(Verbeek1979).VerbeeksuggestedthatallfoodeatenbyGreatBlack-backedGullsatalocalgarbagedumpwasobtainedthroughkleptoparasitism.OtherinstancesofkleptoparasitismbyGreatBlack-backedGullsincludeabirdthatrobbedaPeregrineFalcon(Falcoperegrinus)ofaCommonTeal(Anascrecca)(Tinbergen1953),othersthatstolepreyfromherons(Ardea [HarrisonandHurrell1933,Carlsson1979,Forsberg1979] and--A.herodias[Quinneyetal.1981]),and onethatevenstoleafish(Pollachiusvirens)fromthemouthofashark(probablyLamnanasus[French1982]).AttemptedkleptoparasitismofOspreys(Pandion haltaetUs) hasalsobeenreported(Leck1973).309

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GreatBlack-backedGullFoodsobtainedthroughkleptoparasitismvarywidelyandmayincludeshellfish,crustacea,birds,orotheritems.Tolonen(1976)identified20itemsstolenfromHerringGullsinRhodeIsland,Connecticut,andNewJerseyduringastudyconducted1968-72.Halfwereflatfishandmostoftherest(8)werecrabs(Callinectes,Cancer,Carcinus,Libinia).Gullsreadilyscavengefood,andsuchfoodmayformamajorpartofthediet.Theextenttowhichthisspeciesscavengesfromfishingvesselsvariesconsiderably.Kock 1974(inVerbeek1979)reportedthatGreatBlack-backedGullsintheNorth Sea atHelgolandfeedbehindfishingvesselstoalargeextent;incontrast,Verbeek(1979)didnotseeanyofthesegullsaroundtheboatsduringatwo-daysojournintheIrishSea.OnseveraloccasionsGreatBlack-backedGullshavebeenseenfoot-paddling(King1976),abehaviorinwhichgullstramponthesubstrate(Buckley1966) andthatmaybringearthwormstothesurface.Thescarcityofrecords,however,suggeststhatGreatBlack-backedGullsfoot-paddleconsiderablylessoftenthandootherlaridssuchastheHerringGull.Likeseveralotherspeciesofgulls,GreatBlack-backedGullsdropobjectstoshatterthemforeasierconsumption.They dothisratherrarely(Ingolfsson1967),buthavebeenrecordeddroppingDovekies(AIleaIle)onahard-topparkinglotinMassachusetts(Snyder1960);anorthernquahog(Mercenariamercenaria)onanice-coveredpondinConnecticut(Tolonen1976);aratinafieldinSussex(HarberandJohns1947);andeggsofthePink-footedGoose(Anserbrachyrhynchus)inIceland(FisherinTinbergen1953).-----Verbeek(1979)mentionedthattheforagingtimesofGreatBlack-backedGullsatWalneyIsland,England,wereaffectedbythetides;priortolaying,fewornobirdsremainedwithinthecolonyduringlowtide.Differencesindietinrelationtotheageofthebirdhavebeennotedinseveralinstances.AdultsandunfledgedyounginnorthernScotlandfedmoreoftenonsandeels,andimmaturestookmore scavepged fishoffal(Beaman1978).AdultsinConnecticutatemoreflatfishthandidimmatures(Tolonen1976);Tolonenattributedthisdifferencetomoreefficienthandlingofpreybyadults.Belopol'skii(1957)indicatedthatyoungGreatBlack-backedGullsfeedtoagreaterextentonfishandmolluscsthandoadults;adultseatmoreberriesandbirdsandtheireggs. Size ofthebreedingcolonyandseasonmayalsohelpdeterminethedietoftheGreatBlack-backedGull.BirdsnestingsolitarilyinnorthernScotlandfed more frequentlyonotherseabirdsthandidthosenestingcolonially(Beaman.1978).Belopol'skii(1957)reportedthattheamountofmolluscseatenintheBarentsSeaareadecreasedtowardstheendofsummer, andthattheproportionofbirdstakenincreasedinmid-summer whenfledgingyoungofotherspeciesbecame moreavailable.GreatBlack-backedGullssometimeseatothermarinebirds,andattimes310

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GreatBlack-backedGullthedepredationshavebeensogreatthatcontrolmeasureswereinstituted(Harris1964a).TheyhavebeenaconsiderablesourceofmortalitytoManxShearwaters(Puffinuspuffinus),AtlanticPuffins(Fraterculaarctica),andyoungHerringandLesserBlackbackedGullson SkomerIsland,Wales(Harris1965).ManxShearwaterswereamongthemostfrequentlyfound(20.1%)fooditemsin174stomachsofadultGreatBlack-backedGullscollectedatSkomerandSkokholmIslands(Harris1965).AtSt.Kilda,Scotland,some35-40pairsofGreatBlackbackedGullskillabout2,700AtlanticPuffinsfromacolonyofabout40,000pairs,butatanotherlocality(NorthRona)onlyabout400ofapproximately8,000pairsaretakenbysome1,800pairsofgulls(Beaman1978).Eggs,chicks,andadultsofthepuffinarealsotakeninIceland(Ingolfsson1967),butpredationonthelatterislight;mostadultstakenarebirdsthatarealreadydeadordisabled.GreatBlack-backedGullsoftenfeedonducklingsintheGullkronaareaofFinland;19of34samplescontainedtheremainsofyoungbirds(Lemmetyinen1963).Bourget(1973)consideredpredationbythisspeciesandtheHerringGulltobethemajorcauseofegglossfornestingCommonEiders(Somateriamollissima)inPenobscotBay,Maine.ThesegullsalsofeedontheeggsandyoungoftheeiderinIceland(Ingolfsson1967)andelsewhere.Gross(1935)believedthatpredationbythisspecieswasthecauseofa markeddeclineinthenumberofLeach'sStorm-Petrels(Oceanodromaleucorhoa)breedingintheBayofFundy.GreatBlack-backedGullspreyonwidevarietyofotherbirds.Apartiallistofthosekilledandeatenisasfollows:StormPetrel(Hydrobatespelagicus)(Seton1931,HarrissonandHurrell1933);Shag(Phalacrocorax aristoteIIs) (Harris1965);BlackDuck (Anasrubripes)(Addy1945);EuropeanWigeon (Anaspenelope)(Halliday1977);CommonGoldeneye(Bucephalaclangula)(Cleghorn1942);aninjuredRuddy Duck(Oxyurajamaicensis)(Cobb1957);Red-breastedMerganser(Mergusserrator)(Kuerzi1937);AmericanCoot(Fulicaamericana)(Armistead1975);Coot(Fulicaatra)(Thomsen andMeltofte1980);Oystercatcher(Haemato ostralegus)[woundedbyGyrfalcon(Falcorusticolus)](Ingolfsson1976);SandwichTern(Tinbergen1953);Black-leggedKittiwake(Rissatridactyla)(Harris1965,Beaman1978);Razorbill(Alcatorda)(Robinson1923,Harris1965);CarrionCrow(Corvuscorone)(Harris1965);andStarling(Sturnusvulgaris)(Verbeek Black-backedGullshavealsobeenseenattackingandkillingHornedGrebes(Podicepsauritus);LesserScaup(Aythyaaffinis)(Mansueti1961);GreaterScaup(Aythyamarila)(AnderssonandFridzen1970);andBlackbirds(Turdusmerula)(Gamble1959,Karmborg1979).Theyalsotaketheeggsofmanyspecies,among themNorthernFulmar(Fulmarusglacialis)(Ingolfsson1967),NorthernGannet(Sulabassanus);HerringGull;Razorbill;Shag;GreatCormorant(Phalacrocoraxcarbo)(Davis1958);andDouble-crestedCormorant(Phalacrocoraxauritus)(Bourget1973).Eggsareapparentlynotasimportantinthedietasareyoungbirds(Davis1958).MuchhasbeenwrittenaboutthefoodhabitsoftheGreatBlack-backedGull,butlittlespecificinformationis fortheeasternUnitedStates.WesummarizeNorthAmericaninformationbyareabelow.311

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GreatBlack-backedGullNewfoundlandThrelfall(1968a)examined32stomachscollectedinJune-August1966andinMay-September 1967.Fishwerefoundinthemoststomachs(34.0%);vegetation(20.8%),miscellaneousmaterial(17.0%),birdremains(5.7%),molluscs(3.8%),seedandberries(3.8%),echinoderms(1.9%),annelids(1.9%),andinsects(1.9%)werefoundindescendingorderoffrequency.Littleofthismaterialwasidentifiedtospecies,butThrelfall(1968a)notedthatSalmon (Salmosalar)hadbeeneaten,ashadCommonMurres(Uriaaalge)thathadbeencaughtingillnets.TheonlyotherfooditemsidentifiedtospeciesweretwoAtlanticPuffinchicksandanAtlantictomcod(Microgadustomcod).MaineObservationsfromablindatLittleDuckIslandin1979revealedthattheyoungarefedon avarietyoffishandcrustaceans,goose-neckedbarnacles(Lepassp.),squid(Loligoborealis),ediblemussels(Mytilusedulis),andthechicksofHerringGullsandCommonEiders(ButlerandTrivelpiece1981).StudiesoffoodhabitsmadeintheOld WorldincludereportsfromWales(Harris1965),England(Verbeek1979),Scotland(Seton1931,HarrissonandHurrell1933,Beaman1978),West Germany (Kock1974)andtheU.S.S.R.(Belopol'skii1957).AdditionalinformationonspecificfoodseatenbyGreatBlack-backedGullsmaybefoundinthesepapersandinotherslistedinthespeciesbibliography.GeneraorspeciesoffisheatenatvariousOld Worldlocalitiesincludecapelin(Mallotusvillosus)(1ngolfsson1976),sandlance(Ammodytesspp.)(1ngolfsson1967,Kock1974,Beaman1978),Atlanticherring(Clupeaharengus)(1ngolfsson1967),cod(Gadusmorhua),poorcod(Trisopterus minUtUS) (Kock1974),Norwaypout (Trisopterus-esmarkIi), haddock(Melanogrammusaeglefinus)(Beaman1978),whiting(Merlangusmerlangus)(Kock1974,Beaman1978),dab(Limandalimanda),seascorpions(Cottusscorpius),lumpfish(Cyclopteruslumpus)(1n-golfsson1967),andeels (AngUilla anguilla)(Kock1974).------Othervertebrateseatenincluderabbits(Oryctolaguscuniculus),voles(Aplodemussylvaticus),andamphibians(Harris1965).Crabs(Hyasareneus[1ngolfsson1967],Carcinusmaenus[Harris1965,Kock1974], Crangon-crangon [Kock1974])andmolluscs [Harris1965,1ngolfsson1967,Kock1974],Modiolusmodiolus[1ngolfsson1967],Helixnemoralis,Arenicolasp.,Patellaspp.[Harris1965],Cardiumedule, BuCCillUm undulatum[Kock1974])areofteneatenandaresometimes important inthe dieC:-Otherinvertebrateseatenincludeinsects(Belopol'skii1957,Harris1965,1ngolfsson1967),polychaetes(1ngolfson1967),oligochaetes(Harris1965),echinoderms(Belopol'skii1957,1ngolfsson1967),balanidandlepadidbarnacles(1ngolfsson1967),cirripedes,lamellibranchs(Harris1965),andland-snails(Verbeek1979).Plantsaresometimesconsumed,particularlyberriessuchascrowberry(Empetrumnigrum)andblueberries(Vacciniumuliginosum)(1ngolfsson1967).BerriesareanimportantfoodintheU.S.S.R.,amountingto9.2%ofallfoodseaten(Belopol'skii1957).312

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GreatBlack-backedGullSUSCEPTIBILITYTOOILPOLLUTIONOiledGreatBlack-backedGullshavebeenreportedfrequentlyinboththeOldandNewWorlds(Table37),butinonlyoneinstancehasalargenumberofbirdsbeenaffectedadversely.ThisoccurredwhenfloatingoilcontaminatedgullsroostingatnightintheMedwayEstuary(HarrisonandBuck 1967inVermeerandVermeer1974).LikeotherspeciesofLarus,thesegullsusuallyareunaffectedbyoilingincidents.Corkhill(1973)reportedthreeinstancesinwhichGreatBlack-backedGullssurvivedoiling.Intwooftheseinstances,however,nosuccessfulreproductionoccurredduringthefollowingbreedingseason,inoneinstancebecausenoattemptwas madetobreed,andintheotherbecausetheeggsfailedtohatch.Corkhillsuggestedthatthehatchingfailurewascausedbyoilingofthe eggs thatcouldhaveoccurredwhentheoiledplumageoftheadultcameincontactwiththeeggs.SeveralrecentexperimentalstudiesexploredtheeffectsofoilontheeggsoftheGreatBlack-backedGull.Coonetal.(1979)foundthatembryonicdeathinafieldsituationinMaine wassignificantlyhigherineggsthathadbeentreatedwith20microlitersofNo.2fueloil.Theseauthorsfoundnodifferenceinhatchingweightsbetweentreatedanduntreatedeggsincubatedinthelaboratory.Ontheotherhand,treatedeggshatchedsignificantlyfeweryoungthantheuntreatedones.Of64eggsineachtreatedsample,71.95%ofthecontrolshatched;43.8%oftheeggstreatedwith5microlitersofoilhatched;and20.3%oftheeggstreatedwith20microlitersofoilhatched.McGillandRichmond(1979)usedadosageof20microlitersofNo.2fueloiloneggsfromothercoloniesinthesameareaandfoundasignificantreductioninhatchingsuccessamongtreatedeggs;78%of83controleggshatchedincontrastto24%of84treatedeggs.Bybackdatingfromanassumedincubationperiodof29days,McGillandRichmondwereabletostatethatalloil-treatedeggsthatsurvivedwereatleast7-9daysoldwhenexposedtooil.Theypointedoutthatoilcontaminationofeggsmaybea moreseriousthreatthanoilingofadults;theyalsocommentedthatoilcontaminationwouldpresentagreaterhazardtoreproductionearlyinthebreedingseason.TheonlyareainthesoutheastwherethisspeciesoccursinanyabundanceisalongtheNorthCarolinacoastinwinter.Consequently,theeffectofoilingonthisspeciesinthesoutheasternUnitedStatesisprobablyminimal.GreatBlack-backedGullsarestillexpandingtheirrangesouthwardanditisconceivablethatthisgullmayonedaybreedinnumberssignificantenoughtowarrantmoreconcern.BIBLIOGRAPHY1982Bergman,G.1982.Populationdynamics,colonyformationandcompetitioninLarusargentatus,fuscusandmarinusinthearchipelagoofFinland.Ann.Zool.Fenn.19:143-164.313

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GreatBlack-backedGullTable37.Numberofoiledbirdsand numberandpercentageofdeadGreatBlack-backedGullsfoundaftermajoroilingincidents.Jan.1966 805 15Sept.19662,748(b)927Jan.-Feb.10,992(a,b)411970Feb.-Apr.1,276(b,c)1970 Mar. 19724,759(b)Dec. 19729,151(b)2AreaIslandBeach,NewJerseyNortheastEnglandMedwayEstuary,Kent,EnglandNortheastBritainOffeasternCanadaNorth-centralKattegat,Denmark Waddensea, DenmarkDatesJan.1945 Numberofoiledbirds92(a)NumberofdeadGreatBlackbackedGulls4PercentageofGreatBlackbackedGullsSource4.35Kramer and Kramer 19451.86Parrack196733.73Bourne 1968a0.37Greenwoodetal.19710.08Brownetal.19730.02Joensenand Hansen 19770.02Joensenand Hansen 1977(a)Totalincludessomebirdsthatwerenotoiled.(b)Totalincludesonlythosebirdsidentifiedtospecies.(c)Totalincludesbothliveanddeadoiledbirds.Butler,R.G.andS.Janes-Butler.1982.TerritorialityandbehavioralcorrelatesofreproductivesuccessofGreatBlack-backedGulls.Auk99:58-66.French,T.W.1982.Piracyby aGreatBlack-backedGullonashark.WilsonBull.94:96.Gotmark,F.1982.ColonialityinfiveLarusgulls:acomparativestudy.OrnisScand.13:211-224.Hudson, A.1982.GreatBlack-backedGullsonGreatSalteeIsland,1980.IrishBirds2:167-175.314

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GreatBlack-backedGullMudge,G.P.andP.N.Ferns.1982.Thefeedingecologyoffivespeciesofgulls(Aves:Larini)intheinnerBristolChannel.J.Zool.(Lond.)197:497-510.Parsons,K.C.1982.Nest-sitehabitatandhatchingsuccessofgulls.ColonialWaterbirds5:131-138.1981Andrews,R.1981.ThegullpopulationonMonomoyIsland,Massachusetts,USA.(Abstractonly.)ColonialWaterbirds4:194.Burger,J.andM.Gochfeld.1981c.Discriminationofthethreatofdirectvs.tangentialapproachtothenestbyincubatingHerringLarusargentatusandGreatBlack-backedGullsLarusmarinus.J.CompoPhysiol.Psychol.95:676-684.Butler,R.G.andW.Trive1piece.1981.Nestspacing,reproductivesuccess,andbehavioroftheGreatBlack-backedGull(Larusmarinus).Auk98:99-107.Duncan,R.A.1981.TheGreatBlack-backedGull:aGulfcoaststatusreview.Am.Birds35:233-234.Heer,P.de.1981.OnGreatBlack-backedGullwithorangelegs.DutchBirding3:9.McGill-Harelstad,P.1981.TheeffectsofinterspecificcompetitiononreproductivesuccessofGreatBlack-backedandHerringGulls.(Abstractonly.)ColonialWaterbirds4:197.Portnoy,J.W.andM.A. Soukup.1981.Gulluseofafreshwaterkettleholepond:quantificationandseasonalvariation.(Abstractonly.)ColonialWaterbirds4:200.Quinney,T.E.,B.N.MillerandK.R.S.Quinney.1981.GullsrobbingpreyfromGreatBlueHerons(Ardeaherodias).Can.Field-Nat.95:205-206.Raffaele,H.A.1981.theWestIndies.NewrecordsofbirdspeciesforPuertoRicoandoneforAm.Birds35:142-143.1980Burger,J.,M.Fitch,G.ShugartandW.Werther.1980.PiracyinLarusgullsatadumpinNewJersey.Proc.1979Conf.ColonialWaterbirdGroup3:87-98.Grant.P.J.1980.FieldidentificationofWestPalearcticgulls.Part3.Audouin's,Herring,LesserBlack-backed,GreatBlack-backedandGreatBlack-headedGulls.Brit.Birds73:113-158.315

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GreatBlack-backedGullHjernquist,B.1980.Hackningsframgangforkolni-hackendehavstrutarLarusmarinus.Blacku6:18-28.Kilpi,M., H.Puntiiand T.Toivenen.1980.northerncoastoftheGulfofFinland.EnglishwithFinnishsummary.]NumbersofgullsnestingontheDrnisFenn.57:153-160.[InMiller,J.C.1980.IncreasednestingofGreatBlack-backedGullsinNewJersey.Cassinia58:22.Parkes,K.C.1980.GreatBlack-backedGulltowingfooditemashore.Brit.Birds73:586-587.Plotz,J.1980.ParasitosendurchNematodenbefallbeiSilbermowen(Larusargentatus)und Mantelmowen(Larusmarinus)vonderInselHelgoland.[ParasitosisthroughnematodeinfectioninHerringGulls(Larusargentatus)andGreatBlack-backedGulls(Larusmarinus)fromHelgoland.]Angew.Ornithol.4:195-200.[InGermanwithEnglishsummary.]Pulliainen,E. and A.Marjakangas.1980.EggshellthicknessinelevenseaandshorebirdspeciesoftheBothnianBay.OrnisFenn.57:65-70.Southern,W.E.1980a.ComparativedistributionandorientationofNorthAmericangulls.Pp.449-498inJ.Burger,B.C. 011 a andH.E.Winn(eds.).BehaviorofMarine Vol.4:MarineBirds.PlenumPress,N.Y.xviiand 515pp.Thomsen,J.B. andH.Meltofte.1980.SvartbageLarusmarinusdraeberogaederBlishons [GreatBlack-backedGullsLarusmarinuskillingandeatingCootsFulicaatra.]Dan.Ornithol.Foren.Tidsskr.74:147154.[InDanishwithEnglishsummary.]vanKreuningen,J.1980.GreatBlack-backedGullLarusmarinuswithorangelegs.DutchBirding2:14.Whilde,A.1980.The numbersofgullsbreedinginthewestofIreland.IrishNat.J.19:328.1979Angehrn,P.A.M.,H.BlokpoelandP.Courtney.1979.AreviewofthestatusoftheGreatBlack-backedGullintheGreatLakesarea.Ont.FieldBioI.33:27-34.Carlsson,L.1979.izingheron.1Trutarsomnaringsparasitpahagar.[GullskleptoparasitVarFagelvarld38:50.[InSwedishwithEnglishsummary.]316

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GreatBlack-backedGullCooke, A.S.1979.EggshellcharacteristicsofGannetsSulabassana,ShagsPhalacrocoraxaristotelis,andGreatBlack-backedGullsLarusmarinusexposedtoDDEandotherenvironmentalpollutants.Environ.Pollut.19:47 Coon,N.C.,p.H.Albersand R. C.Szaro.1979.No.2fueloildecreasesembryonicsurvivalofGreatBlack-backedGulls.Bull.Environ.Contam.Toxicol.21:152-156.Forsberg, B. 1979.Havstrutsomnaringsparasitpahagar.[GreatBlack-backedGullLarusmarinuskleptoparasitizingheron,Ardeacinerea.]VarFagel [InSwedishwithEnglishsummary.]Foxall,R. A.1979.PresumedhybridsoftheHerringGullandtheGreatBlackbackedGull.Am.Birds33:838.Kallander,H.1979.Labbenpatrutenochtrutenpalabbenellerhundenpakattenochkattenpahundendel2.[SecondarykleptoparasitismbygullsonArcticSkuas.]Anser18:299.[InSwedishwithEnglishsummary.]Karmborg,T.1979.Trutardodarkoltrast.[HerringGullandGreatBlackbackedGullkillingBlackbird.]VarFagelvarld38:50.[InSwedishwithEnglishsummary.]King,B.1979.Roof-nestingbyGreatBlack-backedGulls.Brit.Birds72:340-343.McGill,P.A. andM.E.Richmond.1979.backedGulleggstreatedwithoil.HatchingsuccessofGreatBlackBird-Banding50:108-113.Nunnally,S.,D.Nunnally,R.Needham andR.Lennon.1979.Nocturnalfeedingofgullsatalightedpier.Chat43:63.Pierotti,R.1979.mixedcolony.ThebehaviorofHerringandGreatBlack-backedGullsina(Abstractonly.)Pac.SeabirdGroupBull.6:46.Szaro,R.C.,N.C. Coon andE.Kolbe.1979.PesticideandPCBofCommonEider,HerringGull,andGreatBlack-backedGulleggs.Bull.Environ.Contam.Toxicol.22:394-399.Verbeek,N.A.M.1979.SomeaspectsofthebreedingbiologyandbehavioroftheGreatBlack-backedGull.WilsonBull.91:575-582.1978 Beaman,M.A.S.1978.ThefeedingandpopulationecologyoftheGreatBlackbackedGullinnorthernScotland.Ibis120:126-127.Burger,J.sey.1978.GreatBlack-backedGullsbreedinginsaltmarshinNewJerWilsonBull.90:304-305.317

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GreatBlack-backedGullFisk, E. J.1978c.Thegrowinguseofroofsbynestingbirds.Bird-Banding49:134-141.Fritze, E. 1978. Kingmaerkning af Solvrnage (Larusargentatus)og Svartbag (Larusmarinus)pakobenhavnsKommunaleLosseplads,Amager i1970-71.Feltornithologen20:7-9.Johansen,o.1978.[DDE, PCB andHgineggsofLarusmarinusandLarusargentatusfromcoloniesinMoreandRomsdal.] Stavenger Mus. Arbok1978:67 NorwegianwithEnglishsummary.]Moller,A.P.1978.MagernesLarinaeyngleudbredelse,bestandsstorelseog aendringeri Danmark,medsupplerendeomforholdeneidetovrigeEuropa.[Distribution,populationsizeandchangesingullsLarinaebreedinginDenmark,withareviewofthesituationinotherpartsofEurope.]Dan.Ornithol.Foren.Tidsskr.72:15-39.Tatarincova,I.P.andF.N.Shklyarevich.1978.[TheheadlengthasadiagnosticfeatureforsexdeterminationintheLaridae.]Zool.Zh.57:1446-1447.[InRussianwithEnglishsummary.]1977 Grimm,P.1977.BeitragzumNahrungserwerbderMantelmowe,Larusmarinus.Beitr.Vogelkd.23:63-63.Halliday,K.C.R.1977.GreatBlack-backedGullkillingWigeon.Scott.Birds9:348.Houde,P.1977a.LowproductivityofternsonHicksIsland,1975.Proc.Linn.Soc.N.Y.73:49-57.Ljunggren,L.175-181.1977.HavstrutensokersigtillOlandsbron.FaunaFlora72:[InSwedish.]McGill,P.A.1977.BreedingecologyandcompetitionbetweenGreatBlackbackedandHerringGulls. M.S. thesis,CornellUniv./lthaca,NY.Monaghan,P.andJ. C. Coulson.1977.Statusoflargegullsnestingonbuildings.BirdStudy24:89-104.Spikkeland,I.1977. Svartbak, Larusmarinus,vedinnsjoerpaSorlandetforekomstogbiologi.[OccurrenceandbiologyoftheGreatBlack-backedGull,Larusmarinus,infreshwaterhabitatsinsouthernNorway.]Fauna(Oslo)30:75-87.[InNorwegianwithEnglishsummary.]Stamm, A.L.1977.GreatBlack-backedGullinKentucky.KentuckyWarbler53:18.Voous, K.H.1977.NortherngullsinAruba,NetherlandsAntilles.Ardea65:80-82.318

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GreatBlack-backedGull1976deRidder,M.1976. Mantelmeeuw -Larusmarinus.Wielewaal42:161.Flegg,J.J.M.and R. A. Morgan.1976.MortalityinBritishgulls.RingingMigr.1:65-74.Fleige,K.D.1976.MantelmoweschlagtBlessralle.Falke23:175.Fritze,E.1976.RingingofHerringGullsLarusargentatusandGreaterBlackbackedGullsLarusmarinuson Copenhagen dump. Ring8:22-24.Ingolfsson,A.1976.ThefeedinghabitsofGreatBlack-backedGulls,Larusmarinus,andGlaucousGulls,L.hyperboreus,inIceland.Acta Nat:-ISf. 24.19pp.King,B.1976.Furtherrecordsoffoot-paddlingbygullsongrassland.Brit.Birds69:180-181.Stromar,L.1976.TheecologicalaspectandconsequencesofbandingtheBlackheadedGull--inYugoslavia.Ring8:19-22.Tolonen,K.E.1976.BehavioralecologyofLarusargentatusandLarusmarinus:agespecificdifferentialinfeedingefficiency,aprobablefactorintheevolutionofdelayedbreeding.Ph.D.thesis,YaleUniv./New Haven,CT.275pp.Wendland,V.1976.Mantelmowe(Larusmarinus)totetBlessralle(Fulicaatra).Vogelwelt97:150-151.1975Armistead,H.T. 1975.BreedingofGreaterBlack-backedGull,HerringGull,andGadwallatSmithIsland,Maryland.Md.Birdlife31:131-134.King,B. 1975.Gullbehaviourduringsea-mistintheBristolChannel.BristolOrnithol.8:111-112.Kury,C.R. andM.Gochfeld.1975.Humaninterferenceandgullpredationincormorantcolonies.BioI.Conserv.8:23-34.Lien,J.1975.AggressionbetweenGreatBlack-backedGullsandBaldEagles.Auk92:584-585.Parnell,J.F.and R. F.Soots.nestinginNorthCarolina.1975b.HerringandGreatBlack-backedGullsAuk92:154-157.319

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GreatBlack-backedGull1974Barth, E. K.1974a.SvartbakdruknetGramake. drowns aHerringGull.]Fauna27:95-96.summary. ][GreatBlack-backedGull[InNorwegianwithEnglishEggers,J.1974.Vorkommen undHerkunftderLechmowe(Larusridibundus)im HamburgerRaumimVergleichzurSturm-,Silber-,und Mantelmowe(Laruscanus,L.argentatus,L.marinus).Hamburg.AvifaunaBeitr.12:95-144.[In Ger;an withEnglishsummary.)Firth, R., Jr.1974.Experimentalcross-fosteringofHerringGullandGreatBlack-backedGullchicks.Auk91:139-144.Gochfeld,M.1974c.Gullpredationonmigrantpasserinebirds.Engelhardtia6:12-13.Hogg,R.H.1974.GreatBlack-backedGullattackingCommonGull.Brit.Birds67:242-243.Kock,K.H.1974.NahrungsokologischeUntersuchungenanMantelmowen(Larusmarinus)aufHelgoland.[StudiesonthefeedingecologyoftheGreatBlackbackedGull(Larusmarinus)onHelgoland.]Helgol.Wiss.Meeresunters.26:88-95.[InGermanwithEnglishabstract.)Love,J.A. Rona.1974.StatusoftheKittiwakeandGreatBlack-backedGullonNorthSeabirdRep.4:29-35.Seymour,N.R.1974.GreatBlack-backedGullsfeedingonliveeels.Can.Field-Nat.88:352-353.1973Andrle,R.F.1973. AsecondprobablehybridofLarusmarinusandL.argentatusontheNiagaraRiver.Can.Field-Nat. 171.Birkhead,T. R., C.LloydandP.Corkhill.1973.Oiledseabirdssuccessfullycleaningtheirplumage.Brit.Birds66:535-537.Bourget,A.A.1973.RelationsofeidersandgullsnestinginmixedcoloniesinPenobscotBay,Maine.Auk90:809-820.Corkhill,P.1973.ManxShearwatersonSkomer:populationandmortalityduetogullpredation.Brit.Birds66:126-143.Drury,W.H.1973-74.PopulationchangesinNewEnglandseabirds.Bird-Banding44:267-313;45:1-15.320

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GreatBlack-backedGullFitzgerald,G.R. andJ.C.Coulson.1973. ThedistributionandfeedingecologyofgullsonthetidalreachesoftheriversTyneandWear. Vasculum 58:29-47.Forsythe,D.M.1973.GullpopulationsatCharleston,S.C.,June1971toJune1972.Chat37:57-62.Godfrey,W.E.1973.Morepresumedhybridgulls:LarusargentatusX L.marinus.Can.Field-Nat.87:171-172.Hunt,G.L.,Jr.andM.W.Hunt.1973.inMaine andnorthwesternEurope.HabitatpartitioningbyforaginggullsAuk90:827-839.King,B.1973.GreatBlack-backedGullattackingasleepingman.BristolOrnithol.6:37.Leek,C.F.1973.AttemptsofgullstorobaCommon Egret andanOsprey.Cassinia54:32.Macdonald,S.M.and C.F.Mason. 1973.Predationofmigrantbirdsbygulls.Brit.Birds66:361-363.Parslow,J.L.F.andW.R.P.Bourne.1973.otherbirdsonAmBalg,WestSutherland.GreatBlack-backedGullsandSeabirdRep.3:15-24.Pasquier,R.1973.RecentChristmasCountincreasesofGreatBlack-backedGullsandMourning DovesintheN.Y.area.Linn.Newsl.26:3.Prys-Jones,o.E.1973.InteractionsbetweengullsandeidersinSt.AndrewsBay,Fife.BirdStudy20:311-313.Taverner,J.H.1973.GreatBlack-backedGullnestinginunusualhabitat.Brit.Birds66:398.Zobbe,B.1973.IagttagelserfraVestgronland.Dan.Ornithol.Foren.Tidsskr.67:65-66.1972Andrle,R.F.1972.AnotherprobablehybridofLarusmarinusand L.argentatus.Auk89:669-671.Dittberner,H.1972.Die Grossmowen(Larusmarinus,L.fuscusandL.argentatus)alsGastvogelmarkischer Gewasser. Beitr.-Vogelkd.18:141-155.Keskpaik,Y.andP.Khorma.1972.[Bodytemperaturesandheartrateofflyingsea-gulls(Larusm.marinusL.).]EestiNSVTead.Akad. Toim.BioI.21:109-116.[InRussianwithEnglishsummary.] 321

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GreatBlack-backedGullNilsson, X. 1972.Fagelrapporter.Narke1971.VarFagelvarld31:199-201.[InSwedish.]Ring,H.1972.TrutattackerandefiskgjusePandionhaliaeetus.ingOspreyPandionhaliaeetus.]VarFagelvarld31:129-130.withEnglishsummary.][Gullattack[InSwedishStenman,0., X. KomuandA.Ermala.1972.Kloraloosiharmaa-jamerilokkimyrkkyna.[FinnishexperienceofpoisoningHerringandGreatHlack-backedGullswithchloralose.]SuomenRiista24:107-116.[InFinnishwithEnglishsummary.] 1971Corkhill,P.1971.CannibalisticGreatBlack-backedGull.Brit.Birds64:30-32.Erwin, X. M.1971.Thebreedingsuccessof,twosympatricgulls,theHerringGullandtheGreatBlack-backedGull.WilsonBull.83:152-158.Harris,M.P.1971.EcologicaladaptationsofmoultinsomeBritishgulls.BirdStudy18:113-118.Schmidt,G.1971.Mantelmowen,Larusmarinus,alsKommensalen beim Kormoran,Phalacrocoraxcarbo.Vogelwelt92:24-25.[InGerman.]Svensson,L.1971.OmhavstrutenLarusmarinussompredator.[TheGreatBlack-backedGullLarus marinus-as-a predator.]VarFagelvarld30:125.[InSwedishwithEnglishsummary.] 1970Andersson,A. andK.E.Fridzen.1970.HavstrutfangarviggpaStockholmsstrom.[GreatBlack-backedGull(Larusmarinus)catchesscaup(Aythyamarila)ontheStreamof Stockholm:r--var Fagelvarld29: 267-269:--TIn Swedish.]Harris,M.P.lations.1970a.RatesandcausesofincreasesofsomeBritishgullpopuBirdStudy17:325-335.Hatch,J.J.1970.PredationandpiracybygullsataterneryinMaine.Auk87:244-254.Ingolfsson,A.1970a.ThemoultofremigesandrectricesinGreatBlack-backedGullsLarusmarinusandGlaucousGullsL.hyperboreusinIceland.Ibis112:83-9-2-.-Weaver, D.K.andJ.A.Kadlec.1970.A methodfortrappingbreedingadultgulls.Bird-Banding41:28-31.322

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GreatBlack-backedGull1969Beer,J.V.1969.ObservationsonthedispersalofgullsmarkedonSteepHolmandtheDenny.SteepHolmGull Nes. Stn.Rep.1968:4-9.Haugaard,N.1969. Et ynglefundafSvartbag(LarusmarinusL.).Dan.Fugle20:53.Ingolfsson,A.1969a.Behaviourofgullsrobbingeiders.BirdStudy16:45-52.1969b.Sexualdimorphismoflargegulls(Larusspp.).Auk86:732-737.1968Barth, E. K.1968a.EggdimensionsandlayingdatesofLarusmarinus,L.argen and NyttMag.Zool.15:5-34.Karpovich,V.N.andI.P.Tatarinkova.1968.[PopulationdynamicsofGreatBlack-backedandHerringGullsattheislandsontheMurmancoastandtheirinfluenceontheefficiencyofthereproductionoftheCommon Eider.] Pp.85-104inE.Kumari(ed.).[TheCommonEider(Somateriam.mollissimaL.)inthe ValgusTallin.[InRussianwithEnglishsummary.]Muller,H.P.1968.Mantelmowe(Larusmarinus)verschlucktRauchschwalbe(Hirundorustica).Ornithol.Mitt.20:252.[InGerman.]Presnall,C.C.1968.GreatBlack-backedGullattacksinjuredwigeon.Atl.Nat.23:107.Stollman,A.1968.[Theseagull(Larusmarinus)--anew memberoftheSlovakavifauna.]Biologia(Bratisl.)23:651-655.[InCzechoslovakian.]Threlfall,W.1968a.Thefoodof speciesofgullsinNewfoundland.Can.Field-Nat.82:176-180.1968e.ThehelminthparasitesofthreespeciesofgullsinNewfoundland.------Can.J.Zool.46:827-830.Wulf,J.1968.MantelmowefrisstBlesshuhu.Falke15:246.1967Barth,E.K.1967.StandardbodymeasurementsinLarusargentatus,L.fuscus, b. canus,andL.marinus.NyttMag.Zool.14:7-83.Harrison,J.G.andW.F.A. Bur.k.1967. Perilinperspective.AnaccountoftheoilpollutiollintheMedwayEstuary.KentBirdRep. :6, Spec.Suppl.24pp.323

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GreatBlack-backedGullIngolfsson,A.1967. lbe feedingecologyoffivespeciesoflargegulls(Larus)inIceland.Ph.D.thesis,Univ.Michigan/AnnArbor,MI.xiiand 186 pp:-McLaughlin,F.L.1967 ms.BehaviorstudyoftheinterrelationshipsofgullsandeidersinmixedbreedingcoloniesonPenobscotBay,Maine.Unpubl.filerep.,Maine Coop.Wildl.Res.Unit,Orono,ME.Peakall,D.B.1967.RecentchangesinthestatusoftheGreatBlack-backedGull.Kingbird17:69-73.Temme,M.1967."Begegnengszeremonien"beiunausgefarbtenMantelmowen(LarusmarinusL.).["Meetingceremonies"ofjuvenileGreatBlack-backedGulls(LarusmarinusL.).]Ornithol.Mitt.19:155-160.[InGermanwithEnglishsummary.]1966Buckley,P.A.1966.Foot-paddlinginfourAmericangulls,withcomments onitspossiblefunctionandstimulation.Z.Tierpsychol.23:395-402.Miller,J.C.1966b.TheGreatBlack-backedGullnestinginNewJerseyandadditionalnotesonnestingHerringGulls.Cassinia49:31.Preuss,N.O.1966.Svartbag.Feltornithologen8:12.[InDanish.]Randazzo,G.R.1966.SudiunacatturadiLarusmarinusLinn.inSicilia.Riv.Ital.Ornitol.36:366-367.[InItalian.]Temme,M.1966.UberdasVerhaltensrepertoirederMantelmowe(LarusmarinusL.),imWinterhalbjahrsowieeinigeBemerkungenuberdasVerhaltenderSilbermowe(LarusargentatusPontopp.)imgleichenZeitraum.J.Ornithol.107:70-84. 1965Andersson,A.1965.Havstruten(Larusmarinus)somhackfageliSkane.Medd.SkanesOrnitol.Foren.5:8-9-.----Bergman,G.1965.[CompetitionandfeedingecologyofgullsinasouthFinnishskerrygard.]Zool.Revy27:58-77.[InSwedishwithGermansummary.]Garrido,O.H. andF.G.Montana.1965.AvesneuvasparaCuba.PoeyanaInst.BioI.Ser.A10:1-6.[InSpanishwithEnglishsummary.]Gerasimova,T.D.1965.Ryboyadn.ptitsyiikhznacheniev rybnomkhoz.[FoodofgullsoftheMurmanCoast.]Moscow (Nauka)1965:194-209.[InRussian.]Harris,M.P.1965.Thefoodofsome gulls.Ibis107:43-53.324

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GreatBlack-backedGullParslow,J.L.F.1965.GreatBlack-backedGullspreyingonstormpetrels.Brit.Birds58:522-523.Vass,S.E.1965.GullbreedingrecordsfromPrinceEdwardIsland.Can.FieldNat.79:152-154.Williams,I.C.andM.P.Harris.1965.TheinfectionofthegullsLarusargentatusPont.,L.fuscusL.,and L.marinusL.withcestodaonthe Wales.Parasitology55:237-256.1964Ammermann,D.1964.BeringungsergenisseanMantelmowen(Larusmarinus)desdeutschenuberwinterungsgebiets.Vogelwarte22: German.]Harris,M.p.1964a.AspectsofthebreedingbiologyofthegullsLarusargentatus,L.fuscusand L.marinus.Ibis106:432-456.1964b. Theincidenceofsomespeciesoftrematodainthreespeciesof-----LarusgullsinWales.Ibis106:532-536.1964d. Measurements andweightsofGreatBlack-backedGulls.Brit.------Birds57:71-75.1963Baillie,J.L.1963. The13mostrecentOntarionestingbirds.OntoFieldBio!.17:15-26.Bruun,B.1963.Svartbagen(LarusmarinusL.)i Danmark. [TheGreatBlackbackedGull(LarusmarinusL.)inDenmark.] Dan.Ornithol.Foren.Tidsskr.57:94-98.[InDanishwithEnglishsummary.]Goethe,F.1963.Das"Wegsehen"(facingaway)beiderMantelmowe(Larusmarin L.).J.Ornithol.104:437-438.[InGerman.] Lemmetyinen, R. 1963.LokkienesiintymisestajaravinnostaGullkronaselankoillisosassa.[OccurrenceandfeedinghabitsofgullsinthenortheasternpartoftheareaofGullkrona.]SuomenRiista16:69-82.[InFinnishwithEnglishsummary.]Saunders,D.R.1963.TheGreatBlack-backedGullon Skomer.Nat.Wales8:99-104.Williams,L.E.,Jr.1963.NewspecimenrecordsforNorthFlorida.Fla.Nat.36: 127.Zelenka,G.1963.Notes onthedepredationoftheBrownRatbytheGreatBlackbackedGullontheCalfofMan, 1960.Proc.IsleManNat.Rist.Antiq.Soc.6:431-433.325

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GreatBlack-backedGull1962Harris,M.P.1962b.TheLarusgullsbreedingon Skomer, Skokho1mandGrassholmislands.Nat.Wales8:56-58.1962d.RecoveriesofringedGreatBlack-backedGulls.BirdStudy9:192-197.Saunders,D.R.1962.TheGreatBlack-backedGullonSkomer.Nat.Wales8:59-66.1961Isenmann,P.andB.Schmidt.1961. argentatusXLarusmarinus.Observationsaproposd'unhybrideLarusOiseauRev.Fr.Ornithol.31: Mansueti,R.J.1961.WaterfowlpredationbyandrecordsoftheGreatBlackbackedGullinChesapeakeBayduringwinterandspring.ChesapeakeSci.2:102-104.Marshall,R.V.A.1961a.backedGullandHeron.Attackandcounter-attackbetweenaGreatB1ackBrit.Birds54:116.Rooke,K.B.inDevon.1961b.GreatBlack-backedGullattackingCoot.Brit.Birds54:243.1961.AprobablehybridGreatBlack-backedGullXHerringGullBrit.Birds54:161-163.Tomkins,I.R.Georgia.1961.AsecondspecimenoftheGreatBlack-backedGullfromOriole26:10-11.1960 Bergman,G.lands.1960.UberneueFuttergewohnheitenderMowenandenKustenFinnOrnisFenn.37:11-28.[InGermanwithFinnishsummary.)Jeh1,J.R.,Jr.1960.AprobablehybridofLarusargentatusand L.marinus.Auk77:343-345.My1ne,C.K.1960.PredationofManxShearwatersbyGreatBlack-backedGullsonSkomer.BirdNotes29:73-76.Snyder,D.E.1960.GreatBlack-backedGullskillingDovekies.Auk77:476477.326

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GreatBlack-backedGull1959Borg,K.1959.Nabblangdochkroppsvikthosgratrut(Larusargentatus),havstrut(L.marinus)ochsilltrut(L.fuscus).[Billlengthsandbodyweights-forHerringGulls(Larus argentatUs), GreatBlack-backedGulls(L.marinus),andLesserBlack-backedGulls(L.fuscus).]VarFagelvarld 18: 311-317.[InSwedishwithEnglishsummary.]------Colston,P.R.,B. Newport andM.J.Carter.1959.GreatBlack-backedGullattackingmigrantStarling.Brit.Birds52:312-313.Gamble,G.1959.GreatBlack-backedGullattackingmigrantBlackbird.Brit.Birds52:164.lIes,D.B.,J.R.GovettandD.Rushforth.1959.InlandbreedingofGreaterBlack-backedGull.Naturalist870:98.King,B.1959.GreatBlack-backedGulldivingfromaheightandsubmerginginaninlandwater.Brit.Birds52:198.Lewis,H.F.1959ms.PredationofeiderducksbyGreatBlack-backedGullsinNovaScotia.Unpubl.Rep. NovaScotiaDep. Lands Forests,Halifax.16pp.Slinn,D.J.1959.BreedingpopulationoftheGreatBlack-backedGullinMann1957.Peregrine3:2-7.Stevenson,H.M.1959.GreatBlack-backedGull(Larusmarinus).Fla.Nat.32:146.1958Davis,T. A.W.1958.ThebreedingdistributionoftheGreatBlack-backedGullinEnglandand Walesin1956.BirdStudy5:191-215.1957 Cobb,S.1957.GreatBlack-backedGull(Larusmarinus).Auk74:498.Lennerstedt,I.1957.AlbinistikhavstrutLarusmarinusiBohuslan.[AlbinisticGreatBlack-backedGullLarus Bohuslan.]VarFagelvarld16:52,58.[InSwedishwithEnglishsummary.]1956Campbell,J.W.1956.StatusofGreatBlack-backedGullinScotland.Brit.Birds49:152.Cormack, D. M.,R.S. CormackandR.H.Poulding.1956.BreedingofGreatBlack-backedGullinGloucestershire.Brit.Birds49:153.327

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GreatBlack-backedGullKrug,H.H.1956.GreatBlack-backedGullsnestingonLittleHaystackIsland,LakeHuron.Auk73:559.1955 Andrew,D.G.1955.StatusofGreatBlack-backedGullinlandinScotland.Brit.Birds48:455-456.Cobb,F.K.1955.NotesonthebehaviourofGreatBlack-backedGulls.Trans.SuffolkNat.Soc.9:263-266.Hamann,D.F.1955.andCanaveral.GreatBlack-backedGull,Larusmarinus,atOrmondBeachFla.Nat.28:59-60.Larsen,A.andF.B.Jorgensen.1955.Svartbag(LarusmarinusL.)somynglefuglpaSaltholm.Dan.Ornithol.Foren.Tidsskr.49:136138.[InDanish.]1954 Gudmundsson,F.1954.Islenzkirfuglar.X.Svartbakur(LarusmarinusL.).[Icelandicbirds.X.TheGreatBlack-backedGull (Larus-marinus L.).]Natturufraedingurinn24:177-183.[InIcelandicwithEnglishsummary.]Nilsson,N.J.1954.Havstrut(Larusmarinus)hackandevidinsjoariBohuslan.[GreatBlack-backedGulls(Larusmarinus)breedingontwolakesinBohusIan.]VarFagelvarld13:38-40.[InSwedishwithEnglishsummary.]Schmidt,G.1954.ZumRaubenderSchmarotzer-Raubmowe.Vogelwelt75:147-151.[InGerman.]1953Cross,A.1953a.GreaterBlack-backedGulldivingforfood.Scott.Nat.65:129.Jonston,T.L.1953.TheGreaterBlack-backedGullontheCumberlandSolway.Trans.CarlisleNat.Hist.Soc.8:8-13.1952 Dawn,W.1952.HerringGullregurgitatesuponBlack-backedGull'sdemand.Linn.Newsl.6(6/7):2-3.1951Barnes,J.A.G.1951.InlandbreedingofGreatB[l]ack-backedGull.Brit.Birds44:36.Stimson,L.A.1951.NotesfromSouthFlorida.Fla.Nat.24:81.328

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GreatBlack-backedGullWilson,W.H.1951.HybridGlaucousXGreatBlack-backedGullatLimerick.Brit.Birds44:286-287.1950 Adams, R.G.1950a.GreatBlack-backedGulldroppingfoodfromheight.Brit.Birds43:382.1950b.Skua-liketacticsofGreatBlack-backedGull.Brit.Birds43:------382.Butt,O.V.,R.G.PettittandA.E.Vine.1950.GreatBlack-backedGullsroostinginland.Brit.Birds43:61.Davis,T.A.W.1950.GreatBlack-backedGullwithyellowlegs.Brit.Birds43:164.1949 Wilmahn, B.1949.Ringmerkings-resultaterforsvartbak(LarusmarinusL.).StavengerMus. Arbok 58:123-128.[InSwedish.]1948 Anon.1948.GreatBlack-backedGullsdivingforfood.Brit.Birds41:93-94.Conder,P.J.1948.GreatBlack-backedGulldroppingprey.Brit.Birds41:250.1947Barnes,A.M.1947. TheBlack-backedGull.Annu. Rep. BowdoinSci.Stn.8:18-21.Harber,D.D.andM.Johns.1947.GreatBlack-backedGulldroppingrat.Brit.Birds40:317.1946Hare,M.1946.AvianleukosisandtheGreatBlack-backedGull.Auk63:245246.Mayfield,H.1946.ThestatusofGreatBlack-backedandGlaucousGullsinMichigan.Jack-PineWarbler24:19-22.1945 Addy, C. E.1945.GreatBlack-backedGullkillsadultBlackDuck.Auk62: 329

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GreatBlack-backedGullBrooks,M.1945.GreatBlack-backedGullinMonongaliaCounty,WestVirginia.Auk62:634.Gross,A.O.1945.coastofMaine.ThepresentstatusoftheGreatBlack-backedGullontheAuk62:241-256.Svendsen,T.1945.Svartbagens,Larusmarinus,YngleforekomstpaaSandrevetSydforLaeso1944.Dan.Ornithol.Foren.Tidsskr.39:54-55.1944Wilcox,L.1944.GreatBlack-backedGullbreedinginNewYork.Auk61:653654.1943Fitter,R.S.R.1943.StatusoftheBlack-backedGullsintheLondonarea.Brit.Birds36:163-164.Gerber,R.1943.EineMantelmowe,LarusmarinusL.,beiPeniginSachsen.Ornithol.Monatsber.51:44.Mayfield,H.1943.ofLakeErie.GlaucousandGreatBlack-backedGullsatthewesternendWilsonBull.55:129-130.Salomonsen,F.1943.Svartbagen(LarusmarinusL.)iLaesoomradet.Dan.Ornithol.Foren.Tidsskr.37:221-226.[InDanish.]1942Cleghorn,J.D.1942.GreatBlack-backedGullkillingAmericanGoldeneye.Auk59:584-585.1941Svendsen,L.andR.Horring.1941.Svartbage,Larusmarinus,ynglendevedLaeso.Dan.Ornithol.Foren.Tidsskr.35: 1938Johnson,R.A.1938.Predationofgullsinmurrecolonies.WilsonBull.50:161-170.Jourdain,F.C.R.1938.OntheoccurrenceofLarusmarinusinSpitsbergen.Ibis(14thSer.)2:539-540.Marshall,A.J.1938.OntheoccurrenceofLarusmarinusinSpitsbergen.Ibis(14thSer.)2:341-342.330

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GreatBlack-backedGull1937Bodenstein,G.1937.VonderWanderungenderSeemowen(Larushyperboreus,L.marinus,L.argentatus,L.fuscus,undL.canus)inOstlichenOstsee nach denBeringungsergebnissen.Schr. Konigsberg69:223-234.Broekhuysen,G.J.,Jr.1937.GedragingenvangeslachtsrijpeennognietgeslachtsrijpeZilver-enGroteMantelmeeuwen(LarusargentatusPont.etLarusmarinusL.)buitendebroedtijd.Ardea26:159-172.[InDutchwithEnglishsummary.]Kuerzi.R. G.1937. lbe Black-backedGullasapredator.Proc.Linn.Soc.N.Y.48:97-98.Lockley,R.M.1937.Black-backedandHerring-GullsandRavensfeedingonants.Brit.Birds30:325-326.Loppenthin,B.1937.EtBesogpaaknotten.NoteromSvartbag,Larusmarinus,somdanskynglefugl.Dan.Ornithol.Foren.Tidsskr.31:142-146.1936Cottam,C.1936a.EarlymigrationoftheGreatBlack-backedGull.Auk53:79.1936b.StatusoftheBlack-backedGull.Auk53:332-333.Timmermann,G.andF.Gudmundsson.1936.EinBesuchderMantelmowenkolonieAufderInselSandey imThingvallavatn(Sudwest-Island).Beitr.Fortpfl.Vogel12:14-21.1935Broekhuysen,G.J.,Jr.1935.GedragingenvannognietgeslachtsrijpedochreedszelfstandigeZilver-enGroteMantelmeeuwen(LarusargentatusPont.etLarusmarinusL.).Ardea24:239-250.[InDutchwithGermansummary.]Gross,W.A.O.1935.ThelifehistorycycleofLeach'sPetrel(Oceanodromaleucorhoaleucorhoa)ontheouterseaislandsoftheBayofFundy.Auk52:382399.Sprunt,A.,Jr.1935.TheBlack-backedGullontheSouthCarolinacoast.Auk52:447-448.1934Barth,E.K.1934.HegrerofsvartbakostenforLindesnes.Naturen58:28-30.[InDanish.]Cottam,C.1934.PossibleextensionofregularwinterrangeoftheGreatBlack-backedGull.Auk51:376.331

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Lockley,R.M.1932.Herring-Gulls.GreatBlack-backedGullRoberts,R.1934.Notes onthebirdsofcentralandsoutheastIcelandwithspecialreferencetofood-habits.Ibis(13thSer.)4:239-264.Shelley,L.O.1934b.GullnotesfromtheNewHampshireand Mainecoasts.Auk51:83.1933Harllee,H.L. 1933.GreatBlack-backedGullontheSouthCarolinacoast.Auk50:217.Harrisson,T.HandH.G.Hurrell.1933.NumericalfluctuationsoftheGreatBlack-backedGull(LarusmarinusLinn.)inEnglandandWales.Proc.Zool.Soc.Lond.103:191-209.Holt,E.G.1933.GreatBlack-backedGullinMaryland.Auk50:216-217.Oldham, C. 1933.GreatandLesserBlack-backedGullsinNorthWales.Brit.Birds27:38-41.Taylor,W.R.1933.EarlybreedingofGuillemot,Razorbill,Shag andGreatBlack-backedGullinCornwall.Brit.Birds27:52.1932Haworth,A.G.1932.ColourofirisinGreatBlack-backedGull.Brit.Birds26:100.Jackson,C.F.andP.F.Allan.1932.AdditionalnoteonthebreedinginMaineoftheGreatBlack-backedGull(Larusmarinus).Auk49:349-350.Incubation-periodsofLesserandGreatBlack-backedandBrit.Birds25:310-313.Robinson,H.W.1932.ColourofirisinGreatBlack-backedGull.Brit.Birds Birds26:140.1931Eaton,R.J.1931.GreatBlack-backedGull(Larusmarinus)breedinginEssexCounty,Massachusetts.Auk48:588-589.-----Harrisson,T.H.andH.G.Hurrell.1931.NumericalstatusoftheGreatBlack-backedGullinDevon.Brit.Birds25:136-138.Lonnberg,E.1931.Fargeninnabbenavhavstruten,Larusmarinus.FaunaFlora26:183-184.332

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GreatBlack-backedGullMeans,W.G.and R.J.Eaton.1931.GreatBlack-backedGull(LarusmarinusL.)breedinginEssexCounty,Mass.Bull.EssexCo.Ornithol.Club13:4-6.Norton,A.H.andR.P.Allen.1931.BreedingoftheGreatBlack-backedGullandDouble-crestedCormorantinMaine.Auk48:589-592.Rasmussen,S.1931.Svartbag,LarusmarinusL.,somDanskYnglefugle.Dan.Ornithol.Forens.Tidsskr.25:125-127.Seton,G.1931.GreatBlack-backedGullpreyingonStorm-Petrels.Brit.Birds25:60.1925Robinson,H.W.1925.Black-backedGull.ColouroftheeyesintheHawfinch,PochardandGreaterIbis(12thSer.)1:949-951.1924Blathwayt,F.L. 1924.BreedingoftheGreatBlack-backedGullinDorset.Brit.Birds18:61.1923Ingram,G.C.S.andH.M.Salmon. 1923.GreatBlack-backedGullbreedinginSomerset.Brit.Birds16:214-215.Lewis,S.1923.GreatBlack-backedGullbreedinginSomerset.Brit.Birds17:67.Robinson,H.w.1923.GreatBlack-backedGullkillingRazorbill.Brit.Birds17:67-68.1922Robinson,H.W.1922.GreatBlack-backedGullnestinginWestmoreland.Brit.Birds16:139.1920Lewis,H.F.1920.AmongtheCoffinCarriers(Larusmarinus).KansasFieldNat.34:101-106.1919Lonnberg,E.1919.Hybridgulls.Ark.Zool.12:1-22.333

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Great Black-backedGull Palmgren.R.1918.LarusargentatusBrunn[male]Xmarinus(L.)[female].HelsingforsMedd.Soc.FaunaFloraFenn.44:125-127. Kihlen. G.1917.Tillfrarganomalfagelnsruggningarochhafstrutenshackningvidsotvatten.FaunaFlors12:229-230.Deffay,M.1913. Le Goelandmarin.HavreSoc.Linn.1:17-19.Robinson,H.W.1911a.Slaughterof Manx ShearwatersbyBlack-backedGulls.Brit.Birds5:55.1911b. andManxShearwaters.Brit.Birds5:88.Stubbs. F. J.1910.BreedingoftheGreaterBlack-backedGullinLakeland.LancashireNat.3:266.Warren,R.1908.UnusualsiteforaGreatBlack-backedGull'snest. Zooloj;ist (Lond.)12:396. Ridgway.R. 1882.TheGreatBlack-backedGullfroma newlocality.Bull.Nuttall Ornithol. Club7:60.AdultGreatBlack-backedCullinbreedingplumage.PhotographbyPatLynch.334

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GULL-BILLEDTERN(Sternanilotica)IDA:Sandterne,DU:Lachstern,FI:Hietatiira,FR:Sternehansel,GE:Lachseeschwalbe,IC:Hlaturperna,IT:Rondinedimare,zampenere,NW:Sandterne,PO:Rybitwykrotkodziobej,SP:Picodegaviota,Pagazapiconegra;SW:Engelsktarna,Sandtarnal GENERAL DISTRIBUTIONNorthAmericaAlongtheAtlanticcoast,Gull-billedTernsnestirregularlyinNewYork andNewJersey(Buckleyetal.1975) andregularlyfromMarylandsouthtoFlorida,withoccasionalinlandbreedingonfreshwaterinthelatterstate.TheyalsobreedalongtheGulfcoastfromFloridatoTexaswheretheyaremostabundant(Map9).AnisolatedcolonyispresentontheSaltonSeainCalifornia,andothersbreedonthewestcoastofMexico.EasternbirdswinterlocallyalongtheAtlanticandGulfcoasts(Map10),primarilyinthewesternportionoftheGulf.Mostbirds,however,winterfarthersouthalongthecoastsofCentralAmerica andalongthenortherncoastofSouthAmerica. Birds fromthewesternpopulationwinterinCentralAmericaandcoastalSouthAmerica(AOU1957, Blake 1977).WorldDistributionGull-billedTernsarenearlycosmopolitanintheirbreedingdistribution.TheybreedintheCaribbeanintheBahamasandVirginIslands(AOU1957),inSouthAmericaonthecoastsofBrazil,Uruguay,andnorthernArgentina(Blake1977),andinsouthwesternEcuador(Marchant1958).In1972theybredlocallyinEuropeinDenmark, West Germany,France(intheCamargue),Spain,Italy,Romania,Greece,Turkey,andthesouthEuropeanU.S.S.R.(Moller1975d).TheyformerlybredinPoland(AOU1957),England,theNetherlands,Portugal,Austria,Hungary,andBulgaria(Moller1975d).InAfrica,Gull-billedTernsbredin1972alongthenorthwesterncoastinSenegal,intheBancd'ArguinoffMauritaniaandalongtheMediterraneancoastinTunisia.EarlierbreedingrecordsalsoexistforMorocco andAlgeria 1975d).BreedingpopulationsarefoundinAsiafromthePersianGulftoIndiaandtheMalayArchipelago.Gull-billedTernsalsobreedonmainlandAustralia,and(rarely)onTasmania(Dement'evand Gladkov1951,Vaurie1965,Serventyetal.1971).PopulationsineasternEuropeprimarilywinterinnorthwesternAfricafromSenegaltoNigeriawhilepopulationsinwesternEuropewinterineasternAfricafromtheSomaliRepublictoMozambique and Botswana(Moller1975b).PopulationsfrommoresouthernbreedinglocalitiesinAsia,Australia,andEcuadorapparentlywinterlargelyincoastalareasnearthebreedingrange.Taxonomicnote:ThisspeciesisfrequentlyplacedinthemonotypicgenusGelochelidon.335

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GULL-BILLEDTERNBIRDNAME:\Map988" ..GULFOFMEXICO "" ,. \ :r-,J --.. ".1'...r-?I < ) / IsA !X'TEMAX.NO.YEARNorth Carolina colony sites of lessthan25breedingadultsnot censused. Texas colony sites of lessthan50breedingadultsnot censused.BreedingRangeMapforSoutheasternUnitedStatesDALLAS-----Numbersinboxesdenotemaximum estimates of breeding birdsatcoloniesinrecent years. First figure indicates mcucimumnumberof birds. Second figure indicatesyearinwhichestimatewasobtained.

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-iiii GULL-BILLEDTERNBIRONAME: ,) __ i-,'---N88" GULFOFMEXICO I (.:.---fi ,.-.J'r sX' A EWinterDistributian lap forSoutheastI'DUnitedStates< DALLAS BIRDSPER10PARTY-HOURSIILess t"'an 0.1 IIIIIIII!.m 0.1-0.5 0.5-2 .Wd.More than2 (Adaptedfrom IYI.rak, 1974) TINDIVIDUALSOBSERVEDDURINGCHRISTMASBIRDCOUNTS,1973-1977(ARITHMETICMEAN) @Number of individuals o Lellthanone individual None observedH-------+-------\::-0---+----+--,-000.. Map10

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Gull-billedTernDISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESGull-billedTernswereoncecommonalongthecentralandthesoutheasternAtlanticcoastoftheUnitedStates,althoughearlyrangeandstatustrendsofpopulationsarenotwelldocumented.Inthe19thcentury,theybredcommonlyinNewJersey(Stone1909),Virginia(Bailey1913),andGeorgia(Burleigh1958),andprobablyinothercoastalstatesaswell.Thepopulationdeclineinthelate1800'swasapparentlytheresultofplumehuntingandperhapsthelossofmarshhabitat(Bent1921,Burleigh1958).Consideredanuncommonspeciesinthesoutheastearlyinthiscentury,theGull-billedTernhasrecentlyreoccupiedmuchofitsformerrange,atrendthatisstillcontinuing(Savell1972,Buckleyetal.1975).AlthoughthespeciesisnowcommonlocallyalongtheAtlanticcoast,populationsarenotyetstable(Therresetal.1978).NorthCarolinaGull-billedTernsareseenlocallyalongthecoastofNorthCarolinaandareoccasionallyseeninland(Teulings1975d).Nestinghasbeenreportedatmanyplacesalongthecoast(Map9).Thespeciesispresentprimarilyinsummer,buta fewbirdsmaybepresentinfallandwinter(Pearsonetal.1942).TheseternsusuallyreturntotheircoloniesinNorthCarolinafrommid-ApriltoearlyMay. Eggsarelaidfromaboutmid-Maytoaslateasmid-July,anddependentyoungmaybepresentintoearlyAugust(ParnellandSoots1979ms).Gull-billedTernsarenowconsiderablymoreabundantasnestingbirds(Table38)inNorthCarolinathantheywereatthebeginningofthiscenturY,buttheystillbreedonlyinsmallnumbers andareregardedasaspeciesofspecialconcerninNorthCarolina.TheynestfromRoanoke SoundsouthtotheCapeFearRiver(ParnellandSoots1979ms).Amajorityofthebreedingpopulationnestson man-madeoralteredsites(65.8%ofca.1,690breedingbirdsin1973[SootsandParnell1975a],83.6%of900breedingbirdsin1976[Portnoyetal.1981],and 84.3%ofca.1,240breedingbirdsin1977[ParnellandSoots1979ms]).Coloniestendtobesmall(1),butalargeproportionofthestatepopulationisconcentratedata fewsites.Thirty-onecoloniescensusedin1976averaged29.0breedingbirds(median20)(Portnoyetal.1981);21coloniescensusedin1977(ParnellandSoots1979 ms)averaged31breedingbirds(median30).In1976thefourlargestcoloniesheld36%ofthebreedingpopulation.Twooftheselargecolonieswith130and60breedingbirds,respectively,wereondredge-spoilislandsintheCapeFearRiver;onecolonywas on adredge-(1)Old Worldcoloniesarealsosmallbutincreaseinsizeinthesouthernportionsoftherange.ColoniesontheChernomorskiiReserveinthesouthernU.S.S.R.usuallycontain20to40breedingbirds,andonlyrarelyasmanyas100(Borodulina1960).ColoniesinDenmarkaverage72breedingbirdswitha maximumof400 andcoloniesinGermanyaverage10breedingbirdswitha maximumof110)(Moller1981a).Coloniesinsouthernbreedingareasrarelymayholdasmanyas2,000breedingbirds(Moller1982).338

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Gull-billedTernspoilisland'atHatterasInlet(90breedingbirds);andone wa?pn abarrierbeachatPortsmouthIslandontheOuterBanks(62breedingbirds)(Portnoyet.al1981).In1977thefourlargestcoloniescontainedalmost65%ofallnestsfound.Thelargestcolony,locatedon adredgedislandintheCapeFearRiver,heldabout330breedingbirds.Theotherthreelargecolonieswere:onefoundonBrantIsland(adredgedislandintheMoreheadChannel,CarteretCounty),withabout200breedingbirds;oneonadredgedislandatDrumInlet,CarteretCounty,thatheld170birds;andonecolonyonthebarrierbeachonthenorthshoreofDrumInlet,with104breedingbirds(ParnellandSoots1979ms).SouthCarolinaSpruntandChamberlain(1949)consideredtheGull-billedTernacommonsummerresidentandirregularbreedingbirdinthisstate.ThisspecieshasbeenseenasearlyasMarch(Teulings1973c)andaslateasNovember(Teulings1975a)and December(Chamberlain1968),butmostarepresentfromApriltoSeptember(SpruntandChamberlain1949).TheseternsnestregularlynearCharlestonandatCape RomainNWR(Map9),successfullyinsomeyearsandnotsosuccessfullyinothers(Teulings1971d,1972d).EgglayingbeginsinthefirstweekofMayandyounghatchfromJunethroughJuly;youngfledgeasearlyas3July(BlusandStafford1980).Recentnestingpopulationsinthelowhundredshavebeenfoundoncoastalestuarineislands(BlusandStafford1980).Thelargestnestingcoloniesinrecentyearsareoneon RaccoonKeythatheldabout310 and 320breedingbirdsin1972 and1973,respectively(BlusandStafford1980),andonetothesouthonBirdIslandinBullsBaythatcontained220breedingbirdsin1976(Portnoyetal.1981).GeorgiaGull-billedTernsarenowuncommon andirregularsummerresidentsalongtheGeorgiacoast,althoughoccasionalnestinghasbeenreportedinChathamandMcIntoshCounties(Burleigh1958,Dentonetal.1977).Theyalsooccasionallyoccurinland(LeGrand1978).FloridaFewGull-billedTernswererecordedinFloridaintheearlypartofthiscentury(Howell1932).Thefirstnestingwasnotedin1932 (Weston1933),butsincethat time theGull-billedTernhasbecome aregularbreedingbird,mostcommonontheAtlanticcoast.Kale(1979msa)reportedacolonyofabout25pairsattheMayport NavalStationintheSt.JohnsRiverInletinadditiontothecoloniesattheBirdIslandsinNassauSoundandthoseonspoilislandsintheBananaRiveronMerrittIslandNWRindicatedonMap9.ThecolonyatMerrittIslandNWRcontainedover150pairsin1977(Map9).Gull-billedTernsbreedmuchlessabundantlyontheFloridaGulfcoast;Kale(1979msb)consideredthemrarethere,notingthata fewpairsnestonspoilislandsinTampa and BocaCiegaBaysalongthecentralwestcoast,aswellasonSt.JoeIslandalongthenorthernGulfcoast.Mostegglayingoccursinmid-tolateMayorearlyJune(SchreiberandSchreiber1978);dependentyoungareprobablypresentintoAugust.339

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Gull-billedTernAlabamaGull-billedTernsareuncommon summerresidentsinAlabama.BreedingoccursinsouthernMobileCountywhereeggshavebeenreportedfrom22Maythrough16June.TheseternsarecommonmigrantsontheAlabamacoast,witha maximumof75+birdsrecordedatFortMorgan on 6August,butfewarepresentinwinter(Imhof1976b).Twenty-threebreedingbirdswerereportedonDauphinIslandin1976(Portnoy1977),thelargestnumberofGull-billedTernsfoundnestinginAlabamainrecentyears.MississippiTheseternsareuncommoninMississippi,butbreedinsmallnumbers oncoastalislandsandatthemouthofthePascagoulaRiver(Burleigh1944,Jacksonetal.1980).EggshavebeenfoundinlateMayandearlyJune(Jacksonetal.1980),anddependentyoungarepresentatleastthroughJuly.RecordsofoccurrencearefromlateMarchtoearlySeptember(Gandy andTurcotte1970,Jacksonetal.1979msb).LouisianaLowery(1974)consideredGull-billedTernstobefairlycommonpermanentresidentsinLouisiana,leastabundantinsummer.SmallnumbersnestonislandsoffthecoastandneartheRigoletsPass.Portnoy(1977)foundonlyfourcoloniesinLouisianain1976,buthemayhaveoverlookedsomenestingareas.Thelargestbreedingcolony(112birds)containednearlythree-quartersoftheGull-billedTernsfoundnestingin1976.FoundonspoilinAtchafalayaBay(Portnoy1977),thiscolonywasstillactivein1978.Thatyear,afifthcolonywasfoundonGrassyIslandinGrandIslandPass(Portnoy1978ms),butnonebredattwocolonies(anotherinAtchafalayaBay andoneatCurlewIsland)whereGull-billedTernshadbredin1976.AllGull-billedTernsbreedinginLouisiana,Alabama,andMississippiin1976werefoundnestinginBlackSkimmercolonies(Portnoy1977).TexasGull-billedTernsbreedcommonlybutlocallyalongtheTexascoast(Map arepresentthroughouttheyearbutarelessnumerousinwinter.Eggshavebeenrecordedfrom14Aprilthrough17July(Oberholser1974),butthenestingseasonisusuallyshorter;in1977,mostyounghadfledgedby 28June(Chaneyetal.1978).TexasprobablyholdsmorethanhalftheentirenumberofGull-billedTernsnestingintheUnitedStates.Populationsthereareevidentlystable(Blacklocketal.1978ms).Censusesfrom1973through1978revealedpopulationscontainingfrom1,370breedingbirdsin1974toapproximately5,320in1973 (mean=2,975)(Blacklocketal.1978,Blacklocketal.1978ms.).Thesenestingpopulationsweremostabundantalongthecentralcoast.From 1973to1976,thelargestcoloniescontained400ormorebreedingbirds.OnecolonyfoundonspoilinKlebergCountyheldabout1,440and 400birdsin1973 and1975,respectively;anothercolonyonislandsinBaffinBayinKlebergCountyaveragedabout1,020breedingbirdsin1975 and 1976;andathirdcolonyonspoilinCameronCountyheldabout450birdsin1973.However,mostcoloniesareconsiderablysmaller,andonlya fewcontainmorethan100breedingbirds.Of78coloniescensused1973-1976(Blacklocketal.1978),almost54%contained 50orlessbreedingbirds;themediansizeofthesecolonieswas 50birds.340

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Gull-billedTernSYNOPSISOFPRESENTDISTRIBUTIONANDABUNDANCEBreedingTheGull-billedTernisnearlycosmopolitaninitsdistributionandbreedsonallcontinentsexceptAntarctica(AOU1957).Inallareas,however,theyseemtobe uncommon andlocal.InNorth thisspeciesbreedsregularlybutlocallyalongtheAtlanticcoastfromNewJerseysouthtoFloridaandalongtheGulfcoast.AnisolatedgroupnestsattheSaltonSeainCalifornia.Gull-billedTernsalsonestintheWestIndiesandonthecoastofMexico(Bond1961,AOU1957).Americancoloniesnowareusuallysmall,butthespeciesprobablywas moreabundantacenturyagothanitisatpresent.Judgingfromrecentcensuses,mostoftheNorthAmericanpopulationbreedsalongthecoastofthesoutheasternUnitedStates.FigureslistedinTable38probablyunderestimatethenumbersbreedinginsomeareas(e.g.,Florida)becausethisspeciesiseasilyoverlookedoncensusesandbecausethefewrecentsurveysofcoastalwaterbirdsinthesoutheasthavebeenincomplete.Only a fewGull-billedTernswerefoundnestingalongthenortheasternAtlanticcoastin1977(Erwin1979a).Fourcolonieswith38breedingbirdswerefoundinNewJersey,and twocoloniescontaining310birdswerereportedinVirginia.ThepopulationbreedinginMarylandcontainedabout70birdsin1951(Erwin1979a),butonlyonepairwasfoundtherefrom 1974to1976;nonewasfoundin1977(Therresetal.1978).Twopairswerefoundbreedingon LongIsland,NewYork,in1975(Buckleyetal.1975),butthespecieshasnotbeenknowntonesttheresince(Erwin1979a).WefoundfewrecentdataonrecentpopulationsoftheGull-billedTernintheOldWorld.In1972about90bredinDenmark, 6bredinWest Germany,about1,800bredinSpain,400bredinSenegal,about50bredinItaly,about800bredinGreece,6bredinRomania,about800bredinTurkey,andabout230 bred intheCamargueofFrance(Moller1975d);thebreedingpopulationinthelatterareahadincreasedtoabout400by1979(Hafneretal.1980).About13,000bredontheBancd'ArguinoffMauretaniain1959(Moller1975d).In1972another1,000bredinAfghanistan;about2,000bredinIraq,Iran,PakistanandIndia;and no morethan4,000bredintheSovietUnion(Moller1975d).Mollerestimatedthatthetotalnumberbreedingin1972inEurope,AfricaandwesternAsiawasabout24,000.Fromourlimiteddata,thesoutheasternUnitedStatesappearstocontainasignificantproportionoftheglobalworldpopulation.Asa thisareaiscriticaltothewell-beingoftheentirespecies.WinterandMigrationGull-billedTernpopulationsmaybeeithersedentaryormigratorywiththepopulationsinnorthernNorthAmericaandthenorthernPalearcticmovingsouthforthewinter.BirdsbreedingintheUnitedStatesarebelievedtomigrateprimarilyalongthecoasts(Bent1921)toWinteringgroundsincentralandnorthernSouthAmerica,butnostudiesofdispersalandmigrationbasedon markedbreedingbirdsareavailabletoindicatethelocationofparticularlyimportantmigratoryorwinteringareasforpopulationsnestinginthesoutheasternUnitedStates.Smallnumberswinteralongthecoastsof341

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Gull-billedTernTable38.RecentestimatesofGull-billedTernpopulationsnestinginthesoutheasternUnitedStates.PercentofMaximumnumber NumberfoundsoutheasternfoundbreedingbreedinginbreedinginrecentyearsState1976(a)populationandyearNorthCarolina90025.91,690(1973)SouthCarolina2306.6680 (1972)(b)Georgianone noneFlorida-AtlanticCoast(c)2106.01,070(1975)Florida-GulfCoast????Alabama 200.620(1976)Mississippi20.064(1979)Louisiana1504.3150 ( 1976)Texas1,96056.55,320(1973)3,472(a)Figuresabove10areroundedtothenearest10.(b)Totalislowbecauseoneareawasnotcountedduringpeaknesting(BlusandStafford1980).(c)ThefigureslistedfortheFloridaAtlanticcoastfor1976arederivedfromPortnoyet al. (1,981)(inwhichthesedatawerelistedasfromGeorgia)andfrom Map 9.Thetotalnumberbreedinginthisstateispoorlyknown.thesoutheasternUnited (Map10),butlargewinteringpopulationsarefoundonlyalongtheTexascoast.BirdsfromtheeasternPalearcticwinterprimarilyinsouthernAfrica(Moller1975b)withotherswinteringalongtheMediterranean(BOU1971).GullbilledTernsbandedintheOld Worldhavestraggledatleastthreetimesabout7,500km(4,700mi) from DenmarktoBarbadosintheWestIndies(Lincoln1936)andhavewanderedoncetoMadeira(Moller1975b).OtherPalearcticpopulationswinterinArabia,thePersianGulf,India,Ceylon,andtheGreaterSundas(Dement'evand Gla,dkov1951,Vaurie1965,BOU1971).MostmigrantsfromDanishpopulationsfollowtheGermanwestcoastsouthandmigrateoverSpainandalongthenorthwestcoastofAfrica;asmallerproportionmigrateoverItalyandacrossthenorthernAfricancontinent(Moller1975b).MigrantsfrompopulationsalongtheeasternMediterraneanarethoughttomigratesouthalongtheRedSea(Moller1975b).Serventyetal.(1971)regardedAustralianGull-billedTernsasnomadic,suggestingthattheydispersetocoastalareasduringthenonbreedingseasonandreturninlandtobreed.Marchant's(1958)observationsinsouthwestEcuadorsuggestthatthismayalsobetrueofSouthAmericanpopulations.342

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Gull-billedTernHABITATNestingInNorthAmericaGull-billedTernsareprimarilycolonialnestersincoastalareaswheretheytypicallynestwithothermarinebirds.IntheOldWorld,theyoftennestwithotherspeciesofgullsandterns(deWaard1950,Lind1963a).InthesoutheasternUnitedStates,Gull-billedTernsfrequentlynestwithBlackSkimmersand/orLeastTerns(Ericksen1926,Weston1933,Nicholson1948a,Hallman1960,Portnoy1977,Sears1978).Gull-billedTernsbreedinginlandattheSaltonSeainCalifornianestedincaked,wind-blowndunes(Pemberton1927),whilethoseatLakeOkeechobee,Florida,placedtheirnestsonthesandyshoresofsmallislands(Nicholson1948a).Gull-billedTernsbreedingalongthenorthernAtlanticcoastusuallynestinmarshesoronsandybarrierbeaches.Those on LongIsland,NewYorkin1975nestedonexposedfillnearmarshland(Buckleyetal.1975).BirdsatthreesaltmarshcoloniesinNewJerseyin1977placedtheirnestsondriftmaterial(BuckleyandMcCaffrey1978).NestinghabitatsusedinthesoutheasternUnitedStatesincludewindrowsofdeadmarshgrass("drift"or"wrack")(Ericksen1926,ParnellandSoots1979ms),grassyislandsinfreshwaterlakes(Sprunt1940,Nicholson1948a),dredgedmaterialislands(Chaneyetal.1978;Kale1979mSa,1979msb;ParnellandSoots1979 ms;Jacksonetal.1980),naturalestuarineislands(ParnellandSoots1979 ms,BlusandStafford1980),barrierbeaches(ParnellandSoots1979ms),andsitesonthecoastalmainland(Oberholser1974).IntheeasternUnitedStates,Gull-billedTernsoftennestondredge-spoilislandsconsistingofsandmixedwithshellsorsilt(Chaneyetal.1978,Sears1978).Theextenttowhichman-madeorman-alteredsitesareusedfornestingvariesconsiderably.Only28%ofthepopulationsinNewJerseystudiedbyBuckleyandMcCaffrey(1978)nestedondredge-spoil,butabouttwo-thirdstofour-fifthsoftheGull-billedTernsnestinginNorthCarolinawerefoundondredged-materialislands(SootsandParnell1975a,ParnellandSoots1979 ms,Portnoyetal.1981).InareasstudiedalongtheTexascoastin1976 and1977,Chaneyetal.(1978)discoveredthat70%and84%,respectively,ofthebreedingGull-billedTernswerenestingondredged-materialislands.Thedegreetowhichthisspeciesusessuchhabitatsmaybe areflectionoftheextentofman-mademodificationofcoastalenvironmentsandtheamountofremainingnaturalnestinghabitat.OnbeachesinNorthCarolinaGull-billedTernsusuallynestinlow-lyingareassubjecttoflooding(e.g.,flatsnearinlets);onspoilislandstheychooseelevatedslopesordomes.Nestingareasaresparselyvegetated;intheearly1970'stheaveragecoverofvegetationwas 14.7%(SootsandParnell1975a)andtheaverageplantheightwasabout12em(4.7in)(ParnellandSoots1979ms).Plantsfrequentlyassociatedwithnestingsitesareseabeachevening-primrose(Oenotherahumifusa),horseweed(Conyzacanadensis),seasidegoldenrod(Solidagosempervirens),camphorweed(Heterothecasubaxillaris),sea_rocket(Cakileedentulaharperi),andsandgrass(Triplasispurpurea)(SootsandParnell1975a).343

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Gull-billedTernInTexasGull-billedTernsnestprimarilyonsparselyvegetatedsandyislandswithcoarsesand.shell.orserpulidreefmaterial.PlantsusuallyassociatedwithnestingareasinTexasarerushgrasses(Sporobolusspp.),sandspurs(Cenchrusspp.).fingergrass(Eustachyspetraea),forbssuchasbluet(Houstonianigricans),seasideevening-primrose(Oenotheradrummondii).andmarshelder(Ivaangustifolia)(Chaneyetal.1978).Gull-billedTernsnestinmarshesmoreoftenintheOldWorldthantheydoinNorthAmerica.BirdsinDenmarknestprimarilyonsalt-marshisletscoveredwithlowgrass(Moller1981a);nestingareasinGermanyareapparentlysimilar(Gloe1974inMoller1981a).MostpopulationsinthesouthernU.S.S.R.nestin"saltmarshes,floodplains,swamps,lakes,andreeded estuarie-s ..."(Borodulina1960).Gull-billedTernsintheU.S.S.R.nestmostlyonplains,butsomealsonestatelevationsasgreatas2km(1.2mi)onmountainlakesinArmenia(Dement'evandGladkov1951).AttheBaned'ArguininMauritaniatheynestinsandyhabitats(deNaurois1959)similartothoseusedinthesoutheasternUnitedStates.NestsitesinAustraliaareallinlandonthemainland(Serventyetal.1971)andincludeoldnestsofBlackSwans (Cygnusatratus),exposeddrymudincane-grassswamps (Hobbs1975),andbare,earthy,orsandybanksalonginlandlakesand ontheislandswithinthem(Bourkeetal.1973).FeedingGull-billedTernsgenerallyfeedoverlandorinlandmarshes.Cain(1933)characterizedonefeedingareaas"scrub."TheyhavebeenseenfeedingoverabrushyfieldinFlorida(Rohwer andWoolfenden1968)andoverasmallfreshwaterpondinSouthCarolina(Chamberlain1960b).Gull-billedTernsinthesouthernU.S.S.R.feedoverwaterandataridinlandlocalitiesincludinghillocksandsandhills(Borodulina1960).InAustraliatheyfeedinbothfreshwaterandincoastalhabitatssuchasseashores,mudflats,lagoons,andestuaries(Serventyetal.1971).Feedingareasmaybeseveralmilesawayfromnestingsites(Chamberlain1960b,RohwerandWoolfenden1968).InEuropethesizeofthefeedingrangedecreasesfromsmall,northerncoloniestolargesouthernones(Moller1982).InDenmark,wherecoloniesaresmall,mostbirdsfeedwithin20km(12mi)ofthecolony,butsomefeedasfarawayas45km(28mi).InaSpanishcolonyofaboutathousandbirds,mostbirdsfeedwithin5km(3mi)ofthecolony,althoughsomefeeduptoabout10km(6mi)away(Moller1982).NonbreedingandOffshoreThereisvirtuallynoinformationonthehabitat,distributionandfeedingecologyofGull-billedTernswinteringinthesoutheasternUnitedStates.Gull-billedTernswinteringinSouthAmericaapparentlyremainalongthecoast.Wetmore(1965)foundbirdsinPanamaonasandspitonthebayshore.InColombiabirdshavebeenseenrestingonasandbarandfeedingoversaltmarsh(Donahue1974).Gull-billedTernsinAustraliaareseldomseenfarfromshore(Serventyetal.1971).ThosewinteringinNorthAfricapreferlagoonsanddeltasshelteredfromthesea,buttheymayalsobeseenoninlandlakesandrivers(Bannerman1962).344

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Gull-billedTernFOODANDFEEDINGBEHAVIORCain(1933)comparedthehuntingbehavioroftheGull-billedTernoverlandtothatofaharrier(Circus),describingitas"skimmingoverthescrub."Gull-billedTernshuntingmiceinAustraliadriftbackwiththewind,thendiveandseizemicewiththebill(Jensen1946).AternflyingaroundtheedgeofamarshypondinSouthCarolinaoccasionallycutacrossthepondormade afigure"8";thebirdthensetitswingsand swoopedtothesurfaceofthepondtopluckaninsectfromthesurface(Chamberlain1960b).Reynolds(1977)sawGullbilledTernsinKenyadipintothewaterforsmallfishfromaheightofabout4m(13ft).SimilarbehaviorhasbeenwitnessedinAustralia(Serventyetal.1971)andelsewhere(Lind1963a).Thisparticularfeedingbehaviorissometimestermed"aerial-dipping"(Clappeta1.1982b)andusuallyreferstobirdsobtainingfoodoverwater.Gull-billedTernsusethismethodoverbothlandandwaterand,judgingfromtheliterature,obtainalargeproportionoftheirfoodthisway. Theymayalsopickupfoodfromthegroundwhilewalkingaroundtheircolonies(Lind1963a).RohwerandWoolfenden(1968)pointedoutthata numberofauthors(e.g.,Bent1921,Ford1961)believedthatGull-billedTernsneverdive,butalsocitedWetmore(1926)andMeinertzhagen(1954)toindicatethatthesebirdsdoenterthewater.Wetmore'sremarksareequivocalasheonlystatedthatGullbilledTernswere"frequentlydivingforsmallfishesinthetidalchannels"inBuenosAires.Meinertzhagenstatedthatthespecies"divesforfood,oftencompletelysubmerged,andinArabiaonlyonsaltwaterbutneverfarfromland."Inanycase,observationsavailabletoussuggestthatthisspeciesseldomplungesintothewaterandobtainsmostofitsfoodonthewing.InDenmark mostGull-billedTernsfeedsolitarilyoringroupsoftwoorthree;flocksof4-10birdsmake up 15-25%of observationsofGull-billedTernsseenfeedinginthebreedingseason(Moller1982).InsouthernFranceflocksof birdsarecommon(authorscitedinMoller1982),andevenlargerflocksmaybeencounteredinsouthernSpain(authorscitedinMoller1982).Gull-billedTernsinVirginiausuallyfeedsolitarilyandarenotseeningroupsofmorethan2-3birds.Fourteenof16observationsoffeedingbirdsinVirginiawereofeither1or2birds(Erwin1978b).Gull-billedTernskleptoparasitizetheirownandotherspeciesofbirds.Reynolds(1977)observedthreeternspursueaGreenshank(Tringanebularia)andtrytoforceittodropafish(Reynolds1977).Hobbs(1976)observedoneGull-billedTernforceanothertodropahousemouse(Musmusculus),whichthefirstternthencaughtinflight.---TheGull-billedTerndiffersfromothercoastalternsinthatitfeedsinshoreandinlandandpreferstoeatinsectsandsmallmammals.Dietvariesbyregionandaccordingtotherelativeabundanceofprey.Fishmake up asmallproportionofthefoodofthisspecies(Dement'evandGladkov1951,Moller1977).345

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Gull-billedTernLittleinformationisavailableonthedietofGull-billedTernsinthesoutheasternUnitedStates.YoungternsareknowntoeatCnemidophoruslizards(SpruntandChamberlain1949) andanadultwasseenseizinggrasshoppersanddragonfliesfromtheedgeofasmallfreshwaterpondinSouthCarolina(Chamberlain1960b).ThestomachsoffivebirdstakeninAlabamacontainedmolecrabs(Emeritatalpoida),dragonflies(Odonata),and agrasshopper(Orthoptera)(Howell1932).Gull-billedTernscollectednearGulfport,Florida,hadfedongreenanoles(Anoliscarolinensis)apparentlypickedfromshrubs(Rohwer andWoolfenden foodsdocumentedaregrasshoppersandfiddlercrabs(Ucasp.).WinteringGull-billedTernsatLakeCharles,Louisiana,havebeen seen feedingoncrayfish(Clement1946).InthesouthernU.S.S.R.Gull-billedTernsfeedprimarilyoninsects,lizards,amphibians,crustaceans,andsmallrodents(Muridae).Theprimaryfoodinoneareawaslizardsandshrimp,inanotherfrogs,andinsectsweremostimportantinotherareas(Borodulina1960andauthorscitedtherein).InsectsarethefoodmostfrequentlyeateninotherareasoftheOld WorldsuchasNorthJutland,Denmark, andtheCamargue,France(Moller1977),butvertebratesmayformmoreofthedietbyweight.SmallmammalsareparticularlyimportantinNorthJutland,Germany(Moller1977)andAustralia(Hobbs1976).FrogsarethemostsignificantitemofdietintheCamargueandlizardspredominateinonepartofAustralia(Hobbs 1976)andarealsoimportantinDenmark(Andersen1945,Moller1977).InsectsfrequentlyeatenintheOld Worldincludebeetles(Coleoptera),grasshoppersandcrickets(Orthoptera),anddragonflies(Odonata)(Leveque1956,Borodulina1960,Moller1977).MammalianpreyitemsfromtheOld Worldincludeshort-tailedvoles(Microtusagrestis)(Moller1977),housemice(Musmusculus)(Hobbs1976,Moller1977),woodmice [=Sylvaemus]sylvaticus),harvestmice(Micromysminutus),commonshrews(Sorexaraneus),pygmyshrews(S.minutus),andwatershrews(Neomysfodiens)(Moller1977).Amphibiansidentifiedinthedietincludefrogs(Ranaridibundus),atoad(Bufocalamita),and anewt(Triturussp.)(Moller1977).LizardseatenincludeLacertaagilis(Borodulina1960,Moller1977),L.vivipara(Moller1977),Eremiasspp.(Borodulina1960), arguta(SchevchenkoinDement'evand Gladkov1951)andPhrynocephalusspp.(SpangenbergandFeigininBorodulina1960).Youngbirdsmade up 14.5%ofthedietinastudyatNorthJutlandandincludedspeciesasdiverseasLapwings(Vanellusvanellus),Redshanks(Tringatotanus),Avocets(Recurvirostraavosetta),CommonandArcticTerns,Skylarks(Alaudaarvensis),MeadowPipits(Anthuspratensis),andYellowWagtails (MOtaCilla flava).Otherspecies or-brrds eatenintheOld WorldincludetheyoungofWhite-wingedLarks(Melanocoryphaleucoptera)(ZarudnyiinDement'evandGladkov1951)andWinterWrens(Troglodytestroglodytes)(Jensen1946).EggsoftheRedshankandthoseofeithertheskylarkorpipitwerealsoeaten(Moller1977).OtherfooditemsreportedfromtheOld Worldincludecrustaceans,earthworms,spiders,andmaggots(Cain1933,Andersen1945,Jensen1946,deWaard1950,Borodulina1960,Rohwer and Woolfenden1968,Moller1977).346

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Gull-billedTernPaperscitedintheterminalbiblographyforthisspecies,especiallyMoller's(1977)comprehensivereviewoffoodhabitsinthePalearctic,providemoreinformationonspecificfoodseaten.IMPORTANTBIOLOGICALPARAMETERSEggLayingIntheeasternUnitedStatesegglayingoccursfromMaytoearlyJuly;peaklayingoccursfrommid-MaythroughearlyJune.Inmoresoutherncolonies(e.g.,inTexas),layingmaybeginasearlyaslateApril.NestsobservedontheSaltonSeainCaliforniawerewelladvancedinincubationon20May(Pemberton1927),suggestingthatlayingisinitiatedinlateApril.NestsinitiatedlaterthanearlyJulyontheAtlanticcoastareprobablytheresultoflossofafirstclutch(Sears1978).LayingperiodsinnorthernportionsoftheOld WorldaresimilartothoseinNorthAmerica.Gull-billedTernsinDenmark,thenorthernendoftheEuropeanbreedingrange,layeggsfromearlyMaythroughearlyJuly;peaklayingoccursfrommid-MaytoearlyJune.Birdsinthe oouthern U.S.S.R.layfrommid-MaythroughearlyJuly(Borodulina1960).BirdsinthePersianGulfbeginlayingasearlyasthefirstweekofApril(Meinertzhagen1954).Gull-billedTernsnestinginAustraliausuallynestinspringand summer(Serventyetal.1971).InwesternAustralia,eggswerepresentinSeptemberatonecolony,butbirdsatanothercolonywerejustbeginningtonestinMay(ServentyandWhittell1976).AtIvanhoe,NewSouthWales,mostegglayinghadbeencompletedbylateDecember(Bourkeetal.1973).Moller(1981a)foundthatbreedingbirdsinDenmarkarrivedatcolonies5-32daysbeforeegglayingbegins;hereportedasignficantnegativecorrelationbetweenarrivaldateandthelengthoftheprelayingperiod(i.e.,theearlierthebirdsarrived,thelongerbeforetheybegantolayeggs).Thelengthofthelayingperiodata numberofcoloniesinDenmarkvariesfrom25to51days(mean38.7).Largercolonieslayforsignificantlylongerperiods(mean43.1days,n=12)thandosmallerones(mean28.3days,n=16)(Moller1981a).Moller(1981a)foundthategglayinginsmallcoloniesbeginslaterthaninlargeones,buthefoundnodifferenceintheinitiationofegglayingbetweensuccessfulandunsuccessfulcolonies.MeanClutchSizeGull-billedTernslay1-4eggs,butclutchesusuallycontainonly2-3(Nicholson1948a;Leveque1956;Borodulina1960;Serventyetal.1971;Moller1975a,1981a).Inmostareas,threeeggsisthemean,butelsewhere(e.g.,thePersianGulf[Meinertzhagen1954),NewSouthWales [Bourkeetal.1973) afullclutchcontainsonlytwoeggs.Asmanyassix(Glegg1925a,Leveque1956,Borodulina1960)orseveneggs(Dement'evand Gladkov 1951)havebeenfoundinonenest,buttheseclutcheswerebelievedtobetheresultoflayingby morethanonefemale.347

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Gull-billedTernInNorthCarolina,Gull-billedTernsthatbegannestingbefore15Mayproducedlargerclutches(mean=2.5,n=16)thanthosebirdsthatlaideggslater(mean=1.7,n=25)(Sears1978).InDenmark,however,clutcheslaidduringthemiddlethirdofthelayingperiodweresignificantlylarger(mean=2.53,n=183)thanthoselaidduringthefirst(mean=2.15,n=107)andlatter(mean=2.17,n=47)thirds,respectively(Moller1975a,1981a).Clutchsizeappearstodecreasefromnorthtosouth(Table39).IncubationPeriodTheincubationperiodforOld WorldGull-billedTernshasbeenreportedas22-23days(DirckseninWitherbyetal.1941,Taning1944,GeroudetinLeveque1956).Thesedataarebasedonlittleinformation.HatchingSuccessTherearefewfiguresavailableontheproportionofGull-billedTerneggsthathatch.Chaneyetal.(1978)reporteda 27.9%hatchingsuccessfor20nestsinTexas.ClutchesfromlargecoloniesinDenmarkthathatchedatleastoneyounghada 96.6%hatchingsuccess;ifunsuccessfulclutchesareincluded,thefigureis90.3%(Moller1981a).AgeatFledgingAccordingtoTaning(1941),youngGull-billedTernsfledge30daysafterhatching.Otherauthors(e.g.,Dement'evand .Gladkov 1951)recordedafledgingperiodof28-35days,butthismaytracebacktoanindefinitestatementthat"Youngcanflyalittleafter4 weeks andwellafter5"(HeinrothinWitherbyetal.1941).FledgingSuccessFewdataareavailableonfledgingsuccessofGullbilledTerns.Moller(198la)reportedthattheoverallreproductivesuccess(averagenumberfledgeddividedbyaverageclutchsize)inDenmark was 69.1%forlargecolonies.Moller(198lb)alsoreportedahigherrecoveryrateforyoungbandedinDanishcolonieswith51-100pairsthanforsmallerorlargercoloniesandequatedthehigherrecoveryratewithgreaterproductionofyoung.ColoniesexaminedbyBlusandStafford(1980)inSouthCarolinawerebelievedtoproducefewyoung;outof28coloniesforwhichtheseauthorsestimatedthenumberofyoungproducedpernest,only4produced0.51-1.00chicks.Mostcolonies(13)produced0.5youngpernestorless,andmany(11)producedno youngatall.MortalityofEggsandYoungLaughingGullsandHerringGullspreyoneggsandchicksofGull-billedTerns(deWaard1950;Sears1978,1979).Sears(1978)thoughtthathumans anddogsarepredatorsorsourcesofdisturbance,and thatstrayingyoungmaybeattackedbyadultsoftheirownorotherspeciesofterns.Sears(1979)emphasizedtheroleofcolonialnestinginthedeterrenceofpredators.Predationbygulls,disturbanceby man, andlossofneststoinclementweatheraretheprimarysourcesofnestingmortalityinthesoutheasternUnitedStates.Hightides,heavyrain,andstormshavecausedtotal(ornearlytotal) atcoloniesinNewJersey(Savell1972),NorthCarolina(Teulings1974d),andFlorida(Ogden1976,1977).Floodingcausedbyheavyrainsandhightides348

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Table39.Gull-billedTernMeanclutchsizesreportedfortheGull-billedTern(a).MeanclutchsizeNumberofclutchesLocalityLatitudeofareaSource2.39152Denmark,NorthernJutland57Moller1975a2.29186Denmark,WesternJutland57Moller1975a2.00(b)Denmark 57Moller1981a2.36(b)Denmark57Moller1981a2.99(c)303Camargue,France43 Leveque19562.06610Andalusia,Spain38Studier-ThierschandStudier-Thiersch19682.041NorthCarolina35Sears19782.3(d)46SouthCarolina33BlusandStafford1980 1.77 30Texas,Laguna Madre 27 Chaneyetal.1978a)Meanclutchsizesgivenherearesometimescalculatedfromdatagiveninthesourcescited.Someofthesefiguresareevidentlycountsofcontentsofneststhatdonottakeintoaccountwhethertheeggswerealllaidbyonebirdorwhetherlayinghadbeencompleted;asaresultthesefiguresmaynotadequatelyrepresentclutchsize.b)Thefirstsetoffiguresisfor16smallcolonies(2-15pairs);thesecondsetisfor12largecolonies.c)Ifnestscontainingfiveorsixeggs(probablyrepresentingmultipleclutches)areexcluded,themeanis2.89(n = 290).d)Thisfigureisimprecisebecauseitisbasedon acountofnestscontainingbotheggsandyoung.causedconsiderablelossofeggsinSouthCarolinacolonies(BlusandStafford1980).ParnellandSoots(1979ms)suggestedthattheprincipalcausesofnestingfailureinNorthCarolinawerethefloodingoflow-lyingcoloniesanddisturbancebypeopleandvehicles.PredatorswereasignificantsourceofnestfailureinSouthCarolina,whereeggsweretakenbothbyLaughingGullsandbyrats(Rattussp.)(BlusandStafford1980);BlusandStaffordbelievedthatno youngwereproducedatDevaux Bankfrom1972to1975becauseofpredationbyLaughingGulls.RatsarealsopredatorsinNorthCarolina(Sears1978).MortalityfromfloodingiscommoninOld WorldcoloniesandisthoughttobethecauseofadeclineinpopulationsnestingontheChernomorskiiReserveinthesouthernU.S.S.R(Borodulina1960).Moller(1978b)statedthatdesertionofDanishcolonieswascausedmostcommonlybydisturbancebyhumans,rats(Rattusnorvegicus)andfoxes(Vulpesvulpes).349

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Gull-billedTernRenestingSeveralpapers(e.g.,SpangenberginDement'evandGladkov1951,ParnellandSoots1979 ms)indicatedthatGull-billedTernsmayrenestafterthelossofafirstnest.However,thedegreeandextenttowhichthisoccursisnotwellknown.Lind(1963a)reportedthatonerepeatclutchwasinitiated8-10daysafterthefirstclutchwasdisturbed.Coloniesthatsufferheavylossesfromtidesorstormssometimesrelocateandrenest(Sears1978).AgeatFirstBreedingGull-billedTernsinDenmarkfirstbreedatanageoffourorfiveyears(Moller1975a).MaximumNaturalLongevityRydzewski(1978)reportedthatabirdbandedasachickwasrecoveredattheageof15yearsand 10months.AbirdbandedasachickinSouthCarolinaandrecoveredinGuyana wasatleast6yearsand4monthsold(Clappetal.1982a).WeightWefoundlittleinformationontheweightsofGull-billedTerns(Table40).TheaverageweightofadultsinNorthAmericaisabout170 g(6oz)(Table40);Old Worldbirdsweighabout210 g(7.4oz)(Borodulina1960).SUSCEPTIBILITYTOOILPOLLUTIONWefoundlittleinformationtosuggestthatGull-billedTernsarefrequentlyaffectedbyoiling.OnebirdseenatLakeCharles,Louisiana,inmidJanuaryhadoil-soakedplumage(Clement1946).Becauseoftheiraerialfeedinghabitsandtendencytofeedinland,Gull-billedTernsprobablywillnotsuffermuchfromthedirecteffectsofoilinthesoutheasternUnitedStates.ThemajorityofNorthAmericanGull-billedTernsbreedinthesoutheasternUnitedStates.Becausethespeciesnestsinareashighlysusceptibletoenvironmentalperturbationsandbecauseitmaybeeasilyaffectedbyhumandisturbance,developmentofcoastalareasmayseriouslyaffectthewell-beingofthislittle-studiedspecies.BIBLIOGRAPHY1982Gloe,P.1982.ZurJugendentwicklung,EthologieundOkologiederLachseeschwalber(Gelochelidonnilotica).Ornithol.Mitt.34:29-40.[InGerman. ]Moller,A.P.1982.ColonialityandcolonystructureinGull-billedTernsGelochelidonnilotica.J.Ornithol.123:41-53.[InGerman.]1981Gloe,P.1981a.Lachseeschwalben(Gelochelidonnilotica)imFlug.Ornithol.Mitt.33:109.[InGerman.] 350

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Gull-billedTernTable40.Weight(ingrams)ofGull-billedTerns(a).MeanweightNumber Range weighedSample andseasonAreaSource176 166 183 150 205 210166-187130-2613129males,Mar.-Maymale,Mar.male,Oct.(b)malemalesmalesMexicoBritishHondurasHondurasFloridasouthernU.S.S.R.U.S.S.R.LSUcoll.Russell1964LSUcoll.Hartman 1955Borodulina1960Dement'evandGladkov 1951 187 169168-205161-17724females,Mar.-Apr.Mexicofemales,Mar.BritishHondurasLSUcoll.Russell1964213172-260femalessouthernU.S.S.R.Borodulina1960 166 213150-190 195-233118adultmalesandTexasfemales,Jun.-Aug.fledglings(Aug.)southernU.S.S.R.Zusi1962;Zusi,pers.comm.Borodulina1960 2429.327.5-32.321recentlyhatchedyoungfreshorslightlyincubatedeggssouthernU.S.S.R.U.S.S.R.Borodulina1960Dement'evandGladkov1951(a)Weightsabove100 gareroundedtothenearestgram.WeightsofbirdsfromtheOld Worldarethoseofthenominaterace,Sternan.nilotica.ThosebirdsfromtheNewWorldareprobablylargelytheeasternS.n.aranea,butthosefromMexicomightbethewesternS.n. BirdsfromBritishHonduraswereidentifiedasaranea by Russellandwereconsideredratherfat.------(b)Thisisprobablythebirdidentifiedby Monroe(1968)asG.n.aranea.351

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Gull-billedTernGloe,P.1981b.ZurfeldornithologischenUnterscheidungvonLachseeschwalbe(Gelochelidonnilotica)undBrandseeschwalbe(Sternasandvicensis).Ornithol.Mitt.33:322324.[InGerman.]Moller,A.P.1981a.BreedingcycleoftheGull-billedTernGelochelidonniloticaGmel.,especiallyinrelationtocolonysize.Ardea69:193-198.1981b.Enanalyseafringmaerknings-oggenmeldingsoplysingerforSandterneGelochelidonnilotica(Gmel.).[AnanalysisofringingandrecoveryinformationfortheGull-billedTernGelochelidonnilotica(Gmel.).]Dan.Ornithol.Foren.Tidsskr.75:127-130.[InDanishwithEnglishsummary.]Sears,H.B.1981.ThedisplaybehavioroftheGull-billedTern.J.FieldOrnithol.52:191-209.1980Blus,L.J.andC.J.Stafford.1980.BreedingbiologyandrelationofpollutantstoBlackSkimmers andGull-billedTernsinSouthCarolina.U.S.FishWildl.ServoSpec.Sci.Rep.Wildl.230. 18pp.Gloe,P.1980.ZurVariabilitatdesDunenkleidesderLachseeschwalbe(Gelochelidonnilotica).Ornithol.Mitt.32:227-228.[InGerman.]Moller,A.P.1980.FlokstorrelsehosSandternenGelochelidonnilotica.[FlocksizeintheGull-billedTernGelochelidonnilotica.]Dan.Ornithol.Foren.Tidsskr.74:152-153.[InDanishwithEnglishsummary.]1979Gloe,P.1979.VegetabileNahrungsrestebeiLachseeschwalben(GelochelidonniloticaGmel.).Vogelwelt100:107-111.Sears,H.F.1979.Colonialnestingasananti-predatoradaptationintheGull-billedTern.Auk96:202-203.1978Davis,A.H.andK.E.Vinicombe.1978.FieldidentificationofGull-billedTerns.Brit.Birds71:466-468.Erwin,R.M.1978b.Colonialityinterns:theroleofsocialfeeding.Condor80:211-215.Gloe,P.1978a.UberdasNestderLachseeschwalbe(Gelochelidonn.nilotica).[RegardingthenestoftheGull-billedTern(Gelochelidonn.nilotica).]Ornithol.Mitt.30:91-95.[InGerman.] 352

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Gull-billedTernGloe,P.1978b.AggressionenauslosendeSituationenanBrutplatzenderLachseeschwalbe(Gelochelidonn.nilotica).Ornithol.Mitt.30:107-115.[InGerman.] 1978c.BeschadigteGrossbeutevonLachseeschwalben-Familien(Gelochel-----idonniloticaGmel.).Corax 6:44-46.Gloe,P.andA.P.Moller.1978.Der ZugnordeuropaischerLachseeschwalben(Gelochelidonn.nilotica)inNord-,Nordwest-undMitteleuropa.[Themigrationof north EuropeanGull-billedTerns(Gelochelidonn.nilotica)inNorth,NorthwestandMiddleEurope.]Ornithol.Mitt.30:-185-202.[InGermanwithEnglishsummary.]Grant,G.S.1978.Foot-wettingandbelly-soakingbyincubatingGull-billedTernsandBlackSkimmers.J.BombayNat.Hist.Soc.75:148-152.Grant,P.J.1978.UpperwingpatternofadultGull-billedand SandwichTerns.Brit.Birds71:468-469.King,K.A.,E. L.FlickingerandH.H.Hildebrand.1978.pesticideresiduesinTexasaquaticbirdeggs,1970.16-21.Shell-thinningandPestic.Monit.J.12:Moller,A.P.1978a.SandternensGelochelidonniloticaadfaerdoverforRovfugle.[AggressivebehaviourofGullbilledTernsGelochelidonniloticatowardsKestrelsFalcotinnunculus.]Dan.Ornithol.Foren.Tidsskr.72: 60.[InDanishwithEnglishsummary.]1978b.SkiftendekoloniplaceringerhosdanskeSandternerGelochelidonn.niloticaGmel.[DesertingflightsintheGull-billedTernGelochelidonn.niloticaGmel.,withspecialreferencetotheDanishpopulation.]Dan.Ornithol.Foren.Tidsskr.72:119-126.[InDanishwithEnglishsummary.]1978c.SandternensGelochelidonn.niloticaGmel.fourageringsomraderiyngletiden.[Feedingareasofthe-Gull-billedTernGelochelidonn.niloticaGmel.duringthebreedingseason.]Dan.Ornithol.Foren. Tidsskr: 72:145-147.[InDanishwithEnglishsummary.]Sears,H.F.1978.NestingbehavioroftheGull-billedTern.Bird-Banding49:1-16.Spaans,A. L.1978.StatusofternsalongtheSurinamcoast.Bird-Banding49:66-76.Therres,G.D.,J.S.WeskeandM.A.Byrd.1978.BreedingstatusofRoyalTern,Gull-billedTern,andBlackSkimmerinMaryland.Md.Birdlife34: 75-77 Vargas,J.M.,A.Antunez andM.Blasco.1978. ComportamientoreproductiveyalimentariodelaPagazapiconegra(GelochelidonniloticaL.)en.laLaguna deFuentePiedradeMalaga.Ardeola24: 227231.353

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Gull-billedTern 1977Gloe,P.1977a.SchlafplatzderLachseeschwalbe,Gelochelidonniloticanilotica(Gmel.)anderWestkusteSchleswig-Holsteins.Ornithol.Mitt.29:107-112.[InGerman.] 1977b.VomBrutplatzabwanderndeLachseeschalben-Familien(Gelocheli-----donnilotica)vorEinsetzendesWegzuges.Ornithol.Mitt.29:231 237. [Tri" German.JJohnstone,R. E. 1977.AnAsianGull-billedTerninWesternAustralia.West.Austr.Nat.14:26.Moller,A.P.1977.SandternensGelochelidonn.niloticaGmel.fodeiyngletideniNordjyllandog Camargue,Frankrig,-medenoversigtoverfodeemineriandredeleafartensudbredelsesomrade.[FoodcompositionoftheGullbilledTernGelochelidonn.niloticaGmel.duringbreedinginNorthJutland,Denmark, and Camargue,France,withareviewoffooditemsinotherareas.]Dan.Ornithol.Foren.Tidsskr71:103-111.[InDanishwithEnglishsummary.]Reynolds,J.F.1977.Attemptedfood-piracybyGull-billedTerns.Brit.Birds70:392-393.Schlenker,R.1977.DeutschenBucht.ZugderLachseeschwalbe(Gelochelidonnilotica)inderOrnithol.Mitt.29:51.1976Gloe,P.1976.Nahrung undZugderLachseeschwalbe(Gelochelidonn.nilotica)imWattenmeer.Ornithol.Mitt.28:117-123.[InGerman.] Hobbs,J.N.1976.Birdsfeedingonhousemice.Emu76:219-220.Sears,H.F.1976.BreedingbehavioroftheGull-billedTern.Ph.D.thesis,Univ.NorthCarolina/ChapelHill,NC.1975Beretzk,P.andA. Keve.1975.DieLachseeschwalbe,Gelochelidonnilotica(Gm.1789),inUngarn.Beitr.Vogelkd.21:245-250.Buckley,P.A.,F.G.BuckleyandM.Gochfeld.1975.Gull-billedTern:NewYorkState'snewestbreedingspecies.Kingbird25:178-183.Hobbs,J.N.1975.NestsoftheGull-billedTern.Austr.BirdWatcher6:117.354

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Gull-billedTernMoller,A.P.1975a.SandternensGelochelidonn.niloticaGm.ynglebiologii Danmark. [Thebreedingbiologyofthe TernGelochelidonn.niloticaGm.inDenmark.] Dan.Ornithol.Foren.Tidsskr.69:9-18. [In-DaTI= ishwithEnglishsummary.]1975b.EuropaeiskeSandternens(Gelochelidonniloticanilotica)traek-----belystvedhjaelpafringmaerking.[MigrationofEuropeanGullbilledTerns(Gelochelidonniloticanilotica)accordingtorecoveries.]Dan.Fugle27:61-77.[InDanishwithEnglishsummary.]1975c.SandternensGelochelidonn.niloticaGmel.bestandsaendringeri Danmark oganalyseafnoglebestandsregulerendefaktorer.[PopulationchangesoftheGull-billedTernGelochelidonn.niloticaGmel.inDenmark,includingananalysisofsomepopulationregulatingfactors.]Dan.Ornithol.Foren.Tidsskr.69:81-88.[InDanishwithEnglishsummary.]1975d.YnglebstandenafSandterneGelochelidonniloticaniloticaGmel. 1972 iEuropa,AfrikaogdetvestligeAsienmedenoversigtoverbestandsaendringeridettearhundrede.[ThebreedingpopulationofGullbilledTernsGelochelidonniloticaniloticaGmel.in1972inEurope,AfricaandWesternAsia,withareviewoffluctuationsduringthepresentcentury.]Dan.Ornithol.Foren.Tidsskr.69:1-8.[InDanishwithEnglishsummary.]1975e.Sandternen(Gelochelidonniloticanilotica)somynglefugli-----Danmark.Enoversightoverdeenkeltekolonier.[TheGull-billedTern(Gelochelidonniloticanilotica)asabreedingbirdinDenmark. Asurveyofthecolonies.]Dan.Fugle27:33-43.[InDanishwithEnglishsummary.]1974 Donahue,P.K.1974.Gull-billedTerninCaribbeanSouthAmerica.Auk91:845.Filonov,K.P.,V.I.Lysenko andV.D.Siokhin.1974.[PeculiaritiesofLimicolaeandLarinestingonislandsofMolochnyEstuary(theSeaofAzov).]Vestn.Zool.1974:52-58.[InRussianwithEnglishsummary.]Gloe,P.1974.DieLachseeschwalbe(Gelochelidonnilotica)imDithmarschen.Ornithol.Mitt.26:47-51.[InGerman.]Lindner,H.1974.Gelochelidonnilotica(Gm.) andLarusmelanocephalusTemm.beiderInselLibitz/Rugen(NSG).Beitr.Vogelkd.20:347.1973Bourke,P.A.,V.T.Loweand T.G.Lowe.1973.NotesontheGull-billedTern.Austr.BirdWatcher5:69.Fien,I.1973.FeedingbehaviorofGull-billedTerns.Sunbird4:57.355

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Gull-billedTernLatour,M.1973.Nidificationdecinqespecesdelaridsau v01s1nage del'embouchuredufleuveSenegal.OiseauRev.Fr.Ornithol.43:89-96.[InFrench.]Petrovic,C.A.andJ.King,Jr.1973.BirdrecordsfromtheDryTortugas.Fla.FieldNat.1:5-8.1972 Anon. 1972. TheGull-billedTern--anewspecies(InACT).CanberraBirdNotes 2:10.Panin,V.Ya. andD.Zhartkanbaeva.1972.[Dicrocoeliumbykhowskajaesp.n.-a newspeciesofdicrocoelidfromGelochelidonnilotica.]Izv.Akad. Nauk Kaz.SSRSer.BioI.1972:46-47. [tn RussianwithKazakh summary.]Savell, 1972.Gull-billedTernsinNewJersey.Cassinia53:47-48.1971Rettig,K.hend.1971.Lachseeschwalben(Gelochelidonnilotica)bei durchzieVoge1kd.Ber.Neiders.3:26.[InGerman.]Schulter,G.C.M.1971.Gull-billedTern,Gelochelidonnilotica.Ostrich42:138.1970Tait,C.S.1970.Gull-billedTernsinStirlingshire.Scott.Birds6:45-46.Wiatr,B.1970.Obserwacjerybitwykrotkodziobej,Gelochelidonnilotica(Gm.)nadjezBukowo.[NewrecordsoftheGull-billedTern,Gelochelidonnilotica(Gm.)inPoland.]ActaOrnithol.12:46-47.[InPolishwithEnglish summary.] 1969Axelsson,H.1969.SandtarnaGelochelidonniloticaochtall-sparvEmberizaleucocephalai Varmland 1967. VarFagelvarld28:46.[InSwedish.]Bates,A.K.andM.L.Bierly.1969. White-rumpedSandpipers,NorthernPhalaropes,RuddyTurnstones,Gull-billedTerns,andStiltSandpipersatMarionFishHatchery.Ala.Birdlife17:52-54.Campbell,N.A.1969.Gull-billedTern(?)atLakeMacIlwaine.Honeyguide59:30.MacDonald,A.1969.Gull-billedTerninEastLothian.Scott.Birds5:284285. 356

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Gull-billedTernManfeldt,E. 1969.Lachseeschwalben-BeobachtungenimostlichenSchleswigHolstein.Corax3:40-41.[InGerman.] Manson, A.J.and C. Manson. 1969.Gull-billedTern--afurthersighting.Honeyguide 59:31.1968Armistead,H. T. 1968.Gull-billedTernsatBlackwater.Md.Birdlife24:94-95.Rohwer, S.A.andG.E.Woolfenden.1968.ThevarieddietoftheGull-billedTernincludesashrub-inhabitinglizard.WilsonBull.80:330-331.Schoennagel,E.1968.Schmarotzerraubmowen(Stercorariusparasiticus)belastigenLachseeschwalben(Gelochelidonnilotica).Ornithol.Mitt.20:4445.[InGerman.]Studer-Tiersch,A. andP.Studer-Tiersch.1968.Sobresavesaquaticusen unalagunadelinteriordeAndalucia(observacionesdemarzo ajuniode1967).Ardeola14:166-174.vanVegten,J.A. 1968.Gull-billedTern on SaoMiguel,Azores.Ardea 56:196.1967 Smout, T. C. 1967.Gull-billedTerninWestLothian.Scott.Birds4:448. 1966Nebelsiek,U.1966.DasSchicksalderFlussseeschwalbe(Sternahirundo)undderLachseeschwalbe(Gelochelidonnilotica)alsBrutvogelBayerns.Anz.Ornithol.Ges.Bayern7:823-846.[InGermanwithEnglishsummary.]Schlenker,R.1966.UberdasVorkommenderLachseeschwalbe(GelochelidonniloticaGmel.)anderWestkusteSchleswig-Holsteins.Corax1:209-216.[InGerman.] 1965Allen,C.R.K.1965. AsightrecordofGull-billedTernsinNovaScotia.Can.Field-Nat.79:211-212.1963 Ahmad,N.-u.-D.1963.AdditiontotherecordedavifaunaofLahore.Pak.J.Sci.15:143-144.357

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Gull-billedTernLind,H.1963a.ThereproductivebehaviouroftheGull-billedTern,SternaniloticaGmelin.Vidensk.Medd. Dan.Naturhist.Foren.125: 1963b.Noglesocialereaktionerhasterner.[Notesonsocialbehav 10ur interns.]Dan.Ornithol.Foren.Tidsskr.57:155-175.[InDanishwithEnglishsummary.]Witkowski,J.ica(Gm.)ica(Gm.) swnmary. ]1963.Wystempowanierybitwykrotkodziobej,GelochelidonnilotwPolsce.[OccurrenceofGull-billedTern,GelochelidonnilotinPoland.]NotatkiOrnitol.4:5.[InPolishwithEnglish1962 Bannerman, D.A.1962. ThebirdsoftheBritishIsles.Vol.XI.Oliverand Boyd, London. 368pp.Forssell,S.G.1962.Sandtarnor(Gelochelidonnilotica)iakttagnanaraGoteborg.[Gull-billedTerns(Gelochelidonnilotica)observednearGothenburg.]VarFagelvarld21:307-308.[InSwedishwithEnglishsummary.]Temme,M.ica).1962.BeitragzurIrnahrungderLachseeschwalbe(GelochelidonnilotVogelwelt83:154-155.[InGerman.] 1961Ford,J.ia.1961.AdditionalrecordsoftheGull-billedTerninWesternAustralWest.Austr.Nat.7:208.Konig,D.1961.Lachseeschwalbe(Gelochelidonnilotica)1960BrutvogelinSchleswig-Holstein.Vogelwelt82:1-6.[InGerman.] Zimmerman,H.1961.LachseeschwalbenundSabelschnableraufder"GrunenInsel"/Schleswig-Holstein.Ornithol.Mitt.13:21-25.[InGerman.] 1960Chamberlain,B. R.1960b.Gull-billedTerntelefeeding.Chat24:98-99.Hallman,R. C.1960.County,Florida.Gull-billedTern(Gelochelidonnilotica)nestinginGulfFla.Nat.33:224. Smout, T. C.1960.Gull-billedTerninEastLothian.Scott.Birds1:335-336.1959 deNaurois,R. 1959.Premieresrecherchessurl'avifaunedesilesduBaned'Arguin(Mauritanie).Alauda 27:241-308.[InFrench.]358

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Gull-billedTern1958Abdulali,H.1958.TheidentityoftheGullbilledTerns[Gelochelidonnilotica(Gmelin)]inIndia.J.BombayNat.Hist.Soc.55:169-170.Marchant,S.1958.ThebirdsoftheSantaElenaPeninsula,S.W.Ecuador.Ibis100:349-387.1957Jakab,A.1957.AkacagoeserfeszkeleseesalocserelofordulasaFonyodon.[AppearanceoftheGull-billedTernandCaspianTernatFonyod(LakeBalaton).]Aquila63/64:284,342.[InHungarianandEnglish.]1956Edfelt,A.1956.Sandtarna(Gelochelidonnilotica)observadvidSkalderviken.[Gull-billedTern(Gelochelidonnilotica)observedinSkalderviken.]VarFagelvarld15:284,287.Konig,D.1956.BeobachtungenanderLachseeschwalbe,Gelochelidonnilotica(Gmel.),nachderBrutzeit.Ornithol.Mitt.8:143-147.[InGerman.]Kumerloeve,H.1956.amNeusiedlersee.ZumVorkommenderLachseeschwalbeinDeutschlandundFalke3:49-51.Leveque,R.1956.DnecoloniedeSterneshanselenCamargue.NosOiseaux23:233-246.[InFrench.]Markgren,M.1956.Iakttagelseravtarnor,bl.a.sandtarnan(Gelochelidonnilotica),vidSjolundautanforMalmo. VarFagelvarld15:126-127.1955Axell,H. E.1955.Gull-billedTerninKent. Brit. Birds48:511-512.Ferguson-Lees,I.J.1955.Gull-billedTerninSussex.Brit.Birds48:512.Harber,D.D.1955.Gull-billedTernsinSussex.Brit.Birds48:511.McKenzie,H.R.1955.A newbirdforNewZealand.Gull-billedTern(Gelochelidonnilotica)nearInvercargill.Notornis6:163-164.1954Meinertzhagen,R.1954.ThebirdsofArabia.OliverandBoyd, London.xiiiand 624pp.359

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Gull-billedTern1952 Brown, C.F.,J.Sheppard,D.D.HarberandA.R.Mead-Briggs.1952.Gu11billedTernsinSussex.Brit.Birds45:262-263.deWaard,S.1952.landin1949.OntheGull-billedTernsat"DeBeer"nearHook-of-Ho1Brit.Birds45:339-341.Ferguson-Lees,I.J.1952.Studiesofsomespeciesrarelyphotographed.XLIV. TheGull-billedTern.Brit.Birds45:357-358.Harber,D.D.1952.Gull-billedTernsinSussex.Brit.Birds45:371-372.Pyman,G.A.andC.B.Wainwright.1952.ThebreedingoftheGull-billedTerninEssex.Brit.Birds45:337-339.1950 de Waard,S.1950.DriebroedparenvandeLachstern,Ge1oche1idonni10tica(Gm.),op"DeBeer",Rozenburg,in1949.Ardea37:161-167.[InDutchwithEnglishsummary.] Remmert,H.1950.Lachseeschwa1beGe1oche1idonn.ni10tica beiBredstedt(Schleswig-Holstein).Ornitho1. Mitt. 2:151.1949Kooijmans,F.P.J.1949. NiewebroedgevallenvandeLachstern,Gelochelidonnilotica(Gm.)inNederland.[NewbreedingcasesofGelochelidonnilotica(Gm.)intheNetherlands.]Limosa 22:342-346.[InDutchwithEnglishsummary.]1948Nicholson,D.J.1948a.Fresh-waternestingoftheGull-billedTerninFlorida.Auk65:139-140.Sprunt,A.,Jr.1948c.Acorrectionoffirstfresh-waternestingofGull-billedTerninFlorida.Auk65:311-312.1946Clement,R. C. 1946.SomeLouisianaobservations.Auk63:97-99.Jensen,P.V.1946.NogleIagttagelseroverSandternens(Gelochelidonnilotica(Gm.biologi.[ObservationsonthebiologyoftheGull-billedTern,Gelochelidonnilotica(Gm.).]Dan.Ornithol.Foren.Tidsskr.40:80-96.[InDanishwithEnglishsummary.] 360

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Gull-billedTern1945Andersen,J.1945.Sandternens(Gelochelidonnilotica(Gm.))fode.[FoodoftheGull-billedTern(Gelochelidonnilotica(Gm.).]Dan.Ornithol.Foren.Tidsskr.39:198-205.[InDanishwithEnglishsummary.]Haverschmidt,F.1945.EennieuwbroedgevalvandeLachstern,Gelochelidonn.nilotica(Gm.)inNederland.Ardea33:117-125.[InDutch.]1944Taning,A.V.1944.YnglefuglenestraektilogfraTipperne.Dan.Ornithol.Foren.Tidsskr.38:168-217.1941Longstreet,R.J.1941.Gull-billedTernnestinginFlorida.Auk58:96.1940Sprunt,A.,Jr.1940.Gull-billedTernbreedinginFlorida.Auk57:251-252.1939Griscom,L.1939.Gull-billedTerninMassachusetts.Auk56:186.1938Craighill,F.H. 1938.Gull-billedTernsbreedingon PeaIsland.Chat2:43-46.1937Emeis,W.1937.DieLachseeschwalbe(Gelochelidonnilotica(Gm.))BrutvogelaufderNordseeinselAmrum.Ornithol.Monatsber.45:170.[InGerman.]Schumann,H.1937.BeobachtungenamletztendeutschenBrutpaarderLachseeschwalbe.Ornithol.Monatsschr.62:184-186.[InGerman.]1936Lincoln,F.C.1936.Trans-AtlanticflightofGull-billedTern. Auk 53:331.1934vanderMeer,G.1934.NogmaalsSternaGelochelidonKl.WaarnemingenvandelachsterninNederland, Club Ned.Vogelkd.6:103-106.361

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Gull-billedTern1933Cain,W.1933.NestingoftheGull-billedTern(Gelochelidonnilotica).S.Austr.Ornithol.12:28-30.Weston,F.M.1933.Gull-billedTernnestingatPensacola,Florida.Auk50:215-216.Wust,W.1933.EinBrutversuchderLachseeschwalbebeiMunchen.Ornithol.Monatsber.41:147-150.1932Dircksen,R. 1932. DieLachseeschwalbe,1931BrutvogelaufNorderoog.Ornithol.Monatsber.40:133-136.[InGerman.]Emeis,W.1932.HerbstdurchzugderLachseeschwalbe(Gelochelidonnilotica(Gm.))imwestlichenSchleswig.Vogelzug3:92-94.[InGerman.] 1931Strijbos,J.P.Nederland.1931.Delachstern,SternaGelochelidonKl.,broedvogelinOrg. Club Ned. 27.Thijsse,J.p.1931a.Alweereennieuwebroedvogel nilotica).LevendeNat.36:97-98.1931b.Onze nieuwegasten[Sterna(Lachstern)].LevendeNat.36:209------211.vanderMeer,G.1931.Waarnemingen van deLachsternSternaGelochelidonKl.,II.Org. Club Ned.Vogelkd.3:136-144.1930 vanderMeer,G.1930. WaarnemingenvandeLachstern,SternaGelochelidonKl.,inNederland,I.Org.Club Ned.Vogelkd.3:70-82.1929Bancroft,G.1929.A newPacificraceofGull-billedTern.Trans.San DiegoSoc.Nat.Hist.5:283-286.1928Ball,W.H.1928.TheGull-billedTern(Gelochelidonnilotica)inWashington,D.C.Auk45:367.362

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Gull-billedTernPemberton.J.R. 1927. TheAmericanGull-billedTernbreedinginCalifornia.Condor29:253-258.Ten Kate, C.G.B.1927.Gelochel1don nilotica(Gm.)bijKampen.Ardea16:138-139.deVries,T.G.1926. (Gm.) bijAmsterdam.Ardea15:1.[InDutch.J Ericksen, W.J.1926.Gull-billedTernsbreedingonthecoastofGeorgia.Auk43:533-534.Wetmore,A.1926.ObservationsonthebirdsofArgenrina,Paraguay,Uruguay,andChile.U.S.Natl. Hus. Bull.133.ivand448pp.Blancher,A.1925.SurIeregimealimentaireduGelochelidonn.nilotica(Gm.).[OnthefoodofGelochelidonn.n!loticaam.) l.{ev:Fr.Ornithol.17:298-299.-----Glegg,W.E.1925a.OnthenestingoftheGull-billedTernintheCarnargue.Brir. Birds 18:202-209.1925b.FurthernotesonthenestingoftheFlamingoandGull-billed--TernintheCamargue. Brit. Birds19:145-148.Waite,H.W.1917.ThebreedingofrheGull-billedTern J.BombaySoc.Nat.Mist.25:300-301.AdultGull-billedTerninbreedingplumage.PhotographbyClaytonTaylor.363

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Gull-billedTernsnestinginaBlackSkimmercolony.PhotographbyJ.A.Spendelow.AdultCaspianTerninbreedingplumage.PhotographbyClaytonTaylor.364

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CASPIANTERN(Sternacaspia)[DA:Rovterne,DU:Reuzenstern,FI:Rayska,FR:Sternecaspienne,GE:Raubseeschwalbe,IC:Randerna,IT:Rondinedimaremaggiore,NW:Rovterne,PO:Rybitwawielkodzioba,SP:Charrancaspica,Pagazapiquirroja;SW:Skrantarna]GENERALDISTRIBUTIONNorthAmericaInCanadaCaspianTernsbreedmainlyin4areasintheinterior.OneareaisinthevicinityofGreatSlaveLakeinsouthernMackenzieand LakeAthabascainnortheasternAlberta;asecondextendsfromDore LakeincentralSaskatchewantoLakeWinnipegand LakeWinnipegosisinsouthernManitoba;thethirdisaroundtheGreatLakesandupperSt.LawrenceRiverinsouthernOntario;andthefourthisinsoutheasternQuebec.TheseternsmaybreedlocallyinNewfoundland(Godfrey1966).InthewesternUnitedStatesthisspeciesbreedslocally,mostlyintheinterior.ThesebirdsbreedinWashington,Oregon,Nevada,Utah,andWyoming, andsouthtotheSaltonSea andcoastalCalifornia(AOU1957).IntheeasternUnitedStatesCaspianTernsmainlybreedinthreedisjunctareas.OneareaisontheGreatLakesinnortheasternWisconsin,Michigan,andnorthwesternPennsylvania(AOU1957);thesecondextendsinpatchesalongthesoutheasternAtlanticcoastfromVirginia(AOU1957,Weskeetal.1977)tonorthernBrevardCounty,Florida(Schreiber1978);andthethirdisalongtheGulfcoast,primarilyinsoutheasternLouisianaandcoastalTexas(AOU1957).Duringthelasttwodecades,scatterednestinghasoccurredoccasionallyonthecentralGulfcoastofFlorida(Schreiber1978),andisolatedinstancesofnestinghavealsobeenrecordedatlakesinnorth-centralMinnesota(Warnerand Beimborn1969)andcentralNorthDakota(Hermanetal.1978).TheonlyotherknownnestingareasintheNewWorldareinBajaCalifornia(AOU1957)andalongthenorthwestcoastofMexico (Voous1960).NorthAmericanpopulationswinterprimarilyalongthePacificcoastfromcentralCaliforniatoBajaCaliforniaandtothesoutherncoastofChiapasin western Mexico(AOU1957,Wetmore1965,GillandMewaldt1979),andalongtheAtlanticcoastfromSouthCarolinatoFlorida(SpruntandChamberlain1949,Sprunt1954,Burleigh1958),alongthenorthernGulfcoast,andtosomeextentintheBahamasandGreaterAntilles(AOD1957,Bond1971,Schreiber1978).Theywintercommonly on'thelowerMagdelenaRiverandCaribbeancoastofColombia(Blake1977),butapparentlynotinlargenumbers onthecoastsofCentralAmerica.Taxonomicnote:AlsoappearsfrequentlyintheliteratureasHydroprognecaspiaorHydroprognetschegrava.365

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CaspianTernAlthoughsomemigrationoccursthroughtheinterior,evidentlymosttakesplacealongthecoasts.ThescarcityofrecordsfortheseternsalongthecoastsofCentralAmericasuggestsadirectoverwaterflighttowinteringareasoffnorthernColombia.WorldDistributionThedistributionofOldWorldbreedingpopulationsisalsofragmented.IneasternEurasia,CaspianTernsbreedalongtheshoresoftheBalticinSweden,Finland,andEstonia(Vaurie1965);onthenorthcoastsoftheBlack Sea andSeaofAzov;andontheCaspianandAralSeas(Borodulina1960).InwesternEurasia,theybreedinnorthwesternMongolia,northernSriLanka,onthecoastofChinanorthtoManchuria,andinUssuriland(Vaurie1965).InAfrica,CaspianTernsnestata numberofscatteredlocalities,mostofthemcoastal,includingtheGulfofSuez;Tunisia;theBancd'ArguinoffnorthernMauretania;isletsoffSenegal,Gambia,andPortugueseGuinea;andNigeria(EtchecoparandHue1964,Vaurie1965,Mackworth-PraedandGrant1970).TheyalsobreedinSouthAfricafromtheCapeofGood HopetothemouthoftheZambeziRiver,andintheinteriorofthecontinentatLakeRudolphinKenya(AOU1957,Vaurie1965).InthesouthwestPacific,CaspianTernsbreedonislandsaroundthecoastofAustralia(exceptforNewSouthWales),ontheNorthandSouthIslandsofNewZealand,andinTasmania(Serventyetal.1971,Condon1975).OldWorldpopulationswinterinavarietyofareasdependingontheirnestingareas.Somemaybesedentary,othersnomadic,yetotherstrulymigratory,butadequateinformationisstilllackingformanypopulations.WesternEurasianbirdswinteralongthecoastoftheMediterraneanbuttoagreaterextentoffthecoastofAfrica.Others,probablythosefrommoreeasternpopulations,winteralongtheRedSeatonorthernIndia,inAfrica,andinIndonesiaandsoutheasternChina(Dement'evandGladkov1951,Borodulina1960).ThosefromthecoastofChinamay movesouthtoIndonesia.PopulationsinAustraliaapparentlyaresedentaryandexhibitonlylocalmovements(Serventyetala1971).DISTRIBUTIONANDABUNDANCEINTHECOASTALSOUTHEASTERNUNITEDSTATESNorthCarolinaCaspianTernsareprimarilyuncommonmigrantsinNorthCarolina.MostrecordshaveoccurredalongthecoastfromAprilthroughNovember(Pearsonetal.1942)witharangeofdatesfrom28Aprilto4January(Pearsonetala1942,LeGrand1978).Inland,whereCaspianTernsareregardedasthemostregularlyseenterninspring(LeGrand1979d),thisspecieshasbeenrecordedasearlyas28 March(Teulings1976c).CaspianTernsbreedirregularlyinsmallnumbersalongtheNorthCarolinacoast.Theywerefirstrecordedbreedinginthestateon25June1972,when twonestswerefoundatOregonInlet(ParnellandSoots1976).Inthefollowingthreeyears,1,4,and4nestswerefoundatthislocality,andanotherwasfoundatHatterasInlet.Alltheobservationsmadefrom1973to1975weremadebetween1and13Juneandnestscontainedeithereggsorchicks(Parnelland366

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CaspianTernSoots1976).Breedingpopulationsduringtheensuingtwoyearsconsistedof18birdsbreedingon aspoilislandinshorefromOregonInletin1976(Portnoyetal.1981,Map11),and 16attwolocalitiesonbarrierislandsin1977(Map11).SouthCarolinaCaspianTernsoccurcommonlyinsmallnumbersthroughouttheyearalongtheSouthCarolinacoast(SpruntandChamberlain1949).TheirstatusasabreederinSouthCarolinaispoorlyknown.SpruntandChamberlain(1949)indicatedthatthisspeciesbredinsmallnumbersamongRoyalTernsintheCape Romainarea,butBurton(1970)statedthatthisternis"apermanentresidentthatdoesnotbreed",addingthathehadseenatleast100,000RoyalTernnestsinSouthCarolinabuthadneverfoundaCaspianTernnestamongthem.Thefirstdocumentednestinginthestateoccurredon 5June1970when anestwithtwoeggswasfoundatCape Romain NWR (McDanielandBeckett1971).Theeggswerestillpresenton 2Julyand achickabout3-weeksoldwascaught,banded,andphotographedon25July.NonewerefoundbreedinginSouthCarolinain1976(Portnoyetal.1981).GeorgiaCaspianTernsarecommonthroughoutandareoccasionallyseenintheinteriorinlate1977).Theyarenotknowntobreedinthestate.usuallyno morethanoneortwobirdsareseenattheyearontheGeorgiacoastsummer andfall(Dentonetal.Thepopulationissmall,and atime(Burleigh1958).Florida-AtlanticCoastCaspianTernsarepresentinsmallnumbers ontheFloridaAtlanticcoastthroughouttheyear,butfewbreedinthisarea.Kale(1979msa)reporteda maximumof31pairsnestingondredge-spoilislandsatMerrittIslandNWR(Map11),theonlybreedingsiteonthiscoast.SurveysoftheFloridaAtlanticcoastin1976foundnonestingCaspianTerns(Portnoyetal.1981).CaspianTernsseeminglyhavebeennestinginFloridamorefrequentlyinrecentyears.Schreiber(1978)suggestedthisistheresultoftheincreasednestinghabitatmadeavailableby man-madespoilislands.Florida KeysCaspianTernsareprimarilywintervisitors(Map12)intheFloridaKeys.Theyareseenfromearlyfallthroughmid-spring(HundleyandHames1960-62).Florida-GulfCoastKale(1979msb)brieflysummarizedCaspianTernnestingalongtheFloridaGulfcoast.Thefirstnestingpairwasfoundin1961 on adredge-spoilislandinBocaCiegaBay.By1978,birdswerenestingondredge-spoilislandsatthreedifferentlocalities:HillsboroughBay(30pairs);Eastport,FranklinCounty(30pairs);andGasparillaSound (1pair).AlongtheGulfcoast,eggshavebeenrecordedfromasearlyas5May(Dunstanetal.1975)to11July(Stevenson1979b),andyounghavebeenrecordedfrom27May(Dunstanetal.1975)to15July(WoolfendenandMeyerriecks1963).Alabama AlongtheAlabamacoastCaspianTernsarecommoninwinterand uncommontorareinsummer(Imhof1976b).AccordingtoImhof(1976b),theyaremorecommoninbrackishorfreshwaterhabitatsandthusreachpeakabundanceinareassuchasMississippiSoundandtheheadofMobileBay.Weknow 367

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BIRDNAME'------------------,\ ..
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CASPIANTERNBIRO NAME'860 ---,\ :Hap 12.-------GULFOFMEXICO "'" I ) v-.-..)r-,J .-I. ),-,r-? J,, ,,I'r ,sX' A DALLASEWinterDistributionMapforSoutheasternUnitedStatesBIRDSPER10PARTY-HOURSIILessthanone 1::i!III:I[I(::iil[IJ 1-5 tcoc"=j 5-20 --'r-] Marethan20(AdoptedfromIyltralr,1974)INDIVIDUALSOBSERVEDDURINGCHRISTMASBIRDCOUNTS,1973-1977(ARITHMETICMEAN) @ Numberof individuals 8 Lessthanoneindividual None observed T {.:.....--t;." ,"#d' I J i \. i O----+----jJ---i--o/l-0,.

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CaspianTernofonenestingrecordforthestate.WhileconductinganaerialsurveyofwaterbirdsalongthenorthernGulfCoastduringthesummerof1976,Portnoy(1977)foundacolonyof132birdsnestingonaspoilislandjustnorthofDauphinIsland.Portnoy(pers.comm.) sawthesebirdsfroma lowaltitudeandhisobservationsofthenestspacing and thebrightredbillsoftheadultsconvincedhimthattheidentificationwascorrect.MississippiBurleigh(1958)regardedthisspeciesasfairlycommonalongthecoast,wheresmallflockscanbeseenalmost daily. Numbersaregreatestinfallandwinterwhenlocalpopulationsareaugmentedbymigrantsfromthenorth.InMississippi,asina numberofothercoastalsoutheasternstates,nestingCaspianTernshavebeenfoundonlyrecently.Weknowoffourrecordsofnesting,fewofwhichweredocumentedinanydetail.Thefirstrecordoccurredin1966when 5nestswerefoundon HornIsland(Portnoy1977).Fifteennestswerefoundthereineachoftwosubsequentyears(Portnoy1977).InJune1976theynestedonaspoilislandinHornIslandPass(Jacksonetal.1979msa).LouisianaCaspianTernsarecommoninLouisianabothinlandandalongthecoast.AsinAlabama,thisspeciestendstopreferfreshor brackish waterandismorecommoninmarshyareasthanalongbeaches(Lowery1974).Lowery(1974)indicatedthatsizablebreedingcolonieswerefoundalongthecoast,butrecentsurveys(Map11)suggestthatonlysmall-numbersbreedwithinthestate.TexasTheseternsoccuralongtheTexascoastthroughouttheyearandasubstantialmajorityoftheCaspianTernsbreedinginthesoutheasternUnitedStatesnestinthisarea(Map11,Table41).Indeed,recentdatasuggestthatseveraloftheTexascoloniescontainmorebreedingCaspianTernsthanalloftherestofthesoutheasternstatescombined.ThebreedingseasoninTexasextendsfrommid-Marchtomid-July.Eggshavebeenrecordedfrom25Marchthrough18June(Oberholser1974).BreedingpopulationsalongtheTexascoastin1977 and 1978contained1,088and1,004pairs,respectively(Blacklocketal.1978ms).Thisrepresentsnochangefromtheprecedingfouryears.Countsfrom1973to1976revealedthatmostofthenestingpopulationoccursalongthenorthandcentralcoasts(Map11).Duringthesefouryearsthenorth,central,andsouthcoastscontained17.3-59.7%,40.3-78.6%,and0.0-9.9%,respectively,ofthepopulationbreedinginTexas.SYNOPSISOFPRESENTDISTRIBUTIONANDBREEDINGBreedingCaspianTernsarebirdsofextremelywidespreadbutpatchydistributionthatarelargelyassociatedwithtemperateandsubtropicalcontinentallandmasses.TheyareabsentasbreedingbirdsonlyfromoceanicandpolarareasandfromSouthAmerica.Theybreedfromabout65 NinEuropetoabout45 SinNewZealand,butmostofthebreedingpopulationsareconcentratedin370

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CaspianTernTable41.RecentestimatesofCaspianTernpopulationsnestinginthesoutheasternUnitedStates.StateNorthCarolinaSouthCarolinaGeorgiaFlorida-AtlanticCoastFlorida-KeysFlorida-GulfCoastAlabamaMississippiLouisianaTexasNumberfoundbreedingin1976 18 none none none nonenone(a)13242822,6403,076Percentofsoutheasternbreedingpopulation0.64.30.19.285.8Maximumnumberfoundbreedinginrecentyearsandyear18(1976)2(1972)none62(1975)none62(1978)132(1976)30(1968)ca.1,000(1967)2,640(1976)(a)Precisedatafor1976arelackingbutwesuspectthatatleastsomenestingoccurredalongtheFloridacoast.Thisspeciestendstonestinisolatedpairsorinsmallcoloniesamongothercoloniallarids,andoftennestsatwidelyscatteredlocalities.Inconsequence,nestsmaybeoverlookedonsurveysandthespeciesmaybreedmorewidelyinsomeareas(e.g.,Florida)thanpresentdataindicate.theHolarcticbetween200and 55N.TheCaspianTernbreedsinthesoutheasternUnitedStates,butthroughoutmuchoftheareaitisarareandirregularbreeder.BreedingpopulationsofathousandormoreformerlycouldbefoundinLouisiana(Portnoy1977),butatthepresenttimesubstantialnestingpopulationsarefoundonlyinTexas.RecentestimatesforthesoutheasternUnitedStatesaregiveninTable41.InNorthAmerica,mostCaspianTerns(about7,500birds)nestintheGreatLakesareaofCanadaandtheUnitedStates.Anestimated1,620breedingpairswerepresentin1964(Ludwig1965a).In1978,about3,740pairsnestedthere,about1,650ofthemonislandsinLakeMichiganwithinU.S.waters(Shugartetal.1979).Thelargest'U.S.coloniesintheGreatLakeswerethoseatGravellyIslandand ShoeIslandwith580 and 350pairs,respectively.SomeoftheotherlargecoloniesofCaspianTernsrecentlyrecordedelsewhereinNorthAmericawereoneswithabout1,240pairsatWhitcombIsland,GraysHarborin1976;1,000pairsatGooseIslandin1973;600pairsatSandIsland,GraysHarborand 400pairsinWallapaBayin1976(allinWashington)(Penland1976a),andonewith500-600pairsonBairIslandinSanFranciscoBay,California,in1975371

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CaspianTern(Gill1977).Forcomparison,thelargestcolonyinrecentyearsinthesoutheasternUnitedStateswasoneinNeucesBay,Texas,thatcontainedabout623pairsin1976(Blacklocketal.1978).ThisTexascolonyheldmorebreedingCaspianTernsthanwerefoundinalltheothersoutheasternstatescombinedandwasthelargestbreedingcolonyintheeasternUnitedStates.TheCaspianTernisalmostunknownasabreederontheU.S.AtlanticcoastnorthofNorthCarolina.OnlyasinglenestingpairinVirginiawasfoundinthisareain1977(Erwin1979a).InnorthernEuropeCaspianTernsnestonlyalongtheBalticSeawithabreedingpopulationestimatedatabout2,300pairs.Nearly1,000ofthesebreedinSweden(Staav1979),andabout1,000pairsbreedalongtheBalticcoastofFinland(Vaisanen1973).RecentdataforcoloniesfrommostoftherestoftheOld Worldarescantorlacking.Dement'evandGladkov(1951)remarkedthatCaspianTernsarecommonontheCaspianSeaandlocallycommonontheAralSeaandineasternKazakhstan.TheyareapparentlyuncommoninthesouthernU.S.S.R.(Borodulina1960).WinterWinteringCaspianTernsinthenorthernportionoftheOld WorldarefoundmostlyoffthecoastofAfrica,fromtheRedSeatonorthernIndia,andinIndonesiaandsoutheasternChina.ThoseinAustraliaevidentlywinternearthebreedingareas.ThosethatbreedinNorthAmericawinterpredominantlyalongthePacificcoastofMexico,ontheAtlanticcoastofFlorida,alongtheGulfcoastfromFloridatoCentralAmerica,andintheCaribbeantonorthwesternSouthAmerica.CaspianTernsfledgingintheGreatLakeswinterpredominantlyinthesoutheasternUnitedStates,theeasternCaribbean,andColombia(Ludwig1965a).ThosefledgingfromcoloniesinSanFranciscoBay,California,winteralongthewestcoastofMexico(GillandMewaldt1979).Thestatusofwinteringpopulationsispoorlyknown.JudgingfromrecentChristmasBirdCounts(Map12),thelargestwinteringpopulationsinthesoutheasternUnitedStatesarefoundonthenorthandcentralAtlanticcoastofFlorida,onFloridaBayinsouthernFlorida,andontheTexascoast.MigrationInformationonthemigrationofCaspianTernsintheOld WorldislimitedandismostcompleteforpopulationswinteringinnorthwestEurope.Detailsofmigrationarewellknownforonlya fewNorthAmericanpopulations.Ludwig(1965a)believedthatadultandjuvenileCaspianTernsmigratingfromtheGreatLakesprimarilyreachtheirwinteringareastotheeastandsoutheastbyfollowingtheMississippiFlywaytotheGulfcoast;somebirdsmovealongLakesErieandOntariotoPennsylvaniaandNewYork,andthenmovesouthalongtheAtlanticcoast.MostyoungfledgefromthecoloniesintheGreatLakesinJulyandAugustandrapidlyproceedsouth.Ludwig(1965a)reportedthattheaveragerecoverydatesinVirginia,NorthCarolina,andLouisianawere,respectively,17August,372

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CaspianTern11September,and12September.MostoftherecoveriesfromMississippi,Georgia,andFloridawereobtainedinNovember andDecember.Almostathird(32.6%)ofthe132recoveriesobtainedinthesoutheasternUnitedStatescamefromFlorida.Louisianahad 18.9%, Alabama andNorthCarolinahad15.2%each,Mississippihad7.6%,GeorgiaandSouthCarolinahadabout4.9%, andonebirdwasrecoveredinTexas.HABITATNestingCaspianTernstypicallynestonsparselyvegetatedislets.Ludwig (1965a) foundCaspianTerncoloniesintheGreatLakes"onlyonislandswithsand-gravelsubstrateswhichhadsparseornovegetation."PreferrednestinghabitatintheBalticareawasdescribedbyBergman(1980)as"small,low,flat,rockyorgravelisletswithouttreesorbushes,inmostcaseslessthan2hainareaandsituatedinaphysiognomicallymarinelandscape."InAustraliamostnestingoccursonsandybeaches,butothernestsarefoundonshingleorheadlands(Serventyetal.1971).Afterreviewingtheliterature,Penland(1976a)concludedthattheonlycommonfactorinallnestingareaswas alimitedamountofvegetation.CaspianTernsnestsolitarily,orinsmalltolargecolonies. In theBaltic,about75%ofthebreedingsitesareoccupiedbysinglepairs,butabout92%ofthebreedingpopulationisfoundincoloniescontainingfrom9to164pairs(Bergman1980).Nestsincoloniesareoftencloselypackedtogether.Distancesbetween50nestsinonecolonyintheU.S.S.R.rangedfrom80cmto4m(ca.32157in)(Borodulina1960).NestingdensitiesforavarietyofnestinghabitatsatWhitcombIsland,Washington,rangedfrom0.39to1.27nests/sqm(3.62-11.80nests/IOOsqft)(Penland1976a).MostoftheCaspianTerncoloniesfoundalongthecoastofLouisianaandMississippiarefoundonbarrierorspoilislands,butthelargestcolony(150birds),representingoverathirdofthebirdsbreedinginthatarea,wasfoundon ashellberminasaltmarshinSt.BernardParish,Louisiana(Portnoy1977).ColoniesinFloridaarefoundmostlyondredge-spoilislands(Schreiber1978).Oberholser(1974)statedthatCaspianTernsnestinginTexasusuallywerefoundin"lessdevelopedandpollutedsegmentsofthecoast,nestingprincipallyonbarrenspoilislandsorshellbeachesfreeofvegetation."FeedingFeedingareasusedbybreedingCaspianTernsmayvaryduringthecourseofthenestingseason.InFinland,Bergman(1953inSoikelli1973b)noted that theseternsfedintheinnerarchipelagoatthebeginningofthebreedingseason,but fedintheouterarchipelagolaterintheseason.CaspianTernsatElkhornSlough,Monterey,California,prefertoforageoverthemainchannel,andfishtoalesserextentovershallowerwatersneartheshore(Baltzetal.1979).CaspianTernsinAustraliaalsousebothdeepandshallowwatersforfishing(Serventyetal.1971).Dement'evandGladkov(1951)remarkedthatfishwerecapturedinshallowclearwaters.373

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CaspianTernNonbreedingandOffshoreWinteringCaspianTernsusuallyarefoundalongbeaches,spits,andonisolatedislets.Theyoftenroostwithotherlarids,particularlythesimilar-appearingRoyalTern.MigrantCaspianTernsareoneoftheNorthAmericanternsmostlikelytobeseeninlandalongwatercoursesorinlargemarshes.ThedistancethatCaspianTernsforageoffshoreinthesoutheastisnotwellknown. LeeandBooth(1979)indicatedthatthisspecieswas"regularlyobserved20ormoremilesoffshore"duringpelagic'observationsconductedoffNorthCarolina.Ontheotherhand,transientCaspianTernsseeninthenorthernChesapeakeBightrarelyrangedoffshorebeyondthesightofland(Rowlett1980).Oberholser(1974)suggestedthatthisspeciesusuallyfeedswelloffshoreinTexas.FOODANDFEEDINGBEHAVIORWhensearchingforfood,CaspianTernsusuallyflyslowlyoverthewater,sometimesbrieflyhovering,andthenplungetothesurfacetoseizepreywiththeirbill.Theymaysubmergeentirely,butonlyforbriefperiods(Penland1976a).Foragingflightsareusuallylow(Dement'evandGladkov1951),butCaspianTernsmayhoverupto40ft(12m)abovethewaterbeforediving(Serventyetal.1971).Penland(1976a)notedthatCaspianTernsforaginginWashingtonusuallydivefromheightsof5to20 m(16-66ft).Dement'evandGladkov(1951)statedthatthisspecies"frequentlyfeedsonthewater,swim minglikeagull."Dement'evandGladkov(1951)indicatedthatbreedingbirdsusuallyforagedonlyshortdistancesfromthenestingareas.Borodulina(1960)reportedthatbirdsinKazakhstanforaged10-12km(6.2-7.5mi)fromthebreedinggrounds.Bergman(1953inSoikelli1973b)indicated,however,thatCaspianTernsinFinlandmayflyover30km(19mi)fromtheirneststoobtainfood.Tagsfromsmoltsofsalmonandsea-troutfoundattwonestsinFinlandhadbeenplacedonthesefishatlocalities70and85km(43to53mi)distant(Soikelli1973b).SimilardataareavailablefromNorthAmericancolonies.Gill(1976)founda numberoftagsfromrainbowtrout(Salmogairdneri)amongfooditemsrecoveredatacolonyinSanFrancisoBay,California.Elevenofthesehadbeenreleasedinareservoir18mi(29km)away, andanotherinareservoir37 mi(60km) away.Gillsuggestedthatthelocationofthetags(sevenatonenest,fiveatanadjacentnest)wastheresultofonepairlearningaboutaforagingsitefromanotherpair.CaspianTernsarealmostentirelypiscivorous.Thekindoffisheatendependslargelyonwhatspeciesofthepropersizeismostreadilyavailable.Thedietmaychangeduringtheyearasonespeciesoranotherbecomes moreabundant.Forexample,inFinland,roach(Rutilusrutilus)predominatedinthedietearlyinthebreedingseasonwhileClupeidswerefavoredattheendofMay(KoliandSoikkeli1974).Perch(Pereafluviatilis)became moresignificantinJuneandJuly,althoughroachremainedveryimportantinthediet.KoliandSoikkeliconcludedthatCaspianTernsareopportunisticfeedersthatfeedprimarilyon 374

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CaspianTernavailablefishinasizerangebetween8and26cm(3-10in)long.Mostpreytakenwerebetween14and20cm(5-8in)long.Borodulina(1960)indicatedthatfishtakenintheU.S.S.R.areusually15-20cm(6-8in)long,butoccasionallysomeaslongas30cm(11.8in)aretaken.ThedietoftheCaspianTernisrelativelywellknownforseveralareasofNorthAmericaandEurope.KoliandSoikkeli(1974)providedetailedinformationonthespeciesoffisheateninFinlandandSweden,andotheritemsofdietconsumedintheOld Worldaregiveninpaperslistedintheterminalspeciesbibliography.LittlespecificinformationisavailableonthedietofCaspianTernsinthesoutheasternUnitedStates. We summarizebyareabelowsomeofthefoodseatenelsewhereinNorthAmericabecausethismayreflectthistern'sdietinthesoutheast.WashingtonSmithand Mudd (1978)examineddeadfishinacolonyatGray'sHarbor,Washington.Thesefishhadtotallengthsrangingfrom9.8cm(3.9in)to29.6cm(11.7in).Becausethesefishhadnotbeeneaten,thesefiguresmaynotbethebestestimateofthesizesoffishnormallyeaten.Thespeciesfoundmostfrequentlywereshinerperch(Cymatogasteraggregata),chumsalmon(Oncorhynchusketa),andPacificstaghornsculpin(Leptocottusarmatus).CaliforniaGill(1976)conductedastudyoffoodhabitsatacolonyinSanFranciscoBayandfoundthatthemajorfooditemswereestuarinefishthatcouldbecaughtclosetotheternery.Althoughatotalof21speciesoffishwereidentified,threepredominated:jacksmelt(Atherinopsiscaliforniensis),shinerperch,andPacificstaghornsculpin.Thesemade up 33%, 19%,and19%,respectively,ofthe605fishexamined.Baltzetal.(1979)reportedonthefoodhabitsoftheCaspianTernsnestingatElkhornSlough,MontereyCounty,basingtheirreportonthecontentsof10stomachscollectedinJuly.Thesebirdshadfedprimarilyonadultshinerperch(80%of25fishidentified),buthadalsoeatenadultNorthernanchovy(Engraulismordax).Martini(1964inKoliandSoikkeli1974)foundthedietatSanDiegoBaytobesimilar.There,88%ofthedietwas made upofshinerperchandtopsmelt(Atherinopsaffinis).AlbertaAt acolonyonLittleGeorgeIslandinLakeWinnipeg,784regurgitatedpelletsallcontainedfishremains,6.0%containedinsects,3.2%containedterneggshells,and0.3%containedbirdbones(Vermeer1973a).U.S.GreatLakesLudwig(1965a)reportedthatternsnestingonLakesHuronandMichiganin1963and1964fedexclusivelyonfish.Of 169fooditemsidentified,73.7%werealewives(Alosapsuedoharengus)and15.1%wererainbowsmelt(Osmerusmordax);thesamespeciesstillmade upthemajorityofthedietin1977and1978(Shugartetal.1979).Alewifesmade up 57.2%(of1,219fooditems)andsmeltmade up34.0%.Mostfishcapturedwere5-15cm(2-6in)inlength.375

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CaspianTernOccasionally(butrarely)crayfish(Cambarussp.)(Shugartetal.1979)andinsects(Borodulina1960)arealsoeaten.Dement'evandGladkov(1951)statedthatthisternalsooccasionallyfeedsonnestlingsandeggsofotherbirds.Rarely,thisternmayfeedoncarrion.Cunningham(1966)foundaCaspianTernfeedingon adeadsnake(EasternCottonmouth,Agkistrodon k' piscivorus)onaroadnearNine-MilePond,EvergladesNationalPark.IMPORTANTBIOLOGICALPARAMETERSEggLayingInAtlanticcoastcoloniesmosteggsarelaidinMayandearlyJune.AlongtheGulfcoast,mostlayingapparentlyoccursduringAprilandMay.Layingregimeselsewhereintheworldvarywithlocality.CaspianTernsintheGreatLakesregionlaideggsfromearlyMayuptothebeginningofJuly(Ludwig1965a).DuringoneyearatGray'sHarbor,Washington,egglayingbeganinthethirdweekofAprilandcontinuedforthreetofourmore weeks(Penland1976a).InnorthernAland,Finland,layingoccursfromearlyMaytolateJuneorearlyJuly,withyoungbeginningtohatchbylateMayorearlyJune(KoliandSoikelli1974).IncentralAsiamostlayearlyinMaybutatmoresouthernlocalitiesmaylayearlier(e.g.,inearlyAprilinMesopotamia)(Dement'evandGladkov1951).EgglayinginsouthernAustraliaoccursfromSeptembertoDecember,butinnorthernAustraliaactivenestshavebeenrecordedyearround(Serventyetal.1971).MeanClutchSizeCaspianTernsusuallylayfromonetothreeeggs,butasmanyasfourorfiveeggshavebeenfoundinonenest(Soikkeli1973a,Penland1976a).Penland(1976a)presentedsomedataindicatingthatsuchclutchesmaybetheresultofeggsbeingtransferredorstolenfromothernests.Shugartetal.(1979)notedthatclutchsizewastypically1-2eggsinManitoba,California,Texas,andFlorida,andtypically2-3eggsintheGreatLakes,Washington,andnorthernEurope.Serventyetal.(1971)reportedthatCaspianTernsnestinginAustraliausuallylay1-2eggs.FirstclutchesintheGreatLakesaveraged2.81eggs(n=225)in19631964 (Ludwig1965a),and2.4eggs(n=2,170)in1975-1978(Shugartetal.1979).Chaneyetal.(1978)foundanaverageof1.6eggsin10nestsatonecolonyinTexas.Penland(1976a)notedmeanclutchsizesof2.05to2.50atcoloniesinWashington.Clutchsizemaydecreaseasthebreedingseasonprogresses.Soikkeli1973a)notedasteadydecreaseinclutchsizethroughtheseasoninFinlandduring1970-73.The meanclutchsizefromthebeginningoftheseasonto20Maywas2.70.Itwas2.04for21-31May,1.79for1-10June,and1.61forclutcheslaidthereafter.IncubationPeriodTarr(1960)reportedthattheincubationperiodatasinglenestofCaspianTernsinAustraliawas21days.Ludwig(1965a)reportedthattheincubationperiodwasabout26daysinCaspianTernsnestingonthe376

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CaspianTernGreatLakesoftheUnitedStatesandCanada.ThreeeggsatWhitcombIsland,Washingtonwereincubated27days each. Dataforelevenothereggssuggestedarangeofincubationperiodsof26-28days,withmost(9)hatchingat27days(Penland1976a).Hatching Success Hatching success was81%at200nestsintheGreatLakes(Ludwig1965a);81.8%hatchedatonecolonyintheTorontoOuterHarbor,Ontario(HaymesandBlokpoel1978a);and12of16 (75%)hatchedatonecolonyinTexas(Chaneyetal.1978).Soikkeli(1973a)didnot report precisefiguresfor hatch ing success, butreportedthat1.78-1.86 chicks hatchedperpairovera3-yearperiod(1970-72)intheAlandArchipelago,Finland.AgeatFledgingBergman (1953inBorodulina1960)indicatedthatCaspianTernsbegintoflyatabout46-48days,and Ludwig(1965a)reportedthatyoungfledgeat6-8weeks(42-56days).Soikkeli(1973a)reportedthatyoungternsintheAlandArchipelago could flyat36daysorless,asdidMartin(inPenland1976a)foryoungternsfromthePotholesReservoirareainWashington.Fledging Success Ludwig(1965a)estimatedaproductionof1.61 chicks perpairatcoloniesintheGreatLakesin1963-64.Thisfigure,dividedby mean clutch size(givenabove),providesafledging success of57.3%.Productioninthisareafrom1975to1978 was calculated at1.0 chicks per pair, anestimated success from39%oftheeggslaid(Shugartetal.1979).Shugartetal.indicatedthatsurvivalofyoung andnestsduringthisperiodwassuboptimalduetoperturbations(i.e.,nestsabandonedorwashedoutbyflooding).RatesofproductionsimilartothoseobservedbyLudwig(1.49-1.64chicksperpair)werefoundattwocoloniesinFinlandin1970-1972(Soikkeli1973a).MortalityofEggsandYoungLossofeggsandyoungbystormwindsandhightidesisperhapsthemostfrequentsourceofnestfailure.Somelocalitiesat whichsuch losseshavebeennotedincludeSouthCarolina(SpruntandChamberlain1949),San Francisco Bay(DeGroot1931),theGreatLakes(Shu-gartetal.1979),andthesouthwest Pacific (Jones1980).During1975-78intheGreatLakes,gale force winds(75+kph)resultedinthelossof12%ofallinitialnestingattempts.Thelargestsinglesourceofnestfailure,however,wastheabandonmentof445outof685nestsatHatIslandin1977astheresultofcannon-nettinginthecolony(Shugartetal.1979).Humandisturbanceofcolonieshasbeenconsideredasourceofegglosselsewhere.Penland(1976a) concluded thateggmortalityinthe Whitcomb Island,Gray'sHarbor,Washington,colonywasverylowin1976-76,"certainlylessthan5%,andprobablynear2%",butbelievedthatmostoftheloss could beascribedtotheeffectsofhumandisturbance. Such disturbancecausedmortalitybyexposingeggsandyoungto cold temperatures,byallowingneststobecoveredbydriftingsand,and byallowingpredatory gulls totakeeggs.Gullssometimespreyoneggs.Penland(1976a)believedthatthelargerthenumberofgulls,theworsetheeffectsofpredation.WhenfewgullswerepresentatacolonythatPenlandstudiedinWashington,theirpredationoneggsusuallyhadnosignficanteffectonnestingsuccess.Thegreatestdangerto chicks atthiscolonywasadultCaspianTernsthatattackedandkilledthe377

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CaspianTernchickswhentheywanderedintoneighboringterritories.KleptoparasitismbyCommonBlack-headedGullsonadultCaspianTernscarryingfoodcausedstarvationandincreasedmortalityinyoungCaspianTernsoffStockholm(Forssgren1980inBergman1980).RenestingSomeCaspianTernsthatlosetheirfirstnestsattempttorenest,butfiguresonratesofrenestingarenotavailable.Shugartetal.(1979)estimatedthat66%of465pairsthatfailed.intheirinitialnestingattemptsin1977atHatIsland,LakeMichigan,triedtonestagain,butthoughtthatthisestimatemayhavebeentoohigh.Therateofsurvivalofchicksfromrenestingattemptstendedtobesimilartothatfromfirstnestingattempts,butproductionofchickswaslowerbecausesecondclutchesweresmaller.AgeatFirstBreedingLudwig(1965a)believedthatmostCaspianTernsdonotbreeduntiltheirthirdyear.Thisaspectoftheirbreedingbiologyhasnotbeenstudiedadequately.MaximumNaturalLongevityTheoldestCaspianTernrecordedinNorthAmer was ajuvenilebandedin Hichigan thatreacheda minimumageof26years2months(Clappetal.1982a).Similarlongevitieshavebeenrecordedelsewhere.TwobirdsinFinlandworebandsfor19and20years,respectively(Soikkeli1970).EstimationshavebeenmadeofmortalityratesandexpectedsurvivalratesofCaspianTerns.Hickey(1952inSoikelli1970)calculatedmortalityratesof44%forbirdsinthefirstyearafterfledging,26%forthesecondyear,and18%forolderbirds.Ludwig(1965a)basedhiscalulationsonthe'samepopulationsintheGreatLakesthatwerestudiedbyHickey,andestimatedthattheaverageadultlifespanforCaspianTernsthereis8.8years,withanaverageannuallossof11.3%forbirdsolderthanthreeyears.Soikelli(1970)calculatedthatCaspianTernsinFinlandthathadsurvivedtoanageof1.5yearshada meanannualmortalityrateof12%,leadingtoafurtherlifeexpectancyof7.8years.WeightsBorodulina(1960)reportedthatthreefemales(presumablycollectedintheU.S.S.R.)rangedfrom 585to673 g(1.29-1.48lb),andaveraged640 g(1.41lb).Dement'evandGladkov(1951)gavetheweightofonefemaleas640g.Sixchicksabout10-15daysoldatWhitcombIsland,Washington,weighed180-257g(6.3-9.1oz)andaveraged233 g(8.2oz)(Penland1976a).Dement'evandGladkov(1951)reportedthatslightlyincubatedeggsweighed57-68g(2.0-2.4oz)andaveraged64g(2.3oz).SUSCEPTIBILITYTOOILPOLLUTIONCaspianTernshaveseldombeenreportedasvictimsofoiling.Threebandedbirdshavebeenrecoveredasaresultofoil--twoinCaliforniaandoneinLouisiana(filesoftheBirdBandingLaboratory,Laurel,MD).SeveralCaspianTernsthatwereoiledonthebreasthavebeenseenflyingintheCorpusChristiareainTexas(F.Buckley,pers.comm.).BecauseCaspianTernsareWidelydis-378

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CaspianTerntributedinsmallnumbersinmuchofthesoutheastandfeedwelloffshore,directeffectsofoilingarelikelytobeslight.Thespeciesmaybeatriskfromdevelopmentofonshorefacilitiesandotherformsoflanddevelopmentbecauseitissusceptibletohumandisturbance.Italsonestsonbarrierisletsandspoilislandsthatareeasilyaffectedbydredginganderosioncontrol.ThestatusofthisspeciesalongtheTexascoast,where asignificantproportionoftheU.S.populationnests,shouldbecarefullymonitored.BIBLIOGRAPHY1982Koonz,W.H.1982.VandalisminaManitobaCaspianTerncolony. Hlue Jay40:48-49.1981 Bergman,G.1981.Single-breedingversuscolonialbreedingintheCaspianTern,Hydroprognecaspia.(Abstractonly.)Mem.Soc.FaunaFloraFenn.57:1.---Blokpoel,H.1981.AnattempttoevaluatetheimpactofcannonnettinginCaspianTerncolonies.ColonialWaterbirds4:61-67.Forssgren,K.1981.ThekleptoparasiticbehaviouroftheArcticSkuaStercorariusparasiticusandtheLesserBlack-backedGullLarusfuscuswiththeCaspianTernHydroprognecaspia.(Abstractonly.) FaunaFloraFenn.57:5.Harvey,T. E.1981.ThebreedingecologyofCaspianandForster'sTernsinElkhornSloughSaltponds,California.(Abstractonly.)Pac. GroupBull.8:93.'"King,B.1981a.CaspianTernssun-bathingwithRing-billedGulls.Brit.Birds74:181.1981b.CaspianTerndroppingandretrievingobjects.Brit.Birds--74:228.Ludwig,J.P.1981.BandwearandbandlossintheGreatLakesCaspianTernpopulationand ageneralizedmodelofbandloss.ColonialWaterbirds4:174-186.Raffaele,H.1981.NewrecordsofbirdspeciesforPuertoRicoandonefortheWestIndies.Am.Birds35:142-143.1980 Bergman,G.1980.Single-breedingversuscolonialbreedingintheCaspianTernHydroprognecaspia,theCommonTernSternahirundoandtheArcticTernSterna OrnisFenn.57:141-152.[InEnglishwithFinnishsummary.]379

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CaspianTernHockey,P.A.R.and C.T.Hockey.1980.Notes onCaspianTernsSternacaspiabreedingneartheBergRiver,SouthwesternCape.Cormorant8:7-10.------Jones,A.1980.Theeffectsofweatheron aCaspianTerncolony.Notornis27:405-406.Pulliainen,E.andA.Marjakangas.1980.EggshellthicknessinelevenseaandshorebirdspeciesoftheBothnianBay.OrnisFenn.57:65-70.Quinn,J.1980.ChickfeedinginCaspianTerns.(Abstractonly.)Proc.1979Conf.ColonialWaterbirdGroup3:259.Staav,R.1980.SkrantarnansSternacaspiaupptradandeiSverigeunderhogocheftersommar.[OccurrenceoftheCaspianTernSternacaspiainSwedenduringhighandlatesummer.]VarFagelvarld 39:139-148:--yyn SwedishwithEnglishsummary.] 1979Baltz,D.M.,G.V.MorejohnandB.S.Antrim.andfoodhabitsoftwoCaliforniaterns.1979.SizeselectivepredationWest.Birds10:17-24.Byard,M.E.1979.CaspianTerninCimarronCounty,Oklahoma.Bull.Okla.Ornithol.Soc.12:21.Caldwell,J.R.and C.Jorgenson.1979.CaspianTernnestsatLastMountainLake,Saskatchewan.BlueJay37:Ill.Gill,R.,Jr.and L. R. Mewaldt. 1979.DispersalandmigratorypatternsofSanFranciscoBayproducedherons,egrets,andterns.N.Am.BirdBander4:4-13.Martin,M.1979.StatusreportonCaspianTern(Sternacaspia)inCanada.Can.Wildl.Serv.,Ottawa,Ontario,Canada.43pp.------Martinez,A.andJ.Muntaner.1979.MigrationdelaSternecaspienneHydroprognecaspiaparIeDeltadel'Ebre.Alauda47:29-33.Shugart,G.W.,W.C.SharfandF.J.Cuthbert.1979.StatusandreproductivesuccessoftheCaspianTern(Sternacaspia)intheU.S.GreatLakes.Proc.1978Conf.ColonialWaterbirdGroup2:146-156.Staav,R.1979.DispersalofCaspianTerns(Sternacaspia)intheBaltic.OrnisFenn.56:13-17.Stevenson,H.M.1979b.FirstnestingoftheCaspianTernintheFloridaPanhandle.Fla.FieldNat.7:8.380

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CaspianTernTombal, C. andJ.C.Tombal.1979.DonneescomplementairessurlamigrationdelaSternecaspienneHydroprognecaspiaparIeDeltadeL'Ebre(Catalogne).[Onthemigrationof Hydroprogne caspiaintheEbraDelta(Catalonia).]Alauda47:303-304.Weseloh,D.V. andL.Cocks.1979.RecentnestingoftheCaspianTernatEggIsland,LakeAthabasca,Alberta.BlueJay37:212-215.1978Blokpoel,H.andP.M.Fetterolf.1978.ColonizationbygullsandternsoftheEasternHeadland,Toronto,OuterHarbour.Bird-Banding49:59-65.Brichetti,P.1978.SullanidificazioneinItaliadi:GabbianocorallinoLarusmelanocephalusTemminck,GabbianoroseoLarusgeneiBreme,RondinedimaremaggioreHydroprognetschegrava(Lepechin).Riv.Ital.Ornitol.48:215-233.Clinning,C.F.1978.BreedingoftheCaspianTerninSouthWestAfrica.Cormorant5:15-16.Haymes,G.T.andH.Blokpoel.1978a.ReproductivesuccessoflaridsnestingontheEasternHeadlandoftheTorontoOuterHarborin1977.OntoFieldBioI.32:1-17.Herman,J.F.,R.A.SchmidtandK.J.Wilson.1978.FirstnestingrecordoftheCaspianTerninNorthDakota.PrairieNat.10:23-24.Hosea,F.1978.Techniquesforternandgullbanding.N.Am.BirdBander3:Ill.Loftin,R.w.1978.CaspianTernsintheGeorgiamountains.Oriole43:66-67.Schreiber,R.w.1978.Speciesofspecialconcern.CaspianTern.P.94inH.W.Kale,II(ed.)Rare andendangeredbiotaofFlorida.Vol.2.Birds.Univ.PressesofFlorida,Gainesville,FL.xixand121pp.Shugart,G.w.1978.ThedevelopmentofchickrecognitionbyadultCaspianTerns.Proc.1977Conf.ColonialWaterbirdGroup1:110-117.Smith,J.L. andD.R. Mudd.1978.FoodoftheCaspianTerninGraysHarbor,Washington.Murrelet59:105-106.Staav,R.1978.[AstudyofCaspianTerns(Sternacaspia)inBlekinge1977.]FaglarBlekinge14:xviii-xx.[In Whitfield,A.K.and S.J.M.Blaber.1978.FeedingecologyofpiscivorousbirdsatLakeSt.Lucia.PartI:divingbirds.Ostrich49:185-198.381

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CaspianTern1977Allen,L.J.1977.ThecomparativefeedingecologyrelativetopopulationdynamicsoftheHerringandRing-billedGulls(LarusargentatusandL.delawarensis)andsomecomparisonswiththeCaspianTern.M.S.thesis.QueensUniv./Kingston,Ontario.196pp.Baglieri,S.andF.Fagotto.1977.LarondinediMareMaggiore--(Hyproprognetschegrava(Lepechin)--nellaSiciliaorientale.[TheCaspianTern(Hydroprognetschegrava)ineasternSicily.]Riv.Ital.Ornitol.47:229-234.tInItalianwithEnglishsummary.]Blokpoel,H.1977.GullsandternsnestinginnorthernLakeOntarioandtheupperSt.LawrenceRiver.Can.Wildl.ServoProg.Notes75.12pp.Fetterolf,P.M.and H.Blokpoel.1977.TernsandgullsnestingonToronto'sEasternHeadland.Ont.FieldBioI.31:51-52.Shugart,G.W.1977c.Nest,nest-site,egg,andchickrecognitionbyadultCaspianTerns.M.S.thesis,N.IllinoisUniv./DeKalb,IL.Staav,R.1977.EtudedepassagedelaSternecaspienneHydroprognecaspiaenMediterraneeapartirdesreprisesd'oiseauxbaguesenSuede.[StudyofthemigrationofCaspianTernsintheMediterraneanbycapturesoftaggedbirdsinSweden.]Alauda45:265-270.[InFrenchwithEnglishsummary. ] Weske,J.S.,R.B.ClappandJ.M.Sheppard.1977.BreedingrecordsofSandwichandCaspianTernsinVirginiaandMaryland.Raven48:59-66.1976Becker,P.1976.Raubseeschwalbe(Hydroprognecaspia)alsBrutvogelanderSudwestafrikanischenKuste.Namib& [InGermanwithEnglishsummary.]Gill, R. E.,Jr.1976.NotesontheforagingofnestingCaspianTernsHydroprognecaspia(Pallas).Calif.FishGame62:155.Isenmann,P.1976.BeweiseineslangerenAufenhalteseinerGruppeRaubseeschwalben(Hydroprognecaspia)wahrenddesWegzugesinderCamargue(SudFrankreich). Vogelwarte-zs:-3l2-3l3. [InGerman.]Parnell,J.F.andR.F.Soots.1976.CaspianTernnestinginNorthCarolina.Chat40:14-15.Penland,S.1976a.ThenaturalhistoryandcurrentstatusoftheCaspianTern (Hydroprognecaspia)inWashingtonState.M.S.thesis,Univ.PugetSound/Tacoma,WA.101pp.1976b.TheCaspianTern:anaturalhistory.Wash.Wildl.28:17-19.382

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Wolinski, R. A.1975.County[Mich.].CaspianTern1975Dunstan,F.M., R. W.SchreiberandJ.J.Dinsmore.1975.CaspianTernnestinginFlorida,1973and1974.Fla.FieldNat.3:16-17.FirstreportedCaspianTernsightingforIsabellaJack-PineWarbler53:75.1974Alcorn,G.D.1974.CaspianTern,seawandereroftheworld.Pac.Search8:4-5.Britton,P.L.andL.H. Brown.1974.ThestatusandbreedingbehaviourofEastAfricanLari.Ostrich45:63-82.Cronert,H.andM.Ny.1974.SkrantarnansforekomstinordostraSkane.Anser13:254-256.Koli,L.andM.Soikkeli.1974.FishpreyofbreedingCaspianTernsinFinland.Ann.Zool.Fenn.11:304-308.Rudebeck,G.1974.RapportfranSilvakratornetvidkrankesjon.1.OvantatrikligforekomstavskrantarnaHydroprognetschegrava.Anser13:149-153.[InSwedishwithEnglishsummary.] 1973Horton,W.1973.CaspianTernsatMountIsa,Q.Austr.BirdBander11:51-55.Isenmann,P.1973.LepassagedelaSterneCaspienneHydroprognecaspiaen1971et1972enCamargue. [The 1971 and 1972passagesoftheCaspianTernHydroprognecaspiaintheCamargue.]Alauda41:365-370.[InFrenchwithEnglishsummary.]Soikkeli,M.1973a.BreedingsuccessoftheCaspianTerninFinland.BirdBanding44:196-204.1973b.Long-distancefishingflightsofthebreedingCaspianTern-----Hydroprognecaspia.OrnisFenn.50:47-48.Vaisanen,R.A.1973.EstablishmentofcoloniesofCaspianTernHydroprognecaspiabydesertingflightsinthenorthernGulfofBothnia.OrnisScand.4:47-53.Vermeer,K.1973a.ComparisonoffoodhabitsandmercuryresiduesofCaspianandCommonTerns.Can.Field-Nat.87:305.383

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CaspianTern1972Gill,R.E.,Jr.1972.SanFranciscoBaybreedingbirdsurvey,1971.Calif.Dep.FishGameWildl.Manage.BranchAdm.Rep.72-6.Schreiber,R.W.andJ.J.Dinsmore.1972.CaspianTernnestingrecordsinFlorida.Fla.Nat.45:161.Staav,R.,B.Almkvistand S.Hedgren.1972.SkrantarnanHydroprognetschegravaiSverige1971. [TheCaspianTernHydroprognetschegravainSweden1971.]VarFagelvarld31:241-246.[InSwedishwithEnglishsummary.]Vermeer,K.1972. ComparisonoftheclutchinitiationofCaspianandCommonTernsatLakeWinnipeg.BlueJay30:218-220.1971Beretzk,P.andA.Keve.1971.las)inUngarn(1953-1969).DieRaubseeschwalbeHydroprognecaspia(PalZool.Abh.(Dres.)30:227German.]Campbell,R.W.1971.StatusoftheCaspianTerninBritishColumbia.Syesis4:185-189.Diamond,A.W.1971.Theecologyofthesea-birdsofAldabra.Philos.Trans.R.Soc.Lond. B.BioI.Sci.260:561-571.McDaniel,T.H.and T.A.Beckett,III. CaspianTernnestinginSouthCarolina.Chat35:39-41.1970Chaniot,G.E.,Jr.1970.NotesoncolorvariationindownyCaspianTerns.Condor 72:460-465.Dobrowolski,K.A.1970.Wystepowanierybitwywielkodziobej,Hydroprognecaspia(Pall.)wPolscewokresieubieglych150lat.[TheoccurrenceofCaspianTern,Hydroprognecaspia(Pall.)inPolandduringthepast150years.]ActaOrnithol.12:209-228.[InPolishwithEnglishsummary.]Evans,R.M.,D.B.KrindleandM.E.Mattson.1970.nearSpruceIsland,LakeWinnipegosis,Manitoba.CaspianTernsnestingBlueJay28:68-71.Schick,W.J.1970.FirstBritishColumbiarecordofCaspianTern.Syesis3:187.Soikkeli,M.1970.MortalityrateofFinnishCaspianTernsHydroprognecaspia.OrnisFenn.47:177-179.Vermeer,K.1970.LargecoloniesofCaspianTernson Lakes WinnipegandWinnipegosis,1970.BlueJay28:117-118.384

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CaspianTern1969Daneel,A.B.1969.BreedingoftheCaspianTernatVaaldam.Ostrich40:16.Dunstan,T.C.fishline.1969.TernmortalityduetoentanglementinnylonmonofilamentLoon41:50-51.Jozefik,M.1969.[CaspianTern,HydroprognecaspiaPall.inPoland--thebiologyofmigrationperiod.]ActaOrnithol.11:381-443.[EnglishtranslationfromthePolish.]Warner,D.W.andD.A.Beimborn.1969.FirstCaspianTernnestinginMinnesota.Loon41:83-84.1968 Ludwig,J.P.1968.DynamicsofRing-billedGullandCaspianTernpopulationsoftheGreatLakes.Ph.D.thesis,Univ.Michigan/AnnArbor,MI. Rohwer,S.A.1968.SecondbreedingrecordoftheCaspianTerninFlorida.Fla.Nat.41:35.Schwarz,R.andP.Kragenow.1968.ZumDurchzugderRaubseeschwalbeimNaturschutzgebiet"GrasserSchwerin"imKreisRobel/Muritz.Falke15:264-267.[InGerman.]1967 Diem,K.L.andD.deL.Condon.1967.BandingstudiesofwaterbirdsontheMollyIslands,YellowstoneLake,Wyoming.YellowstoneLibraryand Mus.Assoc.41pp.Tomialoje,L.1967.Materialydo aWifauny Polski.IV.Hydroprognecaspia(Pall.)--rybitwawielkodzioba.[ContributionstotheavifaunaofPoland.IV.Hydroprognecaspia(Pall.)--CaspianTern.]ActaOrnithol.10:25-82.[InPolishwithEnglishtranslation.]1966 Amadon,D.1966.ThescientificnameoftheCaspianTern.Ibis108:424-425.Cunningham,R.L.1966.CaspianTernfeedingoncarrion.WilsonBull.78:319.Maluquer,S.andJ.A.Albert.1966.LaPagazaPiquirrojaHydroprognecaspiaenelDeltadelEbro.Ardeola11:155.Vaurie,C.1966.Comments onthescientificnamesoftheCaspianTernandPiedWheatear.Ibis108:633-634.385

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CaspianTern1965Bezzel,E. andJ.Reichholf.1965.VomZugderBinnenseeschwalbenChlidoniasundderRaubseeschwalbeHydroprognecaspiainSudbayern.Vogelwarte23:121-128.[InGerman.]Hader, R. J.1965.MoreCaspianTernsatRaleigh,NorthCarolina.Chat29:87.Ludwig,J.P.1965a.BiologyandstructureoftheCaspianTern(Hydroprognecaspia)populationoftheGreatLakes from1896-1964.Bird-Banding36:217-233.1965b.Ring-billedGullnestingwithinCaspianTerncolony.Jack-Pine-----Warbler43:61.Taro,R.1965.Rayskia(Hydroprognecaspia)syyskesalla1963ja-64HauhonselalIa.[ObservationsoftheCaspianTernatHauho.]OrnisFenn.42:63-67.[InFinnishwithEnglishsummary.] 1964Martini,E.1964.OtolitheninGewollenderRaubseeschwalbe(HYdroprognecaspia).[OtolithsinpelletsoftheCaspianTern(Hydroprognecaspia).]--Bonn.Zool.Beitr.15:59-71.[InGermanwithEnglish 1963 Bergman,G.1963.Rovternen.Pp.79-88inN.Bladel(ed.).farver,Vol.4.Munksgard, [InDanish.]NordensfugleiHader,R.J.1963.CaspianTernatRaleigh,NorthCarolina.Chat 27:79-80.Tomkins,I.R.1963b.Skimmer-likebehaviorintheRoyalandCaspianTerns.Auk80:549.Woolfenden,G.E. andA.J.Meyerriecks.1963.CaspianTernbreedsinFlorida.Auk80:365-366.1962 Ludwig,J.P.1962b.AsurveyofthegullandternpopulationsofLakesHuron,Michigan,andSuperior.Jack-PineWarbler40:104-119.Schevareva,J.P. NeuedatenuberWiederfundeberingterTschegraven(HydroprognetschegravaLepechin).MigrationenderTiere,Moskva:92-105.[InGerman.] 336

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CaspianTern1961 Doncev,S.,A.Prostovand Z.Georgiev.1961.DieRaubseeschwalbe,Hydroprognetschegravatschegrava(Lepechin,1770),eineneueArtinderbulgarischenOrnithofauna.Izv.Zool.Inst.Bulg.Akad.N.10:389-391.Dragesco,J.1961b.ObservationsethologiquesdesoiseauxdeBaned'Arguin.Alauda29:81-98.[InFrench.]Lewis,C. B.1961.CaspianTernsinJamaica.Auk78:264-265.Ludwig, C.C.andF.E. Ludwig. 1961.HerringGullsandCaspianTerns--1960recoveries.Inl.BirdBandingNews33:28-29.Ziedler,K.1961.RaubseeschwalbeamGalbenbacherSee.Falke8:75-78.1960Moltoni,E.1960.AltrenotizieriguardantilaRondinedimaremaggioreHydroprognecaspia(Pall.)inItalia.Riv.Ital.Oruitol.30:197-200.[InItalian.-]-----Suchantke,A.1960.ComportementdepechedelaSternecaspienne(Hydroprognecaspia)d'apresdesobservationsenCamargue.Alauda28:38-44.[InFrench.]Tarr,H.E.1960.NestingoftheCaspianTern.1959Austr.Bird Watcher 1:78-81.Briche,L.1959.ObservationdeSternescaspiennesHydroprognecaspia(Pallas)aArras.Alauda27:321.[InFrench.]Davidson,A.R.1959.TheCaspianTern.VictorianNat.16:21.deNaurois,R.1959.Premieresrecherchessurl'avifaunedesilesduBaned'Arguin(Mauritanie).Alauda27:241-308.[InFrench.]Mayaud,N.1959.Hydroprognecaspia(Pallas)aBayonne.Alauda27:234.[InFrench.]Tordoff,H.B.andW.E.Southern.1959.CaspianTernandBlackSkimmerinNewfoundland.WilsonBull.71:385-386.1958Alcorn,E.D.1958.NestingoftheCaspian inGraysHarbor,Washington.Murrelet39:19-20.387

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CaspianTernGeroudet,P.1958.Alauda26:67.CurieusemethodedepechechezlaSternecaspienne.[InFrench.]Mayaud,N.1958.MigrationethivernageenAfriqued'Hydroprognecaspia.Alauda26:151.[InFrench.]Moltoni,E.Italia.1958.LaRondinedimaremaggioreHydroprognecaspia(Pall.)inRiv.Ital.Ornitol.28:218-223.[InItalian.-]------Pettingill,O.S.,Jr.1958b.NestingoftheCaspianTernwithinaRing-billedGullcolony.Jack-PineWarbler36:183-184.Suchantke,A.1958.HydroprognecaspiaenCamargue.Alauda26:306.[InFrench.]1957 Bergman, G.1957.Analyseder"aggressivenBeugungen" unddes"Herabstarrens"derRaubseeschwalbeHydroprognetschegrava.OrnisFenn.34:113-117.[InGerman.]Jakab,A.1957.AkacagoeserfeszkeleseesalocserelofordulasaFonyodon.[AppearanceoftheGull-billedTernandCaspianTernatFonyod(LakeBalaton).]Aquila63/64:284,342.[InHungarianandEnglish.]Stein-Spiess,S.1957.AlocserelsoizbenErdelyben.[FirstoccurrenceoftheCaspianTerninTransylvania.]Aquila63/64:285,343.[InHungarianandEnglish.]Tenovuo,R.1957.Rayskan,Hydroprognetschegrava(Lep.),pesimisestaPeramerella.[OnthebreedingoftheCaspianTerninthenorthernGulfofBothnia.]OrnisFenn.34:140-141.[InFinnishwithEnglishsummary.]1956 Bergman, G.1956.Beteendestudieroverungaravskrantarna(Hydroprognetschegrava).[OnthedevelopmentofthebehaviorinchicksoftheCaspianTern.]VarFagelvarld15:223-245.[InSwedishwithEnglish sunnnary.] Houston,S.1956.TheCaspianTerninSaskatchewan--withfirstnestingrecord.BlueJay14:116-117.Mayaud,N.1956.Etudesurlamigrationetleszonesd'hivernagedesSternescaspiennesd'Eurasie.Alauda24:206-218.[InFrench.]Milon,P.1956.SternescaspiennesenCamargue.Alauda24:232.[InFrench.]Rijsdijk,A.1956.EennieuwewaarnemingvandeReuzenstern,Hydroprognecas (Pall.).Ardea44:243-244.[InDutch.]388

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CaspianTern1955 Bergman, G.1955.DieBeziehungenzWischenBodenfarbederRiviereundFarbederKuckenbeiHydroprognetschegravaundSternamacrura.[OnthecolorvariationofthechicksofHydroprognetschegravaandSternamacrura,andtheinfluenceofthegroundcoloronthechicks'coloration.]OrnisFenn.32:69-83.[InGermanwithEnglishandFinnishsummaries.]Gustavsson,U.andB.J.O.Wahlin.1955.Havstrutar(Larusmarinus)ochskrantarnor(Hydroprognetschegrava)vidRoxen. VarFagelvarld14:123.1954Atkeson,T.Z.,Jr.andD.C.Hulse.1954.NorthernAlabamanotesontheCaspianTern(Hydroprognecaspia).Ala.Birdlife2:35.Beretzk,P.1954.DieRaubseeschwalbe(Hydroprognecaspia[Pall.])inUngarn.AllattaniKozl.44:133-147.Emblen,D.L.1954.CaspianTernsnestingatSan Diego Bay. Condor56:109110.Sharrock,J.T.R.1954.CaspianTerninKent.Brit.Birds47:403-404.Wolff,A.B.H.1954.WaarnemingvaneenviertalReuzensterns,Hydroprognecaspia(Pall.),aandezuidelijkeIJsselmeerkust.Ardea42:345346.[InDutch.]1953 Bergman,G.1953.VerhaltenundBiologiederRaubseeschwalbe(HYdroprognetschegrava).[BehaviorandbiologyoftheCaspianTern.]ActaZool.Fenn.77: 150.[InGerman.]Blomgren,A.1953.Skrantarnansupptradandeiinlandet.NorrbottensNat.2:39.1952Hickey,J.J.1952.Survivalstudiesofbandedbirds.U.S.FishWildl.ServoSpec.Sci.Rep.Wildl.15.177pp.1950 Kux, Z.1950.OccurrenceoftheCaspianTerninSouthMoravia.Sylvia11/12:79. Low, S.H.1950.Atwenty-two-year-oldCaspianTern.Bird-Banding21:115.389

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CaspianTern1949Southern,H.H.1949.CaspianTernandRing-billedGullcolonies,GreenIsland,PrinceEdwardCounty,Ont.Can.Field-Nat.63:115-116.1947Suchsdorff,A.1947.SkrantarnaharhackatvidinsjoiSodermanland.VarFagelvarld6:162-163.Thomas, B. L.1947.ThelifecycleofDiphyllobothriumoblongatum.J.Parasitol.33:107-117.1946Cottam,C.1946.LatenestingofCaspianTerninUtah.Condor48:94-95.1943Miller,A.H.1943. CensusofacolonyofCaspianTerns.Condor45:220-225.1942East,B.1942.CityoftheCaspians.Fauna4:18-21.Ludwig,F.E.1942.MigrationofCaspianTernsintheGreatLakesarea.Bird-Banding13:1-9.1941Witherby,H.F.1941.CaspianTernringedinU.S.A.foundinYorkshire.Brit.Birds34:184.1940Lincoln,F.C.1940a.AnAmericanCaspianTerninEngland.Auk57:569.1940b.CaspianTerninHaiti.Auk57:569.1934 Tomkins,I.1934.Acuriousternaccident.WilsonBull.46:128.1932Beardslee,C.S.1932.CaspianTerninConnecticut.Auk49:85.Potter,J.K.1932.CaspianTern(Sternacaspiaimperator)inCamdenCounty,N.J.Auk49:476-477.390

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CaspianTern1931DeGroot,D.S.1931.HistoryofanestingcolonyofCaspianTernsofSanFranciscoBay. Condor 33:188-192.1930Kitchin,E.A.1930.CaspianTern added totheWashingtonlist.Murrelet11:19-20.1929Emlen,J.T.,Jr.1929.CaspianTerns(Sternacaspiaimperator)atPalmyra,N.J.Auk46:534.1928 Harms,M.1928.DasVorkommenunddieBrutvehaltnissederRaubseeschwalbeinEstland.[TheappearanceandbreedingcircumstancesoftheCaspianTerninEstland.]J.Ornithol.76:420-427.[InGerman.]Janvrin,E.R.P.1928.CaspianTernon LongIsland,N.Y.Auk45:497.1927Yoder,W.andJ.K.Potter.1927.CaspianTernatStoneHarbor,N.J.Auk44:92.1924 Magee,M.J.1924.C
Marine birds of the southeastern United States and Gulf of Mexico: part III: charadriiformes
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Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00000171/00003
 Material Information
Title: Marine birds of the southeastern United States and Gulf of Mexico: part III: charadriiformes
Series Title: Marine birds of the southeastern United States and Gulf of Mexico
Physical Description: 3 v. : ill., maps ; 28 cm.
Language: English
Creator: Clapp, Roger B.
Morgan-Jacobs, Deborah
Banks, Richard C., 1940-
National Coastal Ecosystems Team (U.S.)
United States -- Minerals Management Service
U.S. Fish and Wildlife Service
Denver Wildlife Research Center -- Museum Section
Publisher: Bureau of Land Management, Fish and Wildlife Service, U.S. Dept. of the Interior
Place of Publication: Washington, D.C.
Publication Date: 1983
 Subjects
Subjects / Keywords: Sea birds -- Southern States   ( lcsh )
Sea birds -- Gulf States   ( lcsh )
Birds -- Southern States   ( lcsh )
Birds -- Gulf States   ( lcsh )
Genre: non-fiction   ( marcgt )
 Notes
Statement of Responsibility: prepared for National Coastal Ecosystems Team, Division of Biological Services, Fish and Wildlife Service, U.S. Department of the Interior.
Bibliography: Includes bibliographies.
General Note: "Museum Section, United States Fish and Wildlife Service, National Museum of Natural History."
General Note: Project sponsored by the Bureau of Land Management, Minerals Management Service.
General Note: Pt. 3: "Prepared for National Coastal Ecosystems Team, Division of Biological Services, Fish and WIldlife Service, U.S. Department of the Interior."
General Note: "March 1982" (pt.1); "July 1982" (pt.2); "September 1983"--Pt. 3.
General Note: "Contract no. 14-16-0009-78-917."
 Record Information
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0257
notis - AME1920
alephbibnum - 002436756
oclc - 08359556
lccn - 82601888 //r83
System ID: UF00000171:00003

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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
    Title Page
        Page i
    Disclaimer
        Page ii
    Preface
        Page iii
    Abstract
        Page iv
    Table of Contents
        Page v
        Page vi
        Page vii
    List of maps
        Page viii
        Page ix
    List of Tables
        Page x
        Page xi
        Page xii
        Page xiii
        Page xiv
    Abbreviations used in text
        Page xv
        Page xvi
    Purpose of report and study area
        Page 1
        Page 2
        Page 3
    Methods
        Page 4
        Page 5
        Page 6
    Arrangement and contents of species accounts
        Page 7
        Page 8
        Page 9
        Page 10
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    Oil pollution and marine birds of the southeastern United States
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    Recommendations for future research
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    Acknowledgement
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    Species accounts
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Full Text
























































..










FWS/OBS-83/30
September 1983



MARINE BIRDS OF THE SOUTHEASTERN UNITED STATES

AND GULF OF MEXICO



PART III



CHARADRIIFORMES


by



Roger B. Clapp, Deborah Morgan-Jacobs, and Richard C. Banks



Museum Section
United States Fish and Wildlife Service
National Museum of Natural History
Washington, D.C. 20560


Contract No. 14-16-0009-78-917


Project Officers
Jeffrey A. Spendelow
Cherry Keller
National Coastal Ecosystems Team
United States Fish and Wildlife Service
NASA-Slidell Computer Complex
1010 Cause Boulevard
Slidell, Louisiana 70458

This project was sponsored by the
Bureau of Land Management
(Minerals Management Service)

Prepared for
National Coastal Ecosystems Team
Division of Biological Services
Fish and Wildlife Service
U.S. Department of the Interior
Washington, D.C. 20240










DISCLAIMER


The findings in this report are not to be construed as an official U.S.
Fish and Wildlife position unless so designated by other authorized documents,
nor does the mention of any commercial items indicate an endorsement by the
U.S. Government.


Library of Congress Card Number 82-601888









This report should be cited as:

Clapp, R. B., D. Morgan-Jacobs, and R. C. Banks. 1983. Marine Birds of the
Southeastern United States and Gulf of Mexico. Part III: Charadriiformes.
U.S. Fish and Wildlife Service, Division of Biological Services, Washing-
ton, D.C. FWS/OBS-83/30. xvi and 853 pp.













Part III of the volumes
Gulf of Mexco, published by
synthesis and analysis of information about the marine birds in thU s area. Ac
counts for 22 speces include information on distribution, abundance, and sus-
ceptblity to oil pollution. Also included is info on on the breeding
biology of 16 species abundant in the southeast as breeding birds, winter rest
dents, or migrants. selected bibliographies follow each species account and
include additional sources of information.,

Any suggestions or questions regarding this report should be directed
to:

Information Transfer Speclalrst
National Coastal Ecosystems Team
U.S. Fish -nd Wildlife Ser-..e
NASA-Siadel Computer Complex
d010 ause Bouslevard
Slldell, Loulsiana 70458


A badly oiled ull (Larus sp


.) Photograph by Roger B. Clapp.


iii


PEACE













Information on the seasonal distribution and abundance of 22 speIes of
marine birds of the order Charadriiformes that occur in the coastal south-
eastern United States has been compiled and mapped from the literature. In
many Instances this provides the first synthesis of knowledge about a species
for this region. We also provide information on global distribution, habitat
and food for all species, and include inflation on various aspects of lfe
history for the 16 species that we consider most important in coastal areas.
This information was gathered in an attempt to assess the possible effects
of offshore oil development on populations of marine birds in the southeast.

The susceptibility of birds to oil depends not only on theor 3uxtaposi-
tlon in time and space, but also on currents, clmatic factors, the stage of
the life or annual cycle, and the behavior of the species. Contamination by
oil may result in matted feathers with death following from chilling, star
nation, and ingestion o ooil during preening. Few of the species covered in
this report are at great hazard from the direct effects of "iling, but
populations of most of these speezes are highly susceptible to environmental
change. large concentratons of wintering, breeding, and migrant gulls and
terns occur in the southeast and in some instances make up a large propor-
tion of the global or North ~mercan population. Consequently, this report
includes most of the marine birds that we believe most likely to be detri-
mentally affected by the development of oil resources.

One of the conclusions reached by this report Is that we still know
very little about the status and populations of some of the charadridforms
that occur in the southeast. Mditlonal surveys of colonial marine birds
in the southeast and nearby waters are badly needed to ensure that we now
enough about them to prevent their untimely loss from our coastal areas.




. *i


A Comon Tern nest. Photograph by Roger B. Clapp.










CONTENTS




PREFACE . . . . . . . . . . . . . . . .

ABSTRACT . . . . . . . . . . . . . . . .

MAPS . . . . . . . . . . . . . . . . .

TABLES . . . . . . . . . . . . . . . . .

ABBREVIATIONS USED IN TEXT . . . . . . . . . . . .

PURPOSE OF REPORT . . . . . . . . . . . . .

STUDY AREA . . . . . . . . . . . . . . .

Habitats . . . . . . . . . . . . . . .
Climates . . . . . . . . . . . . . . .

METHODS . . . . . . . . . . . . . . . .

ARRANGEMENT AND CONTENTS OF SPECIES ACCOUNT . . . . . . .

Species Included . . . . . . . . . . . . .
Scientific and Vernacular Names . . . . . . . . .
General Distribution . . . . . . . . . . . .
Distribution and Abundance in the Coastal Southeastern United
States . . . . . . . . . . . . . . .
Synopsis of Present Distribution and Abundance .. . . .....
Habitat . . . . . .. .
Food and Feeding Behavior . . . . . .
Important Biological Parameters . . . . . . . . .
Susceptibility to Oil Pollution . . . . . . . . .
Species Bibliography . . . . . . . . . . . .

OIL POLLUTION AND MARINE BIRDS OF THE SOUTHEASTERN UNITED STATES . .

Variability among Species in Susceptibility to Oil Pollution . .
Regional and Seasonal Differences in Oiling and Mortality of
Beached Birds . . . . . . . . . . . . .
Bird Kills Following Oil Spills in the Southeastern U.S. . . .
Sources of Variation in Mortality from Oil Pollution . . . .
Effects of Oil on Contaminated Birds and Their Eggs . . . .
Potential Hazards to Marine Birds from Offshore Oil Production . .

RECOMMENDATIONS FOR FUTURE RESEARCH . . . . . . . . .


Page

iii

iv

viii

x

xv

1

1

1
3

4

7

8
9
10

10
11
11
12
12
13













Status and Biology of Breeding Species . . . . . . . .
Distribution and Biology of Transient, Wintering, and Pelagic
Populations . . . . . . . . . . . . . .
Effects of Oil on Southeastern Marine Birds . . . . . .


ACKN(

SPECI


Caspian Tern (Sterna caspia)


)WLEDGEMENTS . . . . . . . . .

IES ACCOUNTS . . . . . . . . .

CHARADRIFORMES

Scolopacidae (Phalaropidinae)

Red-necked Phalarope (Phalaropus lobatus) . .


Red Phalarope (Phalaropus fulicaria) . . .


Laridae (Larinae)

Laughing Gull (Larus atricilla) . . . .


Franklin's Gull (Larus pipixcan) . . .


Bonaparte's Gull (Larus philadelphia) . . .


Ring-billed Gull (Larus delawarensis) . . .


Herring Gull (Larus argentatus) . . . .


Great Black-backed Gull (Larus marinus) . .



Laridae (Sterninae)

Gull-billed Tern (Sterna nilotica) . . .


. . . . . . account . .
bibliography .


. account . .
bibli ograph .
S. account . .
bibliography .


Account . .
bibliography .


S. account . .
bibliography ..

S. account . .
bibliography .

. account . .
bibliography ..

S. account . .
bibliography .

S. account . .
bibliography .


S. account . .
bibliography .

. account . .
bibliography .


Page

28

33
35












Royal Tern (Sterna maxima) . .


Sandwich Tern (Sterna sandvicensis)


Roseate Tern (Sterna dougallii)


Common Tern (Sterna hirundo) . .


Arctic Tern (Sterna paradisaea) ..


Forster's Tern (Sterna forsteri)


Least Tern (Sterna antillarum)


Bridled Tern (Sterna anaethetus)


Sooty Tern (Sterna fuscata) . .


Black Tern (Chlidonias niger) . .


Brown Noddy (Anous stolidus)


. . . . account .
bibliography

. . . . account .
bibliography

. . . . account .
bibliography

. . .. . account .
bibliography

. . . . account .
bibliography

. . . .. account .
bibliography

. . . . account .
bibliography

. . . . account .
bibliography

. . .. . account .
bibliography

. . . . account .
bibliography

. . .. . account .
bibliography


Laridae (Rynchopinae)


Black Skimmer (Rynchops niger)


. . . . . account . .
bibliography .


LITERATURE CITED . . . . . . . . . . . . .


Page

393
409

417
436

450
467

483
512

551
559

579
593

599
617

637
651

656
675

689
701

717
724


. .










MAPS


Number

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

21

22


viii


Breeding Range Map for the Laughing Gull . . .. . . .

Winter Distribution Map for the Laughing Gull . . . .

Winter Distribution Map for the Bonaparte's Gull . . .

Winter Distribution Map for the Ring-billed Gull . .

Breeding Range Map for the Herring Gull . . . . . .

Winter Distribution Map for the Herring Gull . . . .

Breeding Range Map for the Great Black-backed Gull . . .

Winter Distribution Map for the Great Black-backed Gull . .

Breeding Range Map for the Gull-billed Tern . . . . .

Winter Distribution Map for the Gull-billed Tern . . .

Breeding Range Map for the Caspian Tern . . . . . .

Winter Distribution Map for the Caspian Tern . . . .

Breeding Range Map for the Royal Tern . . . . . .

Winter Distribution Map for the Royal Tern . . . . .

Breeding Range Map for the Sandwich Tern . . . . .

Winter Distribution Map for the Sandwich Tern . . . .

Breeding Range Map for the Roseate Tern . . . . . .

Breeding Range Map for the Common Tern . . . . . .

Winter Distribution Map for the Common Tern . . . . .

Breeding Range Map for the Forster's Tern . . . . .

Winter Distribution Map for the Forster's Tern . . . .

Breeding Range Map for the Least Tern . . . . . .


r


Page

. 91

. 92

* 143

* 157

. 198

. 199

. 302

. 303

. 336

. 337

. 368

. 369

. 394

. 395

S. 420

. 421

. 451

. 485

. 486

. 581

. 582

. . 601









Number Page

23 Breeding Range Map for the Brow Noddy and Sooty Ter . . . 657

24 Breedig ge Map for the Black Skier . . . . . 732

25 WInter Distribution Map for the Black Skimer . . . . 733


Adult Laughing Gull off Hatteras, North Carolina, in August 1981.
Photograph by Roger B. Clapp.









TABLES


Number Page


1 Number and percentage of beached birds examined that were oiled . 16

2 Susceptibility of various larids to oiling as indicated by banding
recoveries . . . . . . . . .. ... .. ...... 17

3 Comparison of regional and seasonal variation of beached bird
mortality and incidence of oiling in the eastern United States 19

4 Comparison of five-year averages of beached bird mortality for
different areas in the eastern United States . . . . .. 20

5 Oil spills in the southeastern United States 1978-1980 compared
with other states . . . . . . . . ... ..... 23

6 Estimated populations of charadriiform marine birds breeding in the
southeastern United States compared with their status in other areas
administered by the U.S. . . . . . . . . ... . 29

7 Approximate number of Red-necked Phalaropes recorded by month for
the coastal southeastern United States . . . . . ... 48

8 Dates of occurrence for Red-necked Phalaropes in the coastal south-
eastern United States . . . . . . . . ... . . 49

9 Approximate number of Red Phalaropes recorded by month for the
coastal southeastern United States . . . . . . .... 72

10 Dates of occurrence for Red Phalaropes in the coastal southeastern
United States . . . . . . . . ... . . .... 72

11 Recent estimates of Laughing Gull populations nesting in the south-
eastern United States . . . . . . . . ... ..... 93

12 Peak concentrations of migrant and wintering Laughing Gulls in the
coastal southeastern United States . . . . . . ... 96

13 Mean clutch sizes reported for the Laughing Gull . . . ... 103

14 Rates of hatching success reported for the Laughing Gull .... .104

15 Weights (in grams) of Laughing Gulls . . . . . . ... 106

16 Approximate number of Franklin's Gulls recorded by month for the
area from North Carolina to Mississippi . . . . . ... 126









Number Page

17 Dates of occurrence for Franklin's Gulls in the coastal southeastern
United States . . . . . . . . ... . . .... .127

18 Peak concentrations of migrant Franklin's Gulls in coastal Texas 127

19 Weights (in grams) of Franklin's Gulls . . . . . ... 132

20 Peak concentrations of wintering Bonaparte's Gulls in the coastal
southeastern United States . . . . . . . .... .145

21 Mean clutch sizes reported for the Bonaparte's Gull . . ... 149

22 Mean clutch sizes reported for the Ring-billed Gull . . ... 168

23 Rates of hatching success reported for the Ring-billed Gull . .. 171

24 Productivity in Ring-billed Gull colonies . . . . ... 173

25 Weights (in grams) of Ring-billed Gulls . . . . . ... 175

26 Herring Gulls nesting in North Carolina . . . . . . 201

27 Laying periods reported at some North American Herring Gull
colonies . . . . . . . . ... . . . ... . 214

28 Mean clutch sizes reported for the Herring Gull in North America 215

29 Comparison of mean clutch sizes for early and late clutches in
North American Herring Gulls . . . . . . . . ... 218

30 Rates of hatching success reported for the Herring Gull in North
America . . . . . . . . ... . . . . . . 219

31 Comparison of hatching success for early and late nests of North
American Herring Gulls . . . . . . . . ... . .221

32 Comparison of hatching success for three- and two- egg clutches in
some studies of North American Herring Gulls . . . . .. 222

33 Productivity in North American Herring Gull colonies ...... 223

34 Weight (in grams) of North American Herring Gulls . . . ... 227

35 Number of dead birds and number and percentage of dead Herring Gulls
found after major oiling incidents . . . . . . ... 229

36 Dates of occurrence for Great Black-backed Gulls in the coastal
southeastern United States . . . . . . . .... .307










Number Page

37 Number of oiled birds and number and percentage of dead Great Black-
backed Gulls found after major oiling incidents ......... 314

38 Recent estimates of Gull-billed Tern populations nesting in the
southeastern United States .................. 342

39 Mean clutch sizes reported for the Gull-billed Tern ....... 349

40 Weight (in grams) of Gull-billed Terns . . . . . ... 351

41 Recent estimates of Caspian Tern populations nesting in the south-
eastern United States . . . . . . . . .. . . 371

42 Recent estimates of Royal Tern populations nesting in the south-
eastern United States . . . . . . . . ... ..... 399

43 Peak concentrations of wintering and migrant Royal Terns in the
coastal southeastern United States ............... 401

44 Mean clutch sizes reported for the Royal Tern . . . . .. 405

45 Weights (in grams) of Royal Terns. . . . . . . . .408

46 Recent estimates of Sandwich Tern populations nesting in the south-
eastern United States . . . . . . . . ... ..... 424

47 Peak concentrations of wintering and migrant Sandwich Terns in the
coastal southeastern United States . . . . . . ... 426

48 Mean clutch sizes reported for the Sandwich Tern ....... 431

49 Rates of hatching success reported for the Sandwich Tern ... 432

50 Weights (in grams) of Sandwich Terns ............. 435

51 Estimated former and present populations of Roseate Terns in north-
eastern North America . . . . . . . . .. . . 459

52 Approximate number of Roseate Terns recorded by month for the coastal
southeastern United States . . . . . . . .... .462

53 Peak concentrations of migrant Common Terns in the coastal south-
eastern United States . . . . . . . . ... ..... 492

54 Mean clutch sizes reported for the Common Tern . . . ... .500

55 Incubation periods (in days) reported for the Common Tern .... .502








Number

56 Rates of hatching success reported for the Common Tern .. . .

57 Fledging success in Common Tern colonies . . . . .

58 Productivity in Common Tern colonies . . . . . .

59 Weights (in grams) of Common Terns . . . .. . . ...

60 Dates of occurrence for Arctic Terns in the coastal southeastern
United States . . . . . . . . . . . . .

61 Approximate number of Arctic Terns recorded by month for the coastal
southeastern United States . . ................ .

62 Peak concentrations of wintering and migrant Forster's Terns in the
coastal southeastern United States . . . . . .

63 Recent estimates of Forster's Tern populations nesting in the south-
eastern United States . . . . . . . . . . .

64 Mean clutch sizes reported for the Forster's Tern . . . . .

65 Weights (in grams) for Forster's Terns . . . . . . .

66 Recent estimates of Least Tern populations nesting in the south-
eastern United States . . . . . . . . .. .

67 Mean clutch sizes reported for the Least Tern . . . . . .

68 Rates of hatching success reported for the Least Tern . . . .

69 Productivity in Least Tern colonies . . . . . . . .

70 Weights (in grams) of Least Terns . . . . . . . . .

71 Approximate number of Bridled Terns recorded by month for the coastal
southeastern United States . . . . . . .

72 Dates of occurrence for Bridled Terns in the coastal southeastern
United States . . . . . . . . . . . . .

73 Numbers of Sooty Terns found breeding in the Chandeleur Islands .

74 Recent breeding sites of Sooty Terns along the Texas coast . .

75 Peak concentrations of Sooty Terns in the coastal southeastern
United States . . . . . . . . . . . . .


Page

503

505

506

509


555


555


580









Number Page

76 Incubation periods (in days) reported for the Sooty Tern . .. 669

77 Rates of aching success reported for the Sooty Tern . . . 670

78 Weights (in gras) of Sooty Terns . . . . . . . . 674

79 Peak concentrations of migrant Black Terns in the coastal south-
eastern United States . . . . . . . . . . 692

80 Mean clutch sizes reported for the Black Tern . .. . ... 699

81 Weights (in grams) of Black Terns . . . .. . . . 700

82 Recent estimtes of Black Skimer populations nesting in the south-
easteo United States ....................... 737

83 Peak concentrations of migrant and wintering Black Ski ers in the
coastal sontheasteo United States . . . . . . . .. 738

84 Weights (in grams) of Black Sl mers .. ............ 746


An iature Laughing Gull. Photograph by Roger B. Clapp.








ABBREVIATIONS USED IN TEXT


Most of the abbreviations used in the text are in standard use and will be
known to the reader; a few may be less familiar. These are listed below with a
brief indication of their interpretation.


N, S, E, W,

N., S., E.,


(capitalized without period)

W. capitalizedd with period)


compass directions

geographic site designation (e.g.,
S. Padre Island


ac
ad.
AOU
BOU
BLM
ca.
CBC
cf.
coll.
comp.
Co.
CSLP
EBBA
ed./eds.
et. seq.
ha
IBBA
imm.
in litt.
in prep.
ms
n
Natl. Park
Natl. Seashore
NERC
nonad.
NWR
OCS
op. cit.
Par.
pers. comm.
pers. observ.
photogr.
prep.
SD
spec.
sp./spp.
St. Park
subad.


acre
adult
American Ornithologists' Union
British Ornithologists' Union
Bureau of Land Management
(circa) about
Christmas Bird Count
(confer) compare/see
collected
compiler
County
Center for Short-Lived Phenomena
Eastern Bird Banding Association
editor/editors
and the following
hectare
Inland Bird Banding Association
immature
in the letters (of)
in preparation
manuscript
sample size
National Park
National Seashore
National Environmental Research Council
nonadult
National Wildlife Refuge
Outer Continental Shelf
(opere citato) in the work cited
Parish
personal communication
personal observation
photographed
preparer
Standard Deviation
specimen
species (singular/plural)
State Park
subadult









subseq. subsequent
unpubl. unpublished
USWS United States Fish and Wildlife Service
USNFW United States National Fish and Wildlife Laboratory
WAGBI Waterfowlers Association of Great Britain and Ireland
WMA Wildlife Management Area

Units of measurement in the text are presented as they were in the source
from which they were derived, and are followed in parentheses by conversion to
the metric or English systems, as appropriate.


n adult Herring Gull. Photograph by J. A. Spendelow.









PURPOSE OF REPORT


The purpose of this report is to summarize the status of 22 coastal and
marine charadriiform species in the southeastern United States and to explore
the potential effects on these species of the development of petroleum re-
sources on the Outer Continental Shelf (OCS). This entailed a review of avail-
able information in order to:

1) determine where and when coastal birds occur in marine areas that may be
developed for oil and gas production;

2) ascertain which species would be most at risk from oil spills and the devel-
opment of oil resources;

3) evaluate the importance of populations in the southeastern United States
in relation to the global distribution and abundance of the species; and

4) summarize information on the life history of the species most likely to be
adversely affected by the development of oil resources.

This material is presented in a form that enables the Bureau of Land Man-
agement (BLM) to identify aspects of OCS development that might threaten popu-
lations of marine birds. It provides information that will aid managers in
making decisions that minimize damage to these populations during the develop-
ment of energy resources. A corollary objective is to recommend topics for
future research in areas where information is particularly scarce.


STUDY AREA


The study area encompasses the coastal and offshore waters of the south-
eastern United States, from the northern border of North Carolina to the Mexican
border of Texas. Many coastal habitats occur within this area: sandy bar-
rier islands; fresh, salt, and brackish marshes; open beaches; coastal bays;
dredge-spoil islands; mudflats; and mangrove islands. The dominant habitats
are discussed below.


HABITATS

North Carolina is dominated by a series of fringing barrier beaches behind
which lie large estuaries with extensive areas of shallow water and salt marsh.
These fringing islands (the Outer Banks) are farther (30-50 km or 20-30 mi)
from the mainland than are such islands along other areas of the Atlantic coast
(Warinner et al. 1976). Extensive stands of salt marsh with deep tidal channels
are found south of Cape Lookout, North Carolina, through South Carolina and
Georgia. Almost three-quarters of the salt-marsh acreage along the Atlantic
seaboard is found in these three states. The largest areas of salt marsh on
the Atlantic coast are in Georgia, which has 193,000 ha (477,000 ac), North








Carolina (64,000 ha or 158,000 ac), and South Carolina (176,000 ha or 435,000
ac) (West 1977).

Barrier islands are also very important coastal habitat in these three
states. The land areas of the barrier islands for each state are 120,000 ac
(49,000 ha) in North Carolina, 124,000 ac (50,000 ha) in South Carolina, and
153,000 ac (62,000 ha) in Georgia (Warner 1976), for a total of about 397,000
ac (161,000 ha). The area of water behind these islands becomes smaller to
the south (Warinner et al. 1976). North Carolina, South Carolina and Georgia
have about 266 mi (428 km), 199 mi (320 km), and 98 mi (158 km) of open beach,
respectively, along their barrier islands. In other parts of the study area
(e.g., parts of the Florida Gulf coast), beaches are few or nonexistent (Wool-
fenden and Schreiber 1973).

The east coast of Florida is dominated by a chain of barrier islands bro-
ken occasionally by tidal passes. Typically, these islands are sandy along
their outer perimeters. Large areas of marsh and estuarine swamp lie landward
of these islands (Warinner et al. 1976) and salt marshes gradually give way
to mangrove swamp as one moves further landward (Reimold 1977). Much of the
Gulf coast of Florida is dominated by salt marshes and mangrove swamps (Wariner
et al. 1976). Extensive stretches of open beach are found from Naples on the
Florida peninsula north along the panhandle to Alabama (Woolfenden and Schreiber
1973). In Alabama, tidal salt marshes, sandy beaches, and offshore islands are
common coastal landforms. Mississippi's Gulf coast consists almost entirely of
barrier islands that have salt marshes in their centers. The Mississippi shore-
line is extensively developed but still contains fresh, salt, and brackish
marshes (Warinner et al. 1976). Only a limited amount of salt marsh is found
between northern Florida and Mississippi. Most marshes are small, disjunct,
and in alluvial pockets protected by bay shores (West 1977).

Louisiana has more marsh and estuarine area than any of the other United
States except Alaska (Warinner et al. 1976), and contains nearly half the total
acreage of salt marsh in the contiguous United States. In some places the
marshes extend inland as much as 40-50 km (25-30 mi) (West 1977). The coast-
line along the western third of the state is sandy, but the rest of the area
is dominated by barrier islands and marsh that are strongly influenced by the
enormous amounts of mud and silt deposited by the Mississippi River (Warinner
et al. 1976). The Louisiana coast is one of the most productive areas for
marine birds in the continental United States and supports enormous wintering
populations of waterfowl.

The coast of Texas makes up a large portion of the western shore of the
Gulf of Mexico. Sandy beaches and offshore barrier islands are abundant.
Two semi-landlocked lagoons, the Upper and Lower Laguna Madre, and a large
low-salinity estuary, Sabine Lake, are areas of great importance to wintering
waterfowl. An estimated 78% of the world's population of Redhead (Aythya amer-
icana) ducks winters in the Laguna Madre, and 13% of the world's shrimp harvest
comes from Texas waters (Warinner et al. 1976). A limited amount of salt marsh
is present in Texas along bay shores enclosed by offshore bars (West 1977).







CLIMATES


The climatic regime, like the landform, differs widely from one part of
the study area to another. The northeastern portion is the coldest. The low-
est midwinter temperatures along the coast of North Carolina are on the order
of 2C0F (-70C) and the average daily maximum during midsummer along the extreme
southern coast is only 860F (300C), some 60F less than is usually recorded in
the interior. July is the wettest month and October the driest. Along the
coast, snow and sleet usually fall only once or twice a year and are usually
associated with northeasterly winds. Prevailing winds in North Carolina blow
from the southwest most of the year and from the northeast in September and
October (Hardy 1974). The weather along the coast of South Carolina is simi-
lar to that in North Carolina with some variation. Average annual temperatures
along the South Carolina coast are about 68oF (20C), with an average daily
maximum in July of 88F (310C) and average daily minimums in January from 35
F (1.7C) in the northeast to 42F (60C) in the southeast. March is particu-
larly rainy along the coast, and October and November are the driest months.
Prevailing winds in South Carolina are from the southwest and south in spring
and summer, predominantly from the northeast in autumn, and from the northeast
and southwest in winter (Landers 1974).

The climate in Georgia is characterized by short mild winters and warm hu-
mid summers. The coastal area becomes progressively drier and warmer from north
to south. Peak periods of precipitation occur in winter and early spring; the
average annual rainfall ranges from 75 in (190 cm) in the extreme northeastern
part of the state to 53 in (135 cm) along the lower east coast. Average summer
temperatures range from 730F (23 C) in the extreme north to 82F (28C) in parts
of south Georgia; average temperature for the three winter months ranges from
41F (5C) in the north to 56F (13C) on the lower east coast. Areas in north-
ern Georgia have freezing temperatures during the day for almost a third of the
year but the lower coast only has about ten days of freezing temperatures annu-
ally (Carter 1974).

Florida has a wider range of climate than any other state in the southeast.
The climate ranges from temperate to subtropical in the north, to tropical in
the Florida Keys. Summers are warm, humid, and long, and winters are mild and
brief. Rainfall is abundant, especially from June to September. Mean annual
temperatures range from the upper 60's (F) in northern Florida to the mid-70's
in the south and reach nearly 780 F (260C) at Key West. Rainfall varies widely
from area to area and from year to year, with most areas usually receiving be-
tween 50 and 65 in (130-170 cm). The drier Keys have an average annual rainfall
of only about 40 in (100 cm). On the southern part of the peninsula prevailing
winds are from the southeast and east; elsewhere they are more erratic and tend
to be from the north in winter and from the south in summer. Tropical storms
frequently cause great damage; few years pass without a hurricane affecting part
of the state (Bradley 1974).

The Gulf of Mexico has a maritime tropical climate with mean winter temper-
atures of about 70F (21C) and mean summer temperatures of 84F (290C). Rela-
tive to seasons in other parts of the study area, both summer and winter are
hot and humid; humidity is greatest during spring and summer, and lowest during







late fall and winter (BLM 1978a). Rain occurs fairly evenly throughout the
year along the western and northern Gulf, with a peak from June through August
(BLM 1978a). Further east in the Gulf the peak tends to be later and falls in
August and September (BLM 1978b). The area becomes progressively wetter from
the southwest to the north and central portions of the northern Gulf. The
driest area of the Texas coast extends from Brownsville north to about Corpus
Christi; the most humid area from Galveston to the Sabine River (Chaney et al.
1978). Average annual precipitation ranges from about 69 cm (27 in) at Browns-
ville to 137 cm (54 in) at New Orleans (BLM 1978a) and 170 cm (67 in) in Mobile
(BLM 1978b).

Tropical storms and hurricanes regularly occur during late summer and fall
and often ravage coastal habitats. Most of these storms enter the Gulf through
the Yucatan Channel and Straits of Florida (BLM 1978a). Southeasterly winds
predominate over the northern Gulf during the summer. Easterlies are more com-
mon during the winter, and prevailing winds from the west and southwest are
rare at any time of year (BLM 1978a).


METHODS


Most of the information was obtained by a literature search. Additional
information on oiling of individual species of birds and their distribution
was obtained through examination of museum specimens and interviews, but these
were not major sources. Several computerized information retrieval systems
were investigated, but did not meet our needs. These sources were particularly
weak on the local distribution of birds, much of which is to be found in region-
al journals not covered by computer services; the temporal coverage was also
inadequate for this study. As Bartonek and Lensink (1978) pointed out, visual
searches of periodicals "proved far more productive from the standpoint of both
numbers of citations and thoroughness of the search."

We obtained literature citations-primarily by scanning the literature and
by consulting bibliographies in relevant papers. The primary sources for the
journals, books, and papers were the libraries and reprint files of the Bird
Divisions of the Smithsonian Institution, Washington, D.C. and the American Mu-
seum of Natural History, New York. Other major sources of information were the
library of the Department of the Interior, the Library of Congress, and the
Bird Library and reprint files of the Patuxent Wildlife Research Center, Laurel,
Maryland. The Welder Wildlife Fbundation, Sinton, Texas, and the library of
government publications and reports maintained by the National Coastal Ecosys-
tems Team, Slidell, Louisiana, were particularly rich sources of information
otherwise difficult to obtain. Unpublished reports and papers were obtained
from: The Florida Audubon Society, Vero Beach; the Florida Fish and Game Com-
mission, Gainesville; Everglades National Park, Homestead; and individuals
listed in the acknowledgments. Several dozen valuable but unpublished theses
were obtained from educational institutions.








Searches were made of several secondary sources of literature citations.
Literature review sections of major ornithological journals, particularly The
Auk, The Ibis, and the Journal of Field Ornithology (Bird-Banding) were espe-
cially useful, as was Wildlife Review. We also made extensive use of Current
Contents, Oil Pollution Abstracts, and Dissertation Abstracts. The Zoological
Record, Biological Abstracts and Ecological Abstracts were also consulted but
were less efficient sources of information. All state bird journals dealing
with the southeastern United States (see list below) were scanned; these jour-
nals, along with American Birds (Audubon Field Notes in earlier volumes), pro-
vided much of the information on local distribution in each state.

We placed considerable emphasis on recentness of information in the lit-
erature search. A few journals (e.g., Wilson Bulletin, Bird-Banding) were
examined for at least 30 years into the past, The Auk from 1930 to the present.
Many others, depending on the degree to which they yielded useful information,
were scanned for only a few recent years. We covered the foreign literature as
thoroughly as possible. Most of the species treated in this report have a wide
geographic distribution, and much of what is known of their breeding biology is
to be found only in foreign periodicals. The linguistic limitations of the
authors, as well as the temporal and fiscal limitations involved in the produc-
tion of this report, precluded full use of this material.

Listed below are the serial publications covered extensively. Where appro-
priate, those areas of the world that these journals cover most thoroughly are
listed in parentheses.


Acta Ornithologica (Poland, U.S.S.R.)
Alabama Birds
Alauda (France, French Africa)
American Birds [Audubon Field Notes]
(United States, Canada)
Animal Behavior
Ardea (western Europe)
Atlantic Naturalist (Delaware to
Virginia)
Atoll Research Bulletin
Auk (North America)
Australian Bird Watcher

Behaviour
Biologia (Bratislava) (Seria B)
(Czechoslovakia)
Biotropica
Bird Study (Great Britain)
Blue Jay (central Canada)
British Birds
Bulletin of the British Ornitholo-
gists' Club (world)
Bulletin of the Kansas Ornithological
Society


Bulletin of the Oklahoma Ornitholog-
ical Society
Bulletin of the Texas Ornithological
Society

California Fish and Game
Canadian Field-Naturalist
Canadian Journal of Zoology
Chat (North and South Carolina)
Colonial Waterbirds (eastern North
America)
Condor (North America, neotropics)
Corella [Australian Bird-Bander]

Dansk Fugle (Denmark)
Dansk Ornithologisk Forenings Tids-
skrift (Denmark)

Ecology
Ekologia Polska (Poland)
Elepaio (Hawaii)
Emu (Australia, New Guinea)
Estuaries [Chesapeake Science]
(U.S. Atlantic Coast)








Florida Field Naturalist
Florida Naturalist
Florida Scientist
Gerfaut (western Europe, Africa)

Hornero (Argentina)
Ibis (Old World, Africa)
Irish Birds

Jack-Pine Warbler (Michigan)
Journal fur Ornithologie (Germany)
Journal of Animal Ecology
Journal of Applied Ecology
Journal of Ecology
Journal of Field Ornithology [Bird-
Banding] (United States)
Journal of Wildlife Management
(North America)

Kingbird (New York)

Larus (Yugoslavia, eastern Europe)
Limosa (Netherlands)
L' Oiseau et la Revue Francaise
d'Ornithologie (France)
Loon [Flicker] (Minnesota)
Louisiana Ornithological Society News

Marine Pollution Bulletin
Maryland Birdlife
Mississippi Kite
Mississippi Ornithological Society
Newsletter
Murrelet (Pacific northwest, Alaska,
western Canada)

Nos Oiseaux (France, western Europe)
Notornis (New Zealand, Pacific islands)

Oikos (Denmark, Scandinavia)
Oriole (Georgia)
Ornis Fennica (Finland, Baltic area)
Ornis Scandinavica (Scandinavia)


Ornithologische Beobachter
(Switzerland, middle Europe)
Ornithologische Mitteilungen (world)
Ostrich (South Africa)

Proceedings of the Annual Conference,
Southeastern Association of Game
and Fish Commissioners (southeast-
ern U.S.)
Proceedings of the Louisiana Academy
of Science

Revue Suisse de Zoologie (Switzerland,
central Europe)
Ring (Europe)
Ringing & Migration (Great Britain)
Rivista Italiana de Ornitologia (Italy)


Scottish Birds
South Australian Ornithologist
Southwestern Naturalist
(southwestern U.S.)
Soviet Journal of Ecology
Sterna (Norway)
Suomen Riista (Finland, Baltic area)

Texas Journal of Science
Tori (Japan)
Transactions of the North American
Wildlife and Natural Resources
Conference

Var Fagelvarld (Sweden)
Vestnik Zoologii (U.S.S.R.)
Vogelwarte (western and central
Europe)

Western Birds (western U.S.)
Wildfowl
Wilson Bulletin (North America)

Zeitschrift fur Tierpsychologie
Zoologicheskii Zhurnal (U.S.S.R.)


The reprint files of several institutions supplied some less easily ob-
tainable material. The most useful of these were files of the Museum Section
of the Denver Wildlife Research Center (formerly the National Fish and Wild-
life Laboratory), the Bird Division of the National Museum of Natural History,
the American Museum of Natural History, and the Bird Library of the Gabrielson








Laboratory of the Patuxent Wildlife Research Center.


In all, about 10,000 citations dealing directly with the species treated
are included in the three parts of this report. The more general articles
found in the Literature Cited sections at the end of each volume will probably
contain at least an additional 1,000 citatons.

Certain data (e.g., weights) reported in this study were obtained directly
from data in the files and collections of museums, particularly those of the
U. S. National Museum and Louisiana State University.


ARRANGEMENT AND CONTENTS OF SPECIES ACCOUNTS


Each volume of this series covers groups of birds for which the problems
of preparing this report have been different, even though all species included
share the characteristic of being at least occasionally found along the waters
of the southeastern United States. Volume 1 (Clapp et al. 1982b) deals with
pelagic seabirds whose status and occurrence in this area are only now being
documented. These species, for the most part, do not breed commonly in south-
eastern waters, and have received little study.

Volume 2 (Clapp et al. 1982c) deals with ducks, geese, and swans, most
of which are important in southeastern waters because large numbers winter or
migrate through this area. Most of these species are covered extensively in
the North American and foreign literature, and much is known about their breed-
ing biology and distribution. Some of them, primarily species that are seldom
hunted (e.g., mergansers, scoters), have been little studied and are among
those species known to be most adversely affected by oil pollution in other
areas. Information on distribution, status, and breeding biology of many
species of waterfowl has been summarized in a number of recent handbooks
(e.g., Bellrose 1976; Palmer 1976a, 1976b; Cramp et al. 1977).

The species accounts presented in volume 3 vary in length and detail with
the quality and quantity of material examined and with the species abundance
and vulnerability to oiling in southeastern waters. The accounts tend to be
longer than those in the earlier volumes because most of the species included
here have been studied more thoroughly and have a more extensive literature
than the species included in Volume I, because few have received monographic
treatment as have those in Volume II, and because most are regularly or season-
ally abundant in the southeastern United States.

Much of the data on the status and biology in the southeast of the species
included here is of recent origin. Much useful information was published while
this report was being compiled (e.g., Duncan and Havard 1980, Rowlett 1980,
Portnoy et al. 1981); still little is known about offshore distribution for
many species. Although we have tried to make this report as timely and thor-
ough as possible, further research probably will reveal the inadequacy of our
present knowledge of the status of many species in southeastern waters.








SPECIES INCLUDED


This report covers 22 members of the order Charadriiformes: 2 phalaropes,
6 gulls, 13 terns, and the Black Skimmer. The birds dealt with here are spe-
cies for which most of the North American (and sometimes world) population
breeds, winters, or migrates through the southeastern United States. Several
species (e.g., the phalaropes, Arctic and Bridled Terns) occur largely as off-
shore or pelagic migrants, others (e.g., Bonaparte's Gull, Common and Black
Terns) occur largely as coastal migrants. Some species (e.g., Herring and Ring-
billed Gulls) breed mostly or entirely north of the study area, but a large
portion of their North American wintering population occurs in the southeastern
United States. Large portions of the populations of several other species
(e.g., Laughing Gull; Royal, Forster's and Least Terns) breed in the southeast-
ern United States. Other species (Gull-billed, Caspian, Sandwich, and Sooty
Terns; Brown Noddy; Black Skimmer) breed in the southeastern United States in
such large numbers that their populations there make up a significant portion
of the North American population.

Some of the species included (e.g., Laughing Gull; Caspian and Royal
Terns; Black Skimmer) with important breeding populations in the southeast
also use the area as a major wintering ground. The Roseate Tern, uncommon to
rare in the southeast, was included because its populations are declining in
many portions of its range and because it has been suggested as a candidate
for listing as an endangered or threatened species.

We originally intended to include accounts for all species of several sub-
families (Stercorariinae, Larininae, Sterninae, Rynchopinae) of the family
Laridae and for all species of murres, guillemots, and other alcids (Alcidae)
that had been recorded at least once in coastal portions of the southeastern
United States. We also intended to include an account for the American Cpot
(Fulica americana) [Rallidae], a species that is abundant in marshy coastal
habitats in many of the southeastern United States. The loss of personnel,
and the loss of use, increased cost of repair, and deterioration of word pro-
cessing equipment and supplies made it necessary for us to delete many species
from this volume and to complete this report at a scale less than originally
envisioned.

The deleted species included the following: American Coot, Pomarine Jaeger
(Stercorarius pomarinus), Parasitic Jaeger (Stercorarius parasiticus), Long-
tailed Jaeger (S. longicaudus), Skua spp. (Catharacta spp.), Little Gull (Larus
minutus), Common Black-headed Gull (L. ridibundus), Band-tailed Gull (L. bel-
cheri), Mew Gull (L. canus), California Gull (L. californicus), Thayer's Gull
(L. thayeri), Iceland Gull (L. glaucoides), Lesser Black-backed Gull (L. fus-
cus), Glaucous Gull (L. hyperboreus), Black-legged Kittiwake (Rissa tridactyla),
Sabine's Gull (Xema sabini), Elegant Tern (Sterna elegans), White-winged Tern
(Chlidonias leucopterus), Black Noddy (Anous minutes), Dovekie (Alle alle),
Common Murre (Uria aalge), Thick-billed Murre (U. lomvia), Razorbill (Alca
torda), and Black Guillemot (Cepphus grylle). The preliminary bibliogra-pies
for these species total about 170 pages and have been retained in the files
of the Museum Section, Denver Wildlife Research Center.








SCIENTIFIC AND VERNACULAR NAMES


Each species account is headed by the common English and scientific name
of the species, followed by vernacular names in other languages and alterna-
tive common English names used in the United States and in other English-
speaking countries.

Scientific and vernacular names follow the 34th supplement (AOU 1982) to
the AOU Checklist (AOU 1957), as does the sequence in which we list orders,
families, and subfamilies. Widely used alternative scientific names are also
noted. Explanation is made in footnotes where changes in scientific names have
been adopted recently.

Scientific names of other organisms (e.g., plants, fish, crabs, molluscs)
given in the text are either those used in the works cited or are from stand-
ard recent references or regional guides. Missing scientific names have been
supplied only when we were certain what species was meant by the vernacular
name used in the original text.

The primary source for most of the non-English vernacular names is the
Nomina Avium Europaearum (Jorgensen 1958); other sources consulted include
Dement'ev and Gladkov (1951), Austin and Kuroda (1953), Edwards (1972), and
Cramp et al. (1977). The abbreviations for the languages and other geograph-
ical uses are as follows:

DA: Danish IC: Icelandic PR: Portuguese
DU: Dutch IT: Italian RU: Russian
EN: English (Old World) JA: Japanese SP: Spanish
FI: Finnish NW: Norwegian SW: Swedish
FR: French NZ: New Zealand US: United States
GE: German PO: Polish

With few exceptions, the foreign common names given are those in widest
use in the ornithological literature of the countries indicated. In several
instances we include transliterated names from languages in which Roman char-
acters are not used (Japanese, Russian). For Japanese names we relied upon
Austin and Kuroda (1953) and For Russian names we used the names used in the
translation of Dement'ev and Gladkov (1951). In some instances more than one
foreign vernacular name for a species has been used. This is particularly
true for Spanish, in which vernacular names may vary considerably from area
to area.

The primary reason for supplying these alternative names is to assist
future computer-based literature searches. Some of the English translations
of foreign names (which are those entered on computers) imply a different spe-
cies than the name would normally suggest to a reader of English or cannot be
readily associated with an English name (e.g., the translation of the Russian
common name for Larus ridibundus is Laughing Gull, a name that in English in-
dicates the Nort X-merican Larus atricilla). In addition, many of the more
regionally-oriented foreign language journals, like those in the United States,










fail to list the scientific names. As a result, searches of computer litera-
ture systems by scientific name alone may fail to indicate important notes
or papers documenting recent changes in distribution.

Alternative scientific names widely or recently in use are supplied as
another aid to literature searches. The Caspian Tern appears in recent liter-
ature as Sterna caspia, Sterna caspius, Sterna tschegrava, Hydroprogne tsche-
grava, and Hydroprogne caspia. One computer search we made revealed no less
than four different lists of titles when each scientific name was used as a
keyword.


GENERAL DISTRIBUTION

This section is divided into two parts, one giving occurrence in North
America, the other giving occurrence elsewhere in the world. Most of this
information has been taken from standard distributional works, but it has been
supplemented, where possible, with more recent literature. Breeding and win-
tering ranges are emphasized in this section, with less information given about
areas of occurrence during migration. Material relating to North America is
more detailed and more complete than for other areas of the world.


DISTRIBUTION AND ABUNDANCE IN THE COASTAL SOUTHEASTERN UNITED STATES

In this section we present more detailed remarks on distribution and
abundance in the southeast. As much recent information through 1979 as pos-
sible has been incorporated. Some accounts provide even more recent infor-
mation but the accounts were completed over a number of years and, as a result,
the depth of coverage is not consistent. This section is based upon the most
recent state ornithological handbooks and checklists; it also includes much
information on seasonal observations published in American Birds and state
journals, breeding data from the Colonial Bird Register at Cornell University,
and data from unpublished manuscripts. This section also incorporates in-
formation on seasonal occurrence, breeding status and numbers, and habitats.
Coastal areas are emphasized, but in some cases status elsewhere is also men-
tioned. Available data for many species are often unsatisfactory, incomplete,
or extremely scanty. This is particularly true for transients and wintering
populations.

Information is given sequentially by state from North Carolina to Texas.
We have not listed states in which a species has not been recorded. Informa-
tion from Florida is usually presented in two subsections, one dealing with
the Atlantic coast, the other with the Gulf. This was done because the status
of a species may vary considerably from coast to coast. In some instances a
section dealing with the Keys is also included when a species' status there
is different from that on either coast.









SYNOPSIS OF PRESENT DISTRIBUTION AND ABUNDANCE


This section in the species accounts summarizes information given in the
previous sections, often with additional data on world-wide population levels
depending upon our present knowledge of the species. For some species tabular
information is presented about seasonal occurrence, abundance, and areas of
concentration in the southeast. Information on these topics is generally
limited and shows where further data should be obtained.

Two types of maps indicate where concentrations of marine birds have been
reported; one deals with breeding birds, the other with wintering populations.
Breeding colony maps are based on highly diverse sources of information, includ-
ing the Colonial Bird Register and a considerable number of published and un-
published censuses (e.g., Portnoy 1977, Parnell and Soots 1979 ms). Other data
were found in recently published papers or were obtained from local ornitholo-
gists. These maps give an estimate of the number of breeding birds and indi-
cate the year or period for which this estimate applies. The largest estimate
available in the last few years is used instead of a range or mean because
estimates are few and because we wish to emphasize areas known to contain
large colonies or concentrations of colonies. Not all data were plotted be-
cause some species occur in so many colonies that it was not feasible to plot
them on our maps. The maps may contain some inaccuracies. They are not intend-
ed as an atlas; their primary purpose is to provide an overview of where concen-
trations of breeding marine birds occur in the southeastern United States.

Most of the winter distribution maps are derived from Bystrak (1974), who
based his maps on an analysis of National Audubon Society Christmas Bird Counts
for one or more of the years from 1970-1972. We chose 45 of 58 coastal Christ-
mas Bird Counts in the study area and compiled 5-year means for 1973-1977. In
some instances fewer than 5 years of counts were available and the mean is for
a shorter period. Localities were chosen to show geographic variation in num-
bers and to emphasize where the largest concentrations were found. The figures
should not be construed to indicate the true size of local populations. The
Christmas Bird Counts vary considerably in the amount of estuarine, coastal,
and marine habitat covered, but we tried to allow for this by choosing counts
that contained the most marine habitat. The numbers reported in any given
year may not be precise because of the limitations of Christmas Bird Counts.
These maps are intended to serve primarily as an index of where winter concen-
trations are most likely to be found and how this distribution varies through-
out the southeast.


HABITAT

This section deals with the nesting, feeding and nonbreeding habitats and
the offshore habitats used by the species. The extent and detail of informa-
tion reported is usually greatest for nesting habitat, less for feeding habitat,
and least for nonbreeding habitats such as those used by migrating or winter-
ing birds. We often could give only sparse information for poorly studied spe-
cies such as the Bonaparte's Gull, and found little adequate information about










offshore occurrences for many common species. Habitats used in the south-
eastern United States have been emphasized when data were available; other-
wise relevant information from other areas is used.


FOOD AND FEEDING BEHAVIOR

We give at least a brief general statement about foods eaten and the
feeding methods. In some instances we include more detailed information on
food habits, briefly abstracting recent studies and indicating proportions of
different foods eaten. For most species few data on food habits in the south-
eastern United States are given, primarily because little is known about the
birds' diet in this area. Consequently, data from other areas have been given
on the assumption that similar foods are eaten in the southeast. Food habits
have been summarized by geographic area for a few species for which much re-
cent information is available. For species whose food habits are well docu-
mented, we pointed out differences in food habits of adults and young, and
commented on seasonal variation and differences in foods eaten in different
habitats.


IMPORTANT BIOLOGICAL PARAMETERS

This section presents basic information to allow biologists to predict
the effects of development of oil resources on populations. The six species
for which we do not give important biological parameters either occur largely
as offshore migrants (Red and Red-necked Phalaropes, Arctic and Roseate Terns)
or are much more abundant outside the southeastern United States (Great Black-
backed Gull and Brown Noddy). Much of the information compiled in this section
is derived from research conducted elsewhere because few adequate studies of
the breeding biology of coastal seabirds have been made in the southeast. For
one species, the Herring Gull, for which there are many data from both the Old
and New Worlds, we have included only information from North American studies.

The data in this section consist of brief summaries of the egg-laying per-
iod, mean clutch size, incubation period, hatching success, age at fledging,
fledging success, mortality of eggs and young, renesting, age at first breeding
maximum natural longevity, and weight. Data on egg laying, incubation period,
and age at fledging allow one to estimate periods when birds breeding within
the study area are most vulnerable to disturbance. Clutch size and hatching
and fledging success are good indicators of productivity. Information on mor-
tality and renesting indicate the potential for recovery following a large
nesting failure. Figures for known maximum natural longevity allow a crude
comparison between species of their total reproductive potential. The maximum
natural longevity is given in terms of "estimated minimum age" in years and
months following Kennard (1975) and Clapp et al. (1982a), and may list informa-
tion based on banding records from the United States, Canada and the Old World.
Finally, information on weights is included because this, and population data
given elsewhere in the report, will allow planners to compare species in terms
of biomass affected by any given oil-related activity.










SUSCEPTIBILITY TO OIL POLLUTION


We emphasize oiling records from southeastern waters but may have missed
reports of oiling for some species. Much of the Old World literature reports
oiled birds only by species groups (e.g., gulls, ducks) and combines informa-
tion on individual species in these totals. Some information may be found in
Old World regional periodicals unavailable in the United States and not cover-
ed by computer-based literature retrieval systems. For each species we also
provide an estimate of the potential effect of oil pollution and the develop-
ment of oil resources in the southeast, taking into account the known or sus-
pected vulnerability of the species, its abundance in the southeast, and its
abundance elsewhere.


SPECIES BIBLIOGRAPHY

At the end of each species account is a species bibliography which in-
cludes citations that provide additional data on the topics briefly covered
in the text, as well as on various'other aspects of the biology of the species.
All citations in the text appear in the Literature Cited at the end of this
report.

The bibliographies in this volume are not exhaustive, but cover the world
literature because little is known of the biology of marine birds in the coast-
al southeastern United States and because more extensive bibliographies should
be helpful in planning future research on these birds. As in the preceding
volumes in this series (Clapp et al. 1982b, 1982c), the bibliographies stress
distribution, ecology, and behavior of the species but provide more information
on topics such as identification, hybrids, diseases, and the effects of pesti-
cides.

Our search of the literature stressed recentness of information. Each
species bibliography should be relatively complete through 1981 and mid 1982.
Some references published more recently may be included, but these and other
references from 1981 and 1982 may not have been used in writing the account.
We also listed other important papers dealing with the biology of the species
through the early part of the century, and included older references that are
still the major sources of information on the species. We were more complete
with papers written in English.

The species bibliographies are arranged from present to past with authors
listed alphabetically under each year, rather than in the more conventional
alphabetical and chronological listing used in the Literature Cited. We ar-
ranged them this way to make it easier for the reader to find the most recent
information. We checked all references used in the text of the accounts as
well as most of the remaining references. Some citations from secondary
sources remain unchecked. We estimate that the three volumes in this series
contain on the order of 10,000 references, and our temporal and fiscal limita-
tions were too great to allow complete verification of all references.










OIL POLLUTION AND MARINE BIRDS OF THE SOUTHEASTERN UNITED STATES


With the possible exception of marine turtles, marine birds are the ver-
tebrates most severely threatened by oil pollution and the development of oil
resources. The work of Old World biologists presents clear evidence of sub-
stantial damage to several populations of marine birds. Specific information
on the effects of oiling and oil spills on avian populations in the New World
is very limited, especially so for populations in the southeastern United
States. Systematic surveys of beached birds and seabird kills have begun only
recently in the United States. Further, data on oiling of marine birds are
scattered through a diverse body of literature. Many distributional notes re-
porting the first specimen of a species from a geographic locality parentheti-
cally note that the specimen was oiled. Other information is scattered through
regional distributional works and yet more data lie in the banding and recovery
files of the Bird Banding Laboratory of the U.S. Fish and Wildlife Service.

Oil pollution kills many thousands of marine birds each year in Denmark
(Riisgard 1979) and elsewhere. During the winter of 1980-1981, five oiling
incidents in northwest Europe (Mead 1981) led to an estimated 60,000 oiled
birds on the beaches (Mead and Baillie 1981). Most of these birds were vari-
ous species of auks, but many Black Scoters (Melanitta nigra) were killed in
Western Denmark and many live, oiled Black-legged Kittiwakes (Rissa tridactyla)
were seen in Holland and Belgium (Mead and Baillie 1981). These spills proba-
bly killed more marine birds in western Europe than had any incidents during
the previous 12 years (Anon. 1982a). Another recent spill that washed ashore
on the Magdalen Islands in the Gulf of St. Lawrence off eastern Canada killed
more than 1,500 seabirds, most of them evidently Dovekies (Alle alle) and
murres (Uria spp.) (Anon. 1982b).

Oiling has been a major factor in reducing populations of Common Eiders
(Somateria mollissima) in the Danish Waddensea (Joensen 1973), Common Eiders
and Black Scoters in Holland (Swennen and Spaans 1970), and Atlantic Puffins
(Fratercula arctica) in France (Bourne 1976). Other examples of significant
reductions in avian populations due to oil pollution are given in reviews by
Bourne (1968a, 1976), Croxall (1975), Vermeer and Vermeer (1975), and the Food
and Agricultural Organization of the United Nations (1977).

Smaller but significant losses have also been recorded in localized popu-
lations. An estimated 25-50% of the Common Loons (Gavia immer) wintering in
Shetland, off Scotland, died following the ESSO BERNICLA oil spill (Stowe and
Morgan 1979). All local Mallards (Anas platyrhynchos), European Coots (Fulica
atra), and Moorhens (= Common Gallinule, Gallinula chloropus) died following an
oiling of the Amer River in the Netherlands; it was estimated that approximately
88% of the Greylag Geese (Anser anser) and 71% of the Bewick's Swans (Cygnus
columbianus bewickii) also were lost (Belterman 1972). During the winter of
1981-82 an oil spill in Belgium killed about 300 birds and affected as many
as 3,000, mostly gulls and the entire local population of Little Grebes (Tachy-
baptus ruficollis) (Anon. 1982c).










Despite continuing reports of high seabird mortality from oil pollution,
particularly in the colder waters of the North Atlantic, current opinion is
that the long-term effects of oil spills on the environment may be scant. The
Royal Commission on Environmental Pollution (1981 in Bourne 1982) stated that
"We have concluded that oil spills are unlikely to cause long-lasting damage;
the environment has a remarkable capacity for recovering from even the largest
oil spillages. We have found no clear evidence that oil spills have signifi-
cantly affected populations of sea birds or other marine species . The
short term consequences of oil pollution are serious . and are typified
by the pollution of beaches on the one hand, and damage to seabirds on the
other . ." Clark (1982) pointed out that although tens of thousands of
marine birds die annually from oil pollution in western Europe, natural mor-
tality accounts for hundreds of thousands per year. Nonetheless, Nettleship
(1977) believed that oil pollution of the sea still constitutes the single
largest threat to seabirds.


VARIABILITY AMONG SPECIES IN SUSCEPTIBILITY TO OIL POLLUTION

Surveys of beached birds are biased indicators of the proportion of a
population affected by oiling (Bourne 1976, Powers and Rummage 1978). Recent
studies by Manomet Bird Observatory (Powers and Rumage 1978, Powers et al.
1980) documented differences between the species composition of birds found
oiled on beaches and birds found oiled at sea. In December 1976 and in Janu-
ary 1977, 181 oiled birds were found on the beaches of Nantucket Island and
Martha's Vineyard, Massachusetts, following the spill of the ARGO MERCHANT.
Most of these birds were alcids (48.1%), gulls (27.1%) and loons (18.2%), but
7 sea ducks, 2 Northern Gannets (Sula bassanus), 2 cormorants, and 1 grebe
were also found. Most (92% of 1120) of the birds seen offshore were gulls;
of these, 59%, 41% and 9%, respectively, of the Herring Gulls, Great Black-
backed Gulls, and Black-legged Kittiwakes (Rissa tridactyla) were oiled. Both
Northern Gannets and Northern Fulmars (Fulmarus glacialis) were seen offshore
more frequently than observed on the beaches, making up 6% and 1%, respective-
ly, of the birds seen offshore. Twelve percent of the gannets seen and at
least some of the fulmars were oiled. Alcids made up only 12% of the birds
seen, but only 12 oiled Dovekies (Alle alle) were observed. Powers and Rumage
(1978) pointed out, however, that oil on the larger alcids could have been over-
looked easily.

Although beached bird surveys (Table 1) give some idea of differences in
susceptibility to oiling between groups of birds, the recent work by Powers
et al. (1980) suggests how different the situation may be offshore. Offshore
surveys may be more reliable for indicating the magnitude of the oil pollution
problem in a given area and for providing useful data on the incidence and ex-
tent of oil pollution. Species such as loons, grebes, auks, and sea ducks are
frequently killed by oil. Others, such as gulls, are frequently oiled but die
much less often from the direct effects of oiling (Table 2). Terns (Table 1,
Bourne in litt.) are seldom found on beached bird surveys, and if they do die
are perhaps less likely to be found. They evidently seldom die from the direct
effects of oiling (Table 2). Black Skimmers are probably intermediate between










Table 1. Number and percentage of beached birds examined that were oiled (a).


South Oregon-
Great Atlantic Coast Washington California
Kinds of birds Britain United States Coast (b) Coast


Total % Total % Total % Total %
found oiled found oiled found oiled found oiled

Loons (Divers) 152 94 114 4 3 33 175 10
Grebes 54 59 14 64 14 36 798 5
Albatross -- 0 -- 0 -- 8 0
Petrels (c) 337 17 0 -- 2 50 0? -
Northern Fulmar -- 0 -- 570 28 301 4
Shearwaters 14 0 0 -- 623 22
Storm-petrels -- 0 -- 4 25 40 0
Gannets 182 50 6 17 -
Cormorants 218 45 6 0 0 -- 13 0.5
Brown Pelican 17 0 -- 38 0
Wildfowl 1137 76 51 4 26 92 296 7
Phalaropes 0 -- -- 119 3
Jaegers 1 0 0 -- 8 0
Kittiwakes -- 0 -- 105 21 33 24
Gulls 2448 30 131 0 16 31 1197 2
Terns -- 37 0 0 -
Skimmer -1 0 --
Auks 6171 80 0 -- 104 94 2848 19

(a) Data for Great Britain, the south Atlantic coast of the United States,
the Oregon-Washington coast, and the California coast are from Bourne
(1976), Malcolm Simons (in litt.), Harrington-Tweit (1979), and Ainley
(1976), respectively; the periods covered are 1968-1970, December 1977-
August 1978, mid-winter 1976, and 1971-1975, respectively. Data for
the southeastern coast through 1 December 1977 are based on surveys
from Cape Hatteras, North Carolina, to Cape Canaveral, Florida, there-
after south to Jensen Beach, Florida.
(b) Most mortality of fulmars and Black-legged Kittiwakes following this
incident was not believed to be due to oil; most wildfowl and alcid
mortality was attributed to oiling (Harrington-Tweit 1979).
(c) Although Bourne (1976) did not specify, his term 'petrels' probably
included all Procellariidae (e.g., petrels, shearwaters, fulmars)
and may have included Hydrobatidae (storm-petrels). His term 'gulls'
probably included all Laridae (gulls and terns). For other material
summarized here, 'petrels' refers to Pterodroma, shearwaterss' to
Puffinus, 'gulls' to Larus, and 'terns' to Sterninae.










Table 2. Susceptibility of various larids to oiling as indicated by banding
recoveries (a).


Number Percent
Number dead dead
recov- from from Areas from which dead oiled
Species ered oiling oiling larids were reported (b)


Laughing Gull 2,269 4 0.18 New Jersey, Louisiana, Texas,
(Larus atricilla) Panama
Ring-billed Gull 23,531 19 0.08 Ontario (4), New York (3), Florida,
(Larus delawarensis) North Carolina, Michigan (2)
California Gull 2,658 6 0.23 California (4), Washington (2)
(Larus californicus)
Herring Gull 34,713 58 0.17 New York (10), Massachusetts (8),
(Larus argentatus) Ontario (5), Newfoundland, Texas (4)
Western Gull 2,916 8 0.27 California (6), Washington, Oregon
(Larus occidentalis)
Glaucous-winged Gull 18,979 23 0.12 British Columbia (9), Washington (7),
(Larus glaucescens) Oregon (4)
Great Black-backed Gull 1,426 3 0.21 New York (2), Quebec
(Larus marinus)
Caspian Tern 2,390 3 0.13 California (2), Michigan
(Sterna caspia)
Royal Tern 4,230 5 0.12 Virginia, North Carolina,
(Sterna maxima) South Carolina, Florida
Common Tern 56,067 14 0.03 Caribbean (5), Brazil (2),
(Sterna hirundo) Venezuela (2), New Jersey (2)
Sooty Tern 30,461 1 0.003 Florida
(Sterna fuscata)
Brown Noddy 942 1 0.11 Caribbean
(Anous stolidus)
Black Skimmer 927 2 0.22 Florida
(Rynchops niger)

a) This table is derived from a printout produced by the Bird Banding Laboratory
that listed all birds whose recovery through 11 April 1978 was believed due
to oiling, and from Clapp et al. (1982a) which gives the number of recover-
ies for each species processed by the BBL through December 1981. The per-
centages may underestimate the numbers recovered due to oiling, but suggest
the degree to which oiling causes direct mortality. No Franklin's Gulls
(250 recoveries), Roseate Terns (1,286), Arctic Terns (291), or White Terns
(Gygis alba) (376) were reported dead from oiling.
b) For species recovered in large numbers we only list areas from which the
most oiled birds were reported.










these two groups in their susceptibility to oiling.


REGIONAL AND SEASONAL DIFFERENCES IN OILING AND MORTALITY OF BEACHED BIRDS

Although beached bird surveys in the eastern United States have been con-
ducted for only a short time, the extent of oiling among birds found dead along
the southern Atlantic coast appears low compared with other areas in the United
States and Europe. Only 4% of 400 birds found dead along the southeastern
Atlantic coast from January 1976 through August 1978 were oiled. In contrast,
oiling occurred on 82% of 667 birds found along the Polish Baltic coast from
November 1974 to August 1975 (Gorski et al. 1977), on 26% of 162 found along
Irish coasts from December 1977 to March 1978 (O'Keeffe 1978), on 79% of 3,431
found on the international beached bird surveys in Northwest Europe in January-
March 1975 (Lloyd 1976), and on 18% of 2,420 found along the California coast
in 1975 (Ainley 1976).

Bird mortality per mile of beach also tends to be less in the southeast-
ern United States than in other areas (Table 3). Mortality figures for a
heavily polluted area, the Polish Baltic coast (3.2 birds/km or 5.1 birds/mi;
Gorski et al. 1977), are considerably higher than for anywhere in the south-
east. Other areas in northwestern Europe vary considerably in mortality re-
corded during beached bird surveys; these mortalities are usually greater than
those found in the southeastern United States, but resemble those found along
the North Atlantic coast of the United States. Lloyd (1976) reported a range
in the number of dead birds found of 0.17 birds/km (0.3 birds/mi) in part of
France to 4.06 birds/km (6.5 birds/mi) in West Germany during the winter of
1975. For Great Britain, 1968-70, the average was 1.3 birds/km (2.1 birds/mi)
(Bourne 1976), similar to mortality in the southeastern United States. Mortal-
ity during 1976-1980 along the southeastern Atlantic coast and Florida Gulf
averaged 1.3 birds/mi (0.79 birds/km) and 1.2 birds/mi (0.77 birds/km), respec-
tively (Simons 1981 ms). Mortality along the California coast is also greater
than in the southeast; surveys there reported an average of 3.5 birds/mi (2.2
birds/km) from 1971 to 1975 (Ainley 1976). The disparity between beached bird
mortality rates in California and Europe and the southeast may arise partly
from differences in prevailing winds and currents. In parts of North America
where prevailing winds blow offshore, most mortality is found around enclosed
inlets. On islands offshore in western North America and in northwest Europe,
where prevailing winds carry dying birds (and oil) to shore, both chronic oil
pollution and the recorded mortality of marine birds is greater (Bourne 1976).

The colder winter climates in these areas are also almost certainly a fac-
tor in the increased mortality. Both oil pollution and bird mortality are
greatest in northwest Europe during the coldest part of the year (Bourne 1968a).
Oiling (and presumably mortality) of marine birds off the north Atlantic coast
of the United States is greatest during winter and spring (Powers et al. 1980);
mortality is also greater during winter and spring along the south Atlantic
and Gulf coasts (Tables 3, 4).










Table 3. Comparison of regional and seasonal variation of beached bird
mortality and incidence of oiling in the eastern United States (a).


Atlantic Coast Atlantic Coast
N of Cape S of Cape Florida Gulf Texas Gulf
Hatteras Hatteras Coast Coast

Dead Dead Dead Dead
birds/ % birds/ % birds/ % birds/ %
Period mile oiled mile oiled mile oiled mile oiled


SPRING


Mar.-May
Mar.-May
Mar.-May
Mar.-May
Mar.-May
Mar.-May


SUMMER


June-Aug.
June-Aug.
June-Aug.
June-Aug.
June-Aug.


FALL


Sep.-Nov.
Sep.-Nov.
Sep.-Nov.
Sep.-Nov.
Sep.-Nov.


WINTER


Dec.-Feb.
Dec.-Feb.
Dec.-Feb.
Dec.-Feb.
Dec. -Feb.
Dec.-Feb.


1982
1981
1980
1979
1978
1977



1981
1980
1979
1978
1977



1981
1980
1979
1978
1977



81-82
80-81
79-80
78-79
77-78
76-77


2.19
1.48
1.52


2.50



2.10
4.27
4.40
6.37
6.81



2.17
1.48
0.98
1.05
0.24



2.51
1.76

2.19
2.70
9.33


6
34.3
0.0
51.4
66.8 (b)
5.5


4

13.4
0.0
0.0



3.9
12.5

2.3
6.5
5.5


1.32
.57
0.55
0.92
1.58
0.95


1.07
1.04
1.43
1.49
0.60



1.54
.29
.57
1.84
2.87
1.75


0.0
7.7

20.0
0.0
0.0


2

0.0
0.0
0.0



0.9
0.0

1.1
1.4
0.0


2.14
1.00
2.75
.82
1.77
.75



0.27
.39
0.53
1.30
0.50



1.47
1.00
0.59
1.00
1.25


1.39
1.56
1.74
3.40
2.88


0.0

0.0

0.0


0.0

0.0
5.6
0.0



0.0
0.0

0.0

0.0


3.33
2.95


2.00






2.80
3.00






0.40

0.36
4.00
0.75







1.00
1.00


0.0
11.1


0.0


0.0
100.0 (c)


0.0
0.0










Table 3. Concluded.


(a) This comparison is based on Bulletins of the Atlantic and Gulf Coast
Beached Bird Survey Project and Simons (1981 ms, in litt.). These data,
while useful, are incomplete and have sometimes (particularly in Texas)
been based on surveys of so few miles of beach that the results obtained
are probably not comparable from region to region. Dashes indicate
lacking data. Several figures from the first two volumes of this series
have been corrected and some additional information is given.
(b) This high figure is the result of an oil spill in the Chesapeake Bay
in February 1978.
(c) This high figure is presumably the result of the IXTOC oil spill in the
southern Gulf of Mexico.



Table 4. Comparison of five-year averages of beached bird mortality for
different areas in the eastern United States (a).


Dead birds found per mile on the:

Five- Atlantic Atlantic Average
year Coast N Coast S Florida for
period of Cape of Cape Gulf entire
Season ending Hatteras Hatteras Coast area



Spring 1982 2.79 1.38 1.12 2.08
(Mar.-May)
Summer 1980 6.31 0.61 0.83 ---
(June-Aug.)
Fall 1980 0.80 1.22 0.93 1.00
(Sep.-Nov.)
Winter 1981 3.70 2.57 2.25 3.04
(Dec.-Feb.)

(a) These figures are compiled from Bulletins produced by the Atlantic and
Gulf Coast Beached Bird Survey Project. Too few data are available for
the Texas Gulf coast for these to be meaningful. The high average mor-
tality in summer in the northeastern United States is partly the result
of disturbance of nesting colonies.










Data on occurrence and seasonal variation of oiled birds at sea are large-
ly unavailable, but a preliminary report by Powers et al. (1980) of at-sea
observations from February 1976 to December 1979 in the Georges Bank area off
Massachusetts provides some useful information. The majority of oiled birds
seen were gulls. In some months (January and February) gulls made up nearly
all oiled birds seen, but very few oiled gulls were seen during the summer
(June-August). Oiled Northern Fulmars were regularly seen in spring and sum-
mer. Oiled Greater Shearwaters (Puffinus gravis) were seen in June and Octo-
ber. Oiled sea ducks (Common Eider [Somateria mollissima], Red-breasted Mer-
gansers [Mergus serrator], White-winged Scoters [Melanitta fusca] and Old-
squaws [Clangula hyemalis]) were seen in December and March. Northern Gannets
and Pomarine Jaegers (Stercorarius pomarinus) were rarely oiled.


BIRD KILLS FOLLOWING OIL SPILLS IN THE SOUTHEASTERN UNITED STATES

The few reports of large bird kills following oil spills in southeastern
waters are usually inadequate. A typical example occurred in late December
1968, when a barge spilled crude oil along the coast of Wakulla County, Flor-
ida. This resulted in "many ducks snipe and other birds so covered with oil
that they were unable to fly. Smaller birds were unable to walk in the heavy
oil" (Center for Short-Lived Phenomena 1969). Reports of more recent spills
are no better. During the spring of 1981, 39 oiled Common Loons (Gavia immer)
were picked up on the beaches of Pea Island NWR, North Carolina. Despite
treatment, only 2 survived (Simons 1981). How many more died at sea or what
other species were affected is unknown. A "large toll" of wintering Common
Loons and Red-throated Loons (Gavia stellata) was taken by an oil spill near
Myrtle Beach, South Carolina (Anon. 1981). In yet another incident, more than
175,000 gallons of heavy fuel oil spilled into a marsh on the Cape Fear River
in South Carolina. This spill was followed by the indirect oiling of Brown
Pelican (Pelecanus occidentalis) eggs when oiled adults returned to incubate
their eggs. Whether this oiling affected the colony or other waterbirds nest-
ing nearby is not known (Anon. 1982d).

We know of only two instances of major oil spills in or near the south-
eastern United States for which there is even fair information about the num-
ber and species of birds killed. The first of these occurred in early Febru-
ary 1976 in the lower Chesapeake Bay. Following the sinking of a barge near
the mouth of the Potomac River, spilled fuel oil resulted in the widespread
contamination of marshes and beaches, and the death of an estimated 20,000 to
50,000 birds (Roland et al. 1977). Perry et al. (1979) made estimates for
each species that died during this spill, five spills that occurred in the
Delaware River, and another large spill in Chesapeake Bay. Most of the vic-
tims of the seven spills were ducks (69.02% of ca. 52,500 birds), Horned Grebes
(Podiceps auritus) (27.75%), and Common Loons (1.30%). Few charadriiforms
died (315 Herring Gulls, 30 Great Black-backed Gulls, 10 Ring-billed Gulls,
and one American Oystercatcher (Haematopus palliatus).










The second major mortality followed the spill in Tampa Bay of some 80-100
tons of oil when the Greek tanker DELIAN APOLLON ran aground in mid-February
1970 and ruptured its hull (Wallace 1970, Clark 1973). Oil spreading over the
bay resulted in at least 4,500 birds being brought to cleaning and rehabilita-
tion stations (Sims 1970). As many as 9,000 birds may have died (Clark 1973).
The birds brought to cleansing stations were largely ducks, Common Loons, and
Horned Grebes (Sims 1970).

Several of the southeastern states are among those with the greatest num-
ber of oil spills and amount of oil entering the ecosystem yearly (Table 5).
However, so little information on the effects of oiling on seabirds of this
area is being recorded and reported that it is impossible to assess adequately
the overall effect of oil pollution in the southeast.


SOURCES OF VARIATION IN MORTALITY FROM OIL POLLUTION

A large number of factors are involved in determining the magnitude of
detrimental effects of oil pollution on marine birds. Birds oiled in cold
weather or cold water have a much higher fatality rate than do those in warm
weather and warm water. Even small amounts of oil may lead quickly to death
under the stress of cold (Levy 1980), but birds in warmer areas may survive
the same degree of oiling (R. Clapp, pers. observe ; C. Harrison, pers. comm.).
Reports from Europe (Bourne and Bibby 1975, Riisgard 1979) indicate that mor-
tality from oiling is greater in winter than in summer.

Oil spilled in cold water remains liquid longer than in warm water and is
likely to cause more damage as a result. It first forms a "chocolate mousse"
water-in-oil emulsion and then it forms tar balls. Although these forms of
oil may present some hazard to birds (Bourne and Bibby 1975), the hazard is
apparently much less than with fresh oil.

Bourne (1976) summarized some of the changes in daily, annual, and life
cycles of marine birds that may increase their vulnerability to oil pollution.
Local currents and winds may bring drifting slicks into rafts of birds roost-
ing on the water. Bourne and Devlin (1969) suggested that most mortality from
oiling occurs when roosting or feeding birds are trapped by drifting slicks.
Birds that concentrate in flocks of 1,000 or more to scavenge behind fishing
trawlers (e.g., gannets and gulls [Powers et al. 1980]), may be drawn into
the area of an oil spill, causing disproportionately large numbers to be oiled.
Conversely, oiled birds may follow trawlers away from the site of a spill and
thus be underrepresented among the sick and dying coming ashore (Powers and
Rumage 1978).

Bourne (1976) pointed out that marine birds are particularly susceptible
to damage from oil when they are molting. When birds lack their usual insula-
tion, smaller than usual amounts of oil may lead to death from chilling, shock,
and starvation.











Table 5. Oil spills in
with other states (a).


the southeastern United States 1978-1980 compared


Percent
Percent Percent of oil Percent
Number of spills of all Volume spilled of oil
of oil occurring U.S. spilled in the spilled
spills in the oil (gallons) south- in U.S.
State 1978-80 southeast spills 1978-80 east waters


Southeastern
states

North Carolina
South Carolina
Georgia
Florida
Alabama
Mississippi
Louisiana
Texas

Total-Southeast

Other heavily
polluted states

New Jersey
Massachusetts
New York
California
Kansas
Kentucky
Missouri
Rhode Island
Alaska
Pennsylvania

Total-U.S. Waters


297
282
259
2,018
434
151
5,664
3,350


2.4
2.3
2.1
16.2
3.5
1.2
45.5
26.9


1.0
0.9
0.8
6.6
1.4
0.5
18.4
10.9


139,165
105,791
474,206
507,693
518,829
65,750
5,325,747
3,662,896


1.3
1.0
4.4
4.7
4.8
0.6
49.3
33.9


0.4
0.3
1.5
1.6
1.6
0.2
16.5
11.3


12,455 100.1 40.5 10,800,077 100.0 33.4


1,052
792
1,081
2,249
120
508
292
253
798
800

30,777


3,921,308
1,992,655
1,334,970
1,083,129
901,096
861,329
809,987
700,105
639,860
614,129


12.2
6.2
4.2
3.4
2.8
2.7
2.5
2.2
2.0
1.9


- --- 32,207,039


(a) This table is compiled from U. S. Coast Guard (1980, 1981) data. Other
heavily polluted states chosen for comparison were those that had the greatest
amount of oil spilled from 1978 through 1980. The figures for 1980 were pre-
liminary figures only. The figure for Texas probably is somewhat inflated due
to the IXTOC blowout.










Breeding birds are particularly susceptible to oil. The loss of one mem-
ber of a pair may mean complete loss of their reproductive potential for that
year. Although this loss may be recouped in future generations, most marine
birds have low productivity and populations take years to recover from a single
oiling incident. Oil in the vicinity of breeding colonies also may diminish
reproductive success by decreasing the hatching success of contaminated eggs,
by disturbance to the colony from attempts to control pollution (Bourne 1976),
and by debilitating oiled birds so much that they may not attempt to breed
(Stowe 1982).

Ford et al. (1982) recently developed a mathematical model that dealt with
the effects of oil spills on breeding populations of seabirds, using guillemot
and kittiwake populations in the Bering Sea as paradigms. Their model suggests
that a catastrophic mortality of adults requires a longer recovery time for a
population than does a similar mortality of young. A complete breeding failure
by guillemots in one year would have a smaller effect on recovery time than
would a 5% mortality of adults. Although these remarks are based on alcids, a
group highly susceptible to oil pollution, the authors pointed out that their
results probably are applicable to other species with high adult survival rates.
High adult survival rates have been shown for several species that occur in the
southeast, Northern Fulmar, Manx Shearwater (Puffinus puffinus), Herring Gull
(Dunnet 1982), and Common Tern (DiCostanzo 1980), and are likely to occur in
species of gulls and terns breeding in the southeast.

Ford et al. (1982) also modeled the effects of chronic low-level pollu-
tion following a one-time adult mortality (as might be caused by an oil spill).
Their results indicated that small decreases in adult survival or fecundity
greatly increased the amount of time required for the affected population to
recover. They also pointed out that "Chronic low-level pollution may produce
permanent changes in the survivorship and fecundity schedules of a population,
which can alter the recovery time and extinction point for one-time perturba-
tions." Ford et al. (1982) concluded by stating that their model should be
regarded as no more than a preliminary estimate that predicts only the magni-
tude of a population's response to an oil spill. They indicated that more
precise predictions are not possible without adequate data.

Few observations of the behavior of birds encountering oil have been re-
ported. Available information indicates that differences in behavior between
species affect their level of vulnerability to oil. According to the Inter-
nation Council for Bird Protection (1960), Long-tailed Ducks (= Oldsquaw,
Clangula hyemalis) choose to land on oil slicks. If true, this may account
for some of the very high oil-related mortalities reported for this diving
duck. Guillemots (= Common Murres, Uria algae) dive to escape floating oil
but suffer the risk of surfacing into it and thus becoming severely contami-
nated (Bourne 1968b). Hainard (1959) reported that some diving ducks (Tufted
Duck [Aythya fuligula] and Pochard [A. ferina]) avoided patches of oil floating
down a river. Gulls (Bourne 1968b) and Manx Shearwaters (Puffinus puffinus)
(Casement 1966) also actively avoid thicker, more noticeable oil slicks when
flying. Some of these birds also avoid oil when swimming; a Herring Gull and
a Black-legged Kittiwake (Rissa tridactyla) that swam into a patch of floating










oil immediately took flight (Bourne 1968b, Bourne and Devlin 1969). Sander-
lings (Calidris alba) and Willets (Catoptrophorus semipalmatus) oiled on the
Texas coast spent less time feeding and more time roosting and making comfort
movements than did unoiled birds (Chapman 1981).

The number of birds that die following an oil spill is also related to
the type of petroleum spilled and how long it remains in the environment.
Crude oil is less toxic than refined oils (diesel oil, No. 2 fuel oil, Bunker
"C") (Hay 1979), and fresh oil causes more damage than older, weathered oils
(Bourne and Bibby 1975). Some oils may be innocuous enough that oiled birds
are not killed and are even capable of cleaning their plumage (Birkhead et al.
1973, Phillips 1974).

The number of deaths from oiling following a spill is not necessarily
related to the amount of oil spilled; large spills may result in few deaths,
while smaller spills may cause large losses, particularly when substantial
numbers of birds are concentrated in small areas (Croxall 1975, Salomonsen
1979). In addition, large oil spills may cause no greater loss of marine
birds than does chronic oil pollution of the environment (Nelson-Smith 1973,
Croxall 1975, Holmes and Cronshaw 1977).


EFFECTS OF OIL ON CONTAMINATED BIRDS AND THEIR EGGS

The primary effect of oil on birds is a loss of buoyancy and insulation
when the plumage becomes matted. Oiled birds then may quickly chill and die
from exhaustion and exposure. Ingestion of oil by birds also leads to a
variety of physical and physiological disorders that make death more likely
(Clapp et al. 1982c, Eastin and Murray 1981). Experimental studies also have
dealt with the secondary effects of oiling on reproduction of marine birds.
Small amounts of oil will reduce the hatching success of duck, heron, gull,
tern, and auklet eggs (Eastin and Hoffman 1978, Stickel and Dieter 1979, Ain-
ley et al. 1981, Hoffman and Eastin 1981).

Toxicity of oils is greater for new eggs than for those further along in
incubation, and older weathered oils are less toxic than fresh ones. Oiling
of incubating gulls causes significant egg mortality when the oiled feathers
come in contact with the eggs. Oiling of eggs also may result in deformed
chicks. Deformed bills, incompletely ossified wing or foot bones, abnormally
small liver lobes and stunting were the most common abnormalities found in
experimental studies (Stickel and Dieter 1979).

Other studies have shown that the number of eggs laid by Mallards (Anas
platyrhynchos) decreases when they are fed diets containing 2.5% South Louisi-
ana crude oil (Eastin and Hoffman 1978, Stickel and Dieter 1979). Ducklings
fed diets containing 5% of this crude oil grow more slowly than usual and may
develop a number of internal disorders (Eastin and Hoffman 1978). These
amounts of oil seem large, but other work has shown that smaller doses, simi-
lar to what might be encountered in nature, may also have adverse affects on
health and reproductive biology.










Herring Gull chicks at Little Duck Island, Maine were fed 1 ml doses of a
crude oil/corn oil mixture. Chicks fed 0.2 and 0.5 ml of weathered South Lou-
isiana crude oil grew more slowly for 7-9 days after treatment than did con-
trols fed only 1 ml of corn oil (Butler et al. 1978). Chicks fed 0.2 ml of
crude oil recovered within two weeks, but those fed 0.5 ml showed a reduced
gain in weight for about three weeks after the initial dosing. Both groups
fed crude oil also exhibited decreased culmen growth. No differences in be-
havior were noted.

Peakall et al. (1978) tried to determine which compounds in crude oil
inhibited growth. Herring Gull chicks were fed 1 ml of two South Louisiana
crude oils, or the aliphatic or aromatic fractions contained within 1 ml of
each. One crude oil caused no significant reduction in weight gain, but both
the crude oil and aromatic fractions of the other caused significant decreases
in the growth rate. Hallett (in Peakall et al. 1978) found that polynuclear
aromatics with 3 or more rings were present in the oil that caused reduced
growth, but not in the other. This variation in the adverse effects caused
by two presumably similar crude oils suggests that the effects on local breed-
ing populations encountering oil spills may be highly variable and not predict-
able by the degree to which the birds become contaminated. Additional informa-
tion on experimental work showing the effects of oil on marine birds of the
southeast is given in the species accounts for Great Black-backed Gull, Laugh-
ing Gull, and Sandwich Tern.


POTENTIAL HAZARDS TO MARINE BIRDS FROM OFFSHORE OIL PRODUCTION

About two-thirds of the oil in coastal waters comes from runoff and efflu-
ent from terrestrial sources. Tanker operations account for about 26 times
more oil in marine waters of the United States as do offshore operations (Ohlen-
dorf et al. 1978), but may cause a disproportionately large share of avian
mortality. Ohlendorf et al. (1978) suggested that, for the marine environment,
it may be safer to produce oil offshore than to import it. Widespread, severe
effects of chronic small spills at producing offshore oil fields have not yet
been found (Dicks and Hartley 1982) and these fields may cause relatively few
long-term biological problems (Dicks 1982, Dicks and Hartley 1982). The same
may not be true for chronic discharges in coastal waters, particularly at spe-
cific sites where oil is produced, handled, and refined. There the effects may
be local and subtle or severe and long-lasting (Dicks and Hartley 1982).

Longley and Jackson (1980) reviewed the problems caused by the develop-
ment of petroleum resources in brackish marshes and suggested alleviative
measures that could be taken. Effects include direct loss of vegetation and
animals (e.g., by dredging construction of pipelines and roads); addition of
dissolved, particulate and toxic materials to the environment; and changes
in water flows. Changes in water flow were considered the most damaging ha-
zard because they may result in the complete loss of a marsh ecosystem. Such
an event could be accompanied by a reduction or an elimination of the popula-
tions of marine birds that use the habitat for nesting or feeding.










Similar effects are likely when onshore areas and offshore barrier is-
lands are affected by development of oil and gas resources. Changes in water
flow due to dredging could easily change tidal and current patterns, resulting
in the elimination of islands used for nesting. Terrestrial access to larger
islands may result in the introduction of predators (e.g., foxes, raccoons)
that could eliminate an entire bird colony in a season or two. Disturbance
engendered by construction could result in the mass desertion of a traditional
breeding area by some species.

Geological characterizations of the sensitivity of shorelines to oil
spills are not sufficient to determine the overall sensitivity of these areas
(Owens and Robilliard 1981). Owens and Robilliard pointed out that some time-
and site-specific features (e.g., presence of endangered species, migrations
of fish, seabirds, waterfowl, and shorebirds) are largely overlooked in such
considerations and may make a site highly sensitive for only short periods of
time. They stated that "the vulnerability of mobile (whether resident or mi-
gratory) marine bird species is probably more significant than the impact of
oil on the nearby intertidal habitats." They further pointed out that the
threat that the IXTOC I blowout presented to the egg laying of the endangered
Kemp's Ridley sea turtle along an isolated Texas beach would not have been
identified by the usual processes of determining sensitive areas.

Several recent reports reviewed human activities relevant to development
of onshore oil facilities. These reports include Mulvihill et al.'s (1980)
detailed review of the effects of shoreline structures on the coastal environ-
ment, Morton's (1976) review of the ecological effects of dredging, and Buckley
and Buckley's (1976a, 1977a) and Burger's (1981a) reviews of the effects of human
disturbance on colonially nesting birds.

Onshore habitat change or loss resulting from the development of facili-
ties related to offshore oil production in the long run, probably will more
adversely affect waterbirds in the southeastern United States than will the
production of oil itself. The species treated in this volume compose the
group most vulnerable to petroleum development in the southeast. Development
of onshore facilities to produce, transport, and hold oil will probably be a
major source of environmental disruption and disturbance for these birds.
Other forms of coastal development (e.g., recreation and real estate) also may
cause serious declines in populations of the species included here, and may
have more serious consequences than development of petroleum resources. Fur-
ther research into the flexibility in the use of breeding habitat is needed
to determine the extent to which development may damage populations but, as
Buckley and Buckley (1980a) pointed out, such efforts will have little value
unless management efforts include massive habitat protection.

A secondary hazard to breeding seabirds of the southeast is the ingestion
of oil-contaminated food. The impact may not be immediate, but deleterious
effects on the health of adult marine birds and on the production of young can
be severe.










RECOMMENDATIONS FOR FUTURE RESEARCH


This report reveals large gaps in the body of knowledge necessary to
deal effectively with problems relating to marine birds and OCS development.
Below are some of the problems that we believe need most attention.


STATUS AND BIOLOGY OF BREEDING SPECIES

The sizes of breeding populations of marine birds in the southeast are
still poorly known, yet limited data suggest that this area contains signifi-
cant numbers of some species (Table 6). Recent surveys (Soots and Parnell
1975a; Portnoy 1977, 1978 ms; Blacklock et al. 1978, 1978 ms; Parnell and Soots
1979 ms; Portnoy et al. 1981) provide valuable information on the sizes of
populations, but surveys are incomplete for many areas and have been taken
over too short a period to allow for the annual variation usually found in
populations of marine birds. Information from coastal areas of Georgia and
South Carolina is inadequate, and information from Florida is available only
for portions of the state. In addition, the locations and sizes of breeding
colonies of conspicuous breeders (e.g., Sandwich and Royal Terns) that occupy
relatively few nesting sites are better known than those of other species
(e.g., Laughing Gulls, Forster's Terns) whose colonies are more numerous and
widely dispersed, but are found in less easily surveyed habitats. Species
that tend to nest in association with other species (e.g., Gull-billed and
Caspian Terns) are often overlooked on surveys and the actual numbers breed-
ing in the southeast are not well known.

Many recent papers have dealt with the difficulties of censusing water-
birds (Nettleship 1976; Harris and Lloyd 1977; Buckley et al. 1978b; Birkhead
and Nettleship 1980; Drury 1980; Erwin 1980a, 1981; Erwin and Ogden 1980; Hutch-
inson 1980; Portnoy 1980). Perhaps too much emphasis has been placed on obtain-
ing highly accurate censuses when obtaining consistent results over time is
more important. Erwin (1980a) pointed out that "the most important factor in
obtaining systematic sampling results is probably observer consistency" and
that using the same observers over a period of years allows a comparison that
emphasizes relative rather than absolute changes in numbers. In commenting on
censuses conducted in Alaska and New England, Drury (1980) stated that "one
should not draw conclusions based on data gathered in the course of scattered
surveys made over a few years, no matter how precise and thorough the counting
of each sample," a point with which we agree thoroughly. Drury also remarked
that useful conclusions can be drawn from varied surveys if (a) estimates were
made systematically, (b) if the overall change in numbers has been considerable,
and (c) if the biological reasons for periodic changes are known and understood.
He pointed out that variations in numbers between surveys can be dealt with by
calibrating for differences between observers, counting techniques, and counts
taken at different times of day and in different parts of the breeding season.










Table 6. Estimated populations of charadriiform marine birds breeding in
the southeastern United States compared with their status in other areas
administered by the U.S.


Number
breeding World Significance of populations in the
in the breeding southeastern U.S. compared with
Species southeast range other areas under U.S. jurisdiction


Laughing Gull










Herring Gull


Great Black-
backed Gull


Gull-billed
Tern


ca. 263,000


Mostly
North
America


ca. 500 Widespread
in the
Holarctic
regions


ca. 20


Atlantic,
North
America
and eastern
Eurasia


ca. 3,000 Nearly
cosmopolitan


A majority (ca. 65%) of the U.S.
population and probably most of the
world population breeds in the south-
east; most of the remainder breed a-
long the North Atlantic coast. They
also breed in the Caribbean, includ-
the Virgin Islands, but the size of
populations breeding there is poor-
ly known.

Negligible. Well over 100,000 birds
breed along the North Atlantic coast
of North America and in the Great
Lakes region.

Negligible. Most of the North Ameri-
can population breeds along the
North Atlantic coast with the major-
ity breeding in Maine and Massachu-
setts

The southeast contains about 95% of
all Gull-billed Terns breeding in
the U.S. More than half of the
southeastern birds are found in
Texas. Others breed in the Carib-
bean but whether any breed in areas
administered by the U.S. is not known.


Caspian Tern



Royal Tern


ca. 3,000 Widespread The southeast holds over a fifth of
but patchy the U.S. population. Most of the
southeastern birds breed in Texas.


ca. 99,000 Mostly
North
America


Over 90% of the U.S. population
breeds in the southeast.











Table 6. Continued.


Number
breeding World Significance of populations in the
in the breeding southeastern U.S. compared with
Species southeast range other areas under U.S. jurisdiction


ca. 75,000 Mostly
North
America
and
Europe


Roseate Tern ca. 400 (a)


Common Tern


Forster's Tern


Least Tern


Warm and
temperate
waters of
the world


ca. 9,000 Mostly in
eastern
North
America
and the
palearctic
of the
Old World


ca. 24,000


North
America


ca. 26,000 Largely in
the western
and eastern
U.S.


Sandwich Tern


Over 95% of the U.S. population
breeds in the southeast. Some
colonies in Louisiana are among the
largest known. An unknown number
also breeds in the U.S. Virgin
Islands.

Negligible. Most of the Roseate
Terns breeding in areas administer-
ed by the U.S. are in the Atlantic
northeast and in the U.S. Virgin
Islands.

Slight. The only significant popu-
lation breeds in North Carolina and
represents perhaps about 10% of the
birds breeding in the eastern U.S.
Breeding status along the Gulf coast
and in the Caribbean is uncertain.
Most of the U.S. population breeds
around the Great Lakes and in the
Atlantic northeast.

Nearly 70% of the U.S. population
and probably more than half the
world population breeds in the south-
east. About 80% of the southeastern
population breeds in Louisiana.

About 60% of the U.S. population
occurs in the southeast where the
largest known colonies of the
species are found. Considerable
numbers also nest along the U.S.
North Atlantic coast. Others breed
in the Caribbean and in the U.S.
Virgin Islands, but populations
there are poorly known.









Table 6. Concluded.


Number
breeding World Significance of populations in the
in the breeding southeastern U.S. compared with
Species southeast range other areas under U.S. jurisdiction


Bridled Tern None Pantropical Breeds in the U.S. Virgin Islands
except much and elsewhere in the Caribbean but
of Pacific little is known of their populations

Sooty Tern ca. 80,000 Pantropical Florida holds the only sizeable
population breeding in the contig-
uous United States. Thousands breed
in the Caribbean, including the U.S.
Virgin Islands, and in the Mariana
Islands. Hundreds of thousands
breed in Hawaii and American Samoa.

Brown Noddy ca. 3,000 Pantropical The only sizeable population breed-
ing in the contiguous United States
is in Florida. Unknown numbers also
breed in the U.S. Virgin Islands and
many thousands breed in Hawaii,
American Samoa, and the Mariana
Islands.

Black Skimmer ca. 59,000 North Over 85% of the skimmers breeding
and in the U.S. are found in the south-
South east and this area undoubtedly con-
America tains a majority of the North Amer-
ican population. About 70% of the
population in the eastern U.S.
breeds in Louisiana and Texas.

(a) Recent comments by W. B. Robertson, Jr. (pers. comm.) suggest that the
number now breeding in Florida is even less than the figure given in the
species account.









The data currently available, while providing some idea of breeding popula-
tions of southeastern marine birds, do not adequately assess the total number of
birds using the area nor do they provide adequate information about the size
of nonbreeding populations, the seasonal and annual variations in populations,
or the habitats they use. Management decisions based on present information
may err because available data do not reflect conditions applicable when the
decisions need to be made. A continuing commitment to the monitoring of marine
bird populations in the southeast (and elsewhere) is needed. Nothing less will
provide the information necessary to make judicious decisions that will benefit
both the marine bird and human populations that compete for the resources of
the coastal zone.

Data on regional differences in nesting chronology, degree of annual vari-
ation in reproductive success, factors influencing nest-site selection, deter-
minants of colony location (particularly in relation to food resources), and
other demographic parameters, are necessary for satisfactory evaluation of the
effects of managerial decisions on the well-being of populations. Studies
should be undertaken over a period of several years; those conducted during a
single season do not provide enough information for most managerial purposes.
As the National Environment Research Council (NERC) (1977) pointed out, "...
such [long-term studies are] essential as a baseline against which the results
of future studies or environmental impacts can be measured."

The breeding biology of some of the species in this report has been
studied in North America (e.g., Ring-billed Gull), in the Old World (e.g.,
Gull-billed and Black Terns), or in both (e.g., Herring Gull, Common and Rose-
ate Terns), but few of the charadriiforms breeding in the southeast have been
studied there. Only the Sooty Tern and the Laughing Gull have been relatively
well studied in the southeast, and much remains to be learned of the biology
and numbers of the latter. Some information is available on the breeding biol-
ogy of Least Terns and Black Skimmers in the southeast. Royal, Sandwich, For-
ster's, and Gull-billed Terns are largely unstudied in the southeast and the
latter three have been little studied anywhere in the United States. Virtual-
ly nothing is known about the breeding biology of Caspian Terns in the south-
east or of the status and reproductive biology of the Common and Roseate Terns.
The latter is declining in many portions of its range and has been recommended
for endangered species status (Buckley and Buckley 1981).

One way of predicting the effects of oil spills on local populations is
to develop mathematical models that take into account various biological para-
meters that determine the response of populations to spills. A recent attempt
to do this by Ford et al. (1982) indicated that the following parameters are
those most needed for such models:

1) the size of the nonbreeding population,
2) the movement patterns of foraging individuals,
3) the spatial and temporal distribution and availability of food near breed-
ing colonies,
4) the relationship between feeding rates and the age of young in relation
to growth rates and survival probabilities,









5) the degree of density dependence in various population parameters,
6) the probability that a given bird will die as the direct result of an oil
spill,
7) the age-specific mortality schedules of local populations under normal
conditions,
8) the rate at which populations respond to perturbations and regain an
equilibrium distribution at sea, and
9) the effect of an oil spill on the short-term availability of food.

Ford et al. (1982) thought that the last four parameters are the most
important for developing the model, but none are easy to measure and some
require massive logistical support. Future research on species breeding in
the southeast should be directed towards answering some of these questions.


DISTRIBUTION AND BIOLOGY OF TRANSIENT, WINTERING, AND PELAGIC POPULATIONS

Our knowledge of the distribution, numbers, and biology of charadriiform
marine birds occurring offshore or during nonbreeding periods is even more
limited than our knowledge of the status of breeding species. The Bridled
Tern is one of the commonest of offshore species, but little is known of its
areas of origin or biology. Large portions of the North American populations
of Black Terns and Franklin's Gulls migrate through the study area. The area
is also a major wintering ground for Laughing, Bonaparte's, Ring-billed, and
Herring Gulls; Caspian, Royal, and Forster's Terns; and Black Skimmers. We
know little about habitat use by these wintering populations, nor do we know
much about total populations.

With the exception of the Sooty Tern, very little banding has been done
in the enormous marine bird colonies of the southeast; in consequence scien-
tists usually have no idea of the routes by which they disperse from these
breeding colonies. A mass banding program, judiciously undertaken, should
add greatly to our knowledge of their dispersal. Such work should be conduct-
ed over a period of years and be integrated with a marking program that would
allow the recognition of individuals. Such a program could also provide use-
ful data on timing of migration through the area, as well as provide insight
into foraging ranges, local movements, and various aspects of breeding biol-
ogy. Among the charadriiform species nesting in the southeast, Laughing
Gulls; Caspian, Royal, Sandwich, and Forster's Terns; and Black Skimmers would
probably be the species most easily treated by this approach.

Present knowledge of pelagic birds in the southeast is poor. Only a few
pelagic surveys have been conducted in this area with any thoroughness and
regularity. Studies by David Lee in North Carolina, Charles Duncan and Ralph
Havard in Alabama, and by John Johnson in Florida, have been only partially
reported and tend to concentrate on rare birds and those seen well offshore.
Duncan and Havard's recent (1980) study revealed that species like the Blue-
faced Booby (Sula dactylatra) and Bridled Tern, formerly thought rare in the
northern Gulf, are in fact major components of the pelagic avifauna.










Comprehensive and detailed offshore surveys of marine birds, conducted
by boat at least monthly and over a carefully chosen grid, would help immense-
ly in planning contingencies to be taken following oil spills. Such surveys
should last at least 2 years because data from any given year may not repre-
sent typical conditions. Recent aerial studies of this nature (e.g., Fritts
et al. 1982 ms) help delineate the extent to which offshore areas are used
by marine birds and at least should result in useful information about the
amount of biomass in a given area at a given time. One limitation of this
approach is the low reliability of identification to species. Fritts et al.
(1982 ms) were not able to distinguish from the air between four species of
Sterna (Forster's, Common, Arctic, and Roseate Terns), nor could they usually
distinguish between species of storm-petrels, tropicbirds, jaegers, or phala-
ropes. Their report mentions neither Caspian nor Gull-billed Terns although
both must have been present in inshore areas.

An inexpensive way to determine the potential effect of oil development
would be to make observations from oil rigs on species composition and behav-
ior of passing transients. Some species, such as the boobies, are probably
attracted to these platforms, because the rigs may concentrate local food
resources.

International borders are biologically imaginary lines that tend to dis-
tort our knowledge of the distribution of birds. "Foreign" birds may form
a large proportion of the biomass of marine birds in southeastern waters for
part of the year. One of the largest oil spills in history followed the blow-
out on 3 June 1979 of the IXTOC I well of Petroleos Mexicanos in the southern
Gulf of Mexico. Eventually over 3 million barrels of petroleum were lost and
an oil slick that spread across the southern gulf by August washed ashore along
80 km (50 mi) of the Texas coast (Waldichuk 1980). Although only about 20
birds were found dead from oiling in this area (Garmon 1980), several of these
were pelagic species seldom seen along the Texas coast but common in the waters
of the U.S. Gulf of Mexico (Clapp et al. 1982b). These dead birds included
five Blue-faced Boobies and one Brown Noddy (B. Chapman, pers. comm.), neither
of which is common off the Texas coast. The low toll of marine seabirds, one
of the groups known to be most severely affected by oiling, is not surprising
since the Texas coast is some 800 km (ca. 500 mi) north of the blowout site in
the Bay of Campeche (at ca. 19020'N, 92025'W). What is surprising, however,
is that none of the follow-up reports addressed the problem of what happened
to seabird colonies nearer the blowout. Three areas near the IXTOC blowout
(Cayos Arcas [20013'N,91058'W], Cayos Arenas [22007'N, 91024'W] and Arrecifes
Alacran [Alacran Reef] [22023'N, 89040'W]), presently or formerly supported
populations of breeding or roosting seabirds. The closest of these, Cayos
Arcas, is only some 45 mi (75 km) from the wellhead of the IXTOC I spill. It
held some 5,000 Blue-faced Boobies in late 1952. Another 400 were seen on
Cayo Arenas. This species, the Magnificent Frigatebird (Fregata magnificens)
and the Royal Tern are all known to breed on Cayo Arcas (Paynter 1955). Ala-
cran Reef supports breeding populations of Blue-faced Boobies, Brown Boobies
(Sula leucogaster), and Magnificent Frigatebirds. Alacran Reef may also sup-
port nesting populations of Royal and Sandwich Terns (Boswall 1978). Boswall
(1978) reported about 2,000 Blue-faced Boobies and 2,000 Brown Noddies in










September 1975. He noted that thousands of Sooty Terns apparently nest on the
reef. In the early 1950's some 800 Brown Boobies were present and as many as
5,000 Magnificent Frigatebirds bred there (Paynter 1955). The numbers suggest
this area is an important source for the pelagic avifauna of the Gulf of
Mexico.

Several of the marine birds treated here and in Clapp et al. (1982b)
breed in the Caribbean. Audubon's Shearwater (Puffinus Ihermineri) and the
Bridled Tern, periodically common to abundant off our southeastern coasts,
breed in the U.S. Virgin Islands and elsewhere in the Caribbean. The Carib-
bean may be an important source for these as well as for other species (e.g.,
Blue-faced Booby, Magnificent Frigatebird, Sooty Tern) common in the offshore
waters of the southeast. Species of marine birds that breed or have bred in
the U.S. Virgin Islands include Blue-faced Boobies; Brown Boobies; Red-footed
Boobies (Sula sula); Magnificent Frigatebirds; White-tailed Tropiebirds (Phae-
thon lepturus); Red-billed Tropicbirds (Phaethon aethereus); Sandwich, Sooty,
and Bridled Terns; and Laughing Gulls (Dewey and Nellis 1980). The Roseate
Tern population is one of the largest remaining in the Caribbean. The U.S.
Virgin Islands are one of the few (perhaps the only) areas under U.S. juris-
diction where breeding Audubon's Shearwaters, Red-billed Tropicbirds, and
Bridled Terns may be found. Colonies of marine birds in the Virgin Islands
are much in danger from introduced and feral animals and from poachers (Dewey
and Nellis 1980). What information is available makes clear that the unoccu-
pied islands of the Caribbean are one of the main refugia for marine birds in
the warmer waters of the world.

We recommend that efforts be made to initiate cooperative international
surveys of marine bird populations in the southern Gulf of Mexico and in the
Caribbean. International surveys would supply not only a much better under-
standing of the overall status of the species involved, but also would permit
far better insight into the consequences of local managerial decisions on a
species throughout its range. Previous efforts along these lines, particularly
with regard to waterfowl, have been highly effective in producing the informa-
tion needed to manage populations.


EFFECTS OF OIL ON SOUTHEASTERN MARINE BIRDS

We believe that attempted rehabilitation of oiled birds following a major
oil spill is largely a waste of time, money, and other resources. A group of
marine bird experts (NERC 1977) stated that "since the results of attempts to
rehabilitate oiled birds are so poor, it may be more profitable to expend ef-
forts at preventing birds from becoming polluted." It is desirable, however,
to salvage oiled birds to find out what species were affected and to obtain
information that will allow for more prudent responses to future spills.
Satisfactory efforts to adequately document losses to oil spills in the United
States are almost nil. This information is often difficult to obtain because
counts of dead or contaminated birds may not indicate the numbers that were
affected or will die. Seldom is there information on the number of birds pre-
sent in an area before, during, and after contamination. While it may be next










to impossible to do more than estimate numbers present before a spill, rapid
responses to oiling incidents should allow some idea of the numbers present
during and after a spill. Detailed observations made during and after the
spill may allow better estimates of the severity of the spill.

Other factors make estimation of the damage caused by spills more diffi-
cult. For example, the time of an oil slick's passage through an area may be
crucial in determining the degree of contamination and mortality experienced
by each species. During the contamination of the Firth of Forth in February
1978, oil passed near the main feeding area for waterbirds at night; conse-
quently, there was a proportionately greater loss of night-feeding Greater
Scaup (Aythya marila) and Pochards (Aythya ferina) than there was of Common
Goldeneye (Bucephala clangula) and Common Eiders (Somateria mollissima), most
of which had moved elsewhere to roost (Campbell et al. 1978).

The proportion of birds oiled following a pollution incident may vary
widely between species, depending on the habitats used and the habits of the
birds. The probability of finding most oiled birds that roost or loaf onshore
near their offshore feeding areas is greater than it is for finding birds that
spend all or most of their time offshore and that, following oiling, sink from
sight never to be seen again.

Further, wind, offshore currents, and movements by the birds themselves
may take most of the victims of an oil spill far from where they were oiled
before anyone notices their plight. In some parts of Europe and on the west
coast of the United States, prevailing winds bring victims of oiling to shore.
In contrast, on the Atlantic seaboard winds take oiled birds out to sea. Con-
sequently, comparisons of damage from oil pollution incidents between these
areas are not valid. Likewise, we cannot use estimates of mortality from
beached bird surveys in Europe to predict the incidence of mortality in the
western Atlantic. At best, the European reports suggest that damage to wild
birds from oil on the U.S. east coast may be greatly underestimated.

Powers and Rumage (1978) documented one spill off Massachusetts well
enough to demonstrate differences between onshore and offshore observations.
Following the ARGO MERCHANT spill, prevailing winds and tides made it unlike-
ly that oiled birds would wash ashore and led Powers and Rumage to suspect
that the few birds (181) found oiled on the beaches of Nantucket Island and
Martha's Vineyard had made an active effort to get ashore. Hundreds of oiled
Great Black-backed and Herring Gulls were seen offshore during at-sea surveys
following this spill (Powers, pers. comm.). Oiled Northern Fulmars, a species
not found along the beaches, were also seen. Most oiled birds were seen near
the tanker or oil slick, but oiled gulls and Black-legged Kittiwakes were
widely dispersed due to their penchant for following fishing trawlers. Levy
(1980) analyzed the oil found on dead or moribund birds in Atlantic Canada
and suggested that Herring and Great Black-backed Gulls obtained near Sable
Island, Nova Scotia, had been contaminated by oil from the ARGO MERCHANT
spill that had occurred some 840 km (520 mi) away.










Despite the difficulties in obtaining unbiased data on the effects of
oil spills, we still recommend that better efforts be made to monitor and
publish reports of the effects of these spills on marine birds. Much of the
information needed to answer questions relating to oil pollution and marine
birds in the southeastern United States would be available if such efforts
had been made in the past.

We also recommend that more attention be paid to monitoring long-term
effects of oil pollution in the southeast. One of the better and less expen-
sive ways in which this may be accomplished is a periodic census of birds
found dead along the beaches. This lends some objective basis to speculations
about the effects of oil pollution, and also provides information about unusual
or increasing mortality from other causes (e.g., pesticides). Over time, this
may serve as an early warning indicator of serious problems. Such surveys are
conducted in the eastern United States by the Atlantic and Gulf Coast Beached
Bird Survey Project, but the area covered in some regions is very small (e.g.,
2 mi of the Texas coast [M. Simons, pers. comm.]).


ACKNOWLEDGMENTS


Many people contributed to this report in a variety of ways. Gene W.
Blacklock allowed us to use his unpublished manuscript on the occurrence and
status of the birds of Padre and Mustang Islands and supplied us with unpub-
lished material on the Texas Colonial Waterbird Census. William B. Robertson,
Jr. and Herbert W. Kale, II, provided us with unpublished summary material on
Florida birds. Malcolm M. Simons, Jr., Director of the Atlantic and Gulf
Coast Beached Bird Survey Project, supplied unpublished data on incidence
of oiling and mortality of birds along the Atlantic and Gulf coasts.

The library staff of the Smithsonian Natural History Museum was especial-
ly helpful in obtaining copies of most of the theses and many of the papers
that make up the files upon which this and the companion reports are based;
Carolyn S. Hahn, Amy E. Levin, and Jack F. Marquardt were extremely helpful
in this regard. Agnes C. Nalley also aided us in finding and obtaining lit-
erature in the Bird Library of the Gabrielson Laboratory of Patuxent Wildlife
Research Center.

Linda A. Hollenberg helped assemble species bibliographies and the liter-
ature files. Initial versions of the maps were prepared by Martha B. Hays and
completed by the staff of the National Coastal Ecosystems Team, Slidell, Louis-
iana.

We thank Helen L. Harbett, Tracy Chevalier and Linda M. Wolfe who typed
some of the preliminary and final draft. Linda M. Wolfe was also very help-
in proofing final draft. We also thank Charlotte I. Adamson who prepared the
illustrations that intersperse the text and William C. White whose technical
assistance in keeping our faltering word processors functional went far beyond
the requirements of his job and is much appreciated. Jeffrey A. Spendelow,










Jill Parer and Cherry eler provided some much needed editorial assistance
and we thank them for their efforts.

We also thank Francine G. and Paul A. Buckley, R. Michael Erwin, Malcolm
C. Coulter, Martha B. Hys, W. B. Robertson, Jr., alph W. Schrelber, and
Richard L. Zusi who comented on various species accounts and often pointed
out information that we had missed. These individuals often were the source
of much valuable unpublished data.

Early versions of several species accounts were prepared by Wayne A. Hff-
man, who also supplied information on unpublished sightings of marine birds
in the Gulf of Mexico. Charles D. Dncan and Tomas H. Fritts provided un-
published information for this area.

Others also supplied unpublished manuscripts or copies of papers, jour-
nal or reports that would have otherwise been very difficult to obtain, and
provided guidance as to areas where additional information could be acquired.
For these and other services, we thank David G. Ainley, Hans Blokpoel, Eirik A.
T. Blom, W. R. P. Bourn, Danny Bystrak, Brian R. Chapman, Frank Cobb, Jr.,
Mercedes S. Foster, Patrick J. Could, Craig S. Harrson, Jerome A. Jackson,
M. Kathleen Klimkiewicz, Roxie C. Laybourne, Mary K. LeCroy, David S. Lee,
Martin K. MNicholl, lan C. T. Nisbet, Storr L. Olson, Sam Patten, Jr.,
John W. Portnoy, Kevin D. Powers, Chandler S. bobbins, Larry R. Shanks, Fred C.
Sibley, David M. Smzth, Jeffrey A. Spendelow, Judith A. Toups, Jacob Valentine,
George E. Watson, III; John S. Weske, Claudia P. Wilds, and Donald W. Woodard.

This proec was sponsored by the Bureau of Land Manaement's Outer Contin-


An oiled adult Laughing Gull at Flamingo, Florida. Photograph by Roger B. Clapp.
38










RED-NECKED PHALARUPE


(Phalaropus lobatus)


[DA: Udinshane, UU: Grauwe franjepoot, EN: Red-necked Phalarope, FI: Isovesi-
paasKy, FR: Phalarope a bec etroit, Phalarope hyperbore, GE: Odinshuhnchen,
Odinswassertreter, Schmalschnabliger Wassertreter, IC: Odinshani, IT: Falaropo
beccosottile, JA: Aka-eri hire-ashi, NW: Svommesnipe, PO: Platkonog rdzawoszyi,
PR: Falarodo, RU: (Round-nosed Phalarope), SP: Falaropo picofino, Chorlillo
norteno, SW: Smalnabbad simsnappa, US: Northern Phalarope]


GENERAL DISTRIBUTION

North America Ked-necked Phalaropes breed in low Arctic and Subarctic
North America from the Pribilof and Aleutian islands east and north through
northern Alaska, thence east along the northern part of the continent through
southern Victoria Island and central Keewatin to southern Baffin Island. They
breed south to southern Alaska, southwestern Yukon, and northwestern British
Columbia, and from northwestern Mackenzie southeast to northern Saskatchewan,
Manitoba, and Ontario. They also breed east on islands and the coast of James
Bay through northern Quebec to the east coast of Labrador (AUU 1957, Godfrey
1966).

Ked-necked Phalaropes migrate in considerable numbers along and off the
coasts of North America (AUU 1957, Godfrey 1966) and are widespread migrants
in tropical Atlantic waters (Watson 1966). They also migrate through the in-
terior of western North America (AOU 1957) and regularly, but in much smaller
numbers, through the interior farther east. Transients have been recorded in
Central America in Guatemala, Honduras, Costa Rica, and Panama (Blake 1977).

World Distribution Red-necked Phalaropes breed in the Old World in south-
ern Greenland, Iceland (Hohn 1965), the Faeroes, Scotland and Ireland (rarely)
(BOU 1971), in Norway and Sweden south to about 620 N, and in Spitsbergen and
Finland, locally south in the latter to about 640 N. This species also breeds
in the U.S.S.R. in northern Estonia and in northern Russia on the northern Kola
Peninsula and Kandalaksha Bay east to the Taimyr Peninsula (to about 720 N)
and east along the coast to the Chukotski Peninsula. From northern Siberia
it breeds as far south as about 64-660 N, and east to the Commander Islands
(Vaurie 1965). European populations from southeastern Scandinavia and northern
Russia migrate southeast to the Persian Gulf; birds from Old World populations
farther west migrate south along the west coast of Europe to winter off Africa
(Hohn 1966, Hilden and Vuolanto 1972).


Taxonomic note: This species has often been placed in the genus Lobipes.
"Northern Phalarope" is the common name used in most of the recent North Amer-
can literature, but we prefer to use the name "Red-necked Phalarope", following
the usage adopted by the forthcoming AOU checklist.





Ked-necked Phalarope


Both Old and New World populations winter at sea, largely in tropical and
subtropical waters. In the New World, the species winters in large numbers off
the Peruvian coast (Murphy 1936), where it occurs chiefly on the outer fringes
of the Humboldt Current (Blake 1977). Blake (1977) considered it casual in the
coastal waters of Chile south to Santiago, but others (AUU 1957, Vaurie 1965)
thought the wintering range off Pacific South America was from the Galapagos
and Ecuador south to Chile and southern Argentina. These phalaropes also win-
ter commonly off the south Atlantic coast of South America (Hohn 1965, 1969).

Red-necked Phalaropes winter in the Old World off west Africa, Arabia, and
in the waters of southeast Asia (BOU 1971); in the western Pacific they occur
in large numbers off Japan (OSJ 1974), and winter from the Ryukyus south and
east to northern New Guinea (Vaurie 1965), where they are not uncommon (Condon
1975).


DISTRIBUTION AND ABUNDANCE IN THE COASTAL SOUTHEASTERN UNITED STATES

North Carolina Pearson et al. (1942) considered Red-necked Phalaropes
rare transients along the coast, but recent observations show they are actual-
ly numerous offshore during migration. This species has been recorded more
than 70 times in North Carolina (Lee and Booth 1979) with most sightings from


10 August through 25 October and from 3 May through 10 June.
proximately 50 published records of the Red-necked Phalarope
lina, but we list below only those- that involve four or more


1909 23 Sep.


1962 13 Sep.

1966 4 Sep.

1972 27 May

1972 8 Oct.

1973 26 May

1973 2 Sep.

1973 16 Sep.

1974 1 Sep.

1975 12 Oct.

1976 18 May


5 seen (3 coll.) at White Lake, Bladen
County

30 seen 22-25 mi E Hatteras Inlet


5 seen off Hatteras Inlet

30 seen at Pea Island

45 seen off Hatteras

30 seen off Morehead City

4 seen off Hatteras

15 seen off Hatteras

6 seen off Hatteras

70 seen off Cape Hatteras


4 seen
Inlet


feeding at sandflats, Oregon


We know of ap-
from North Caro-
birds.

Pearson et al.
1942


Grant 1963


Parnell 1967a

Teulings 1972c

Teulings 1973a

Teulings 1973e

Teulings 1974d

Teulings 1974d

Teulings 1975a

Teulings 1976a

Teulings 1976d






Red-necked Phalarope


1976 11 Aug. 4 seen off Oregon Inlet Teulings 1977a

1976 5 Sep. 26 seen off Oregon Inlet Teulings 1977a

1976 6 Sep. 27 seen off Hatteras Teulings 1977a

1976 21 Sep. 20 seen off Oregon Inlet Teulings 1977a

1977 9 June 3-6 seen off Oregon Inlet LeGrand 1977b

1978 10 Oct. 13 seen off Oregon Inlet LeGrand 1979a

1979 10 May 91 seen off Oregon Inlet LeGrand 1979d

South Carolina The status of the Red-necked Phalarope in South Carolina
is probably the same as in North Carolina. Although regarded by Sprunt and
Chamberlain (1949) as a rare transient, mostly along the coast, they are almost
certainly common migrants offshore. Wayne (1910) and Sprunt and Chamberlain
(1949) agreed that the species was abundant offshore despite the paucity of
records. We found only 11 records for the state, the majority (7) from the
spring migration. Spring records extend from 17 May through 3 June, and fall
records from 11 September through 25 October.


1880

1885


17 May

25 Sep.


1885? 25 Oct.


1903 3 June

1913 28 May


1933 30 May

1934 29 May

1950 11 Sep.

1956 2 May

1961 22 May

1972 27 Hay


1 coll. in Chester Co., 150 mi inland

seen at Frogmore, Beaufort Co.


seen at "Sea Islands"


1 captured at Charleston

I seen off Mt. Pleasant


1 female seen at Cape Romain

1 ad. female coll. at Bull's Island

1 seen inland in Columbia area

2 seen inland at Aiken

1 female seen inland at Lake Murray

1 seen at Huntington Beach


Loomis 1880

Sprunt and Cham-
berlain 1949

Sprunt and Cham-
berlain 1949

Wayne 1905

Sprunt and Cham-
berlain 1949

Sprunt 1933

Chamberlain 1934

Burton 1970

Burton 1970

Smith 1961

Teulings 1972c


Georgia Red-necked Phalaropes are thought to be rare inshore and offshore
migrants in Georgia and accidental inland (Denton et al. 1977). We know of only






Red-necked Phalarope


about 18 records for the state. Records from the spring migration extend from
5 May through early June; records from the fall migration from 24 July through
3 October. There is one winter record.


1933 24 May

1937 3 Oct.

1950 9 May


s


1958 13 Aug.-
13 Sep.

1959 19 Sep.

1959 23, 24 Sep.

1964 10 Sep.

1968 12 Sep.

1972 27 May

1976 23 May

1976 23 May

1976 early June

1977 24-26 July

1979 11 Feb.


female coll. at mouth of Savannah River Burleigh 1958


1 male coll. in Grady County

1 male coll. inland near Augusta


1 seen at Savannah

2 seen on pond 3 mi E Savannah

1 coll. at Hutchinson's Island, Savannah

een at Savannah


4-11 seen at Hutchinson's Island


2

4,1

1

2+

1

2

2(?)

2

2

3


seen at Hutchinson's Island

seen at Hutchinson's Island

seen at Tybee Island

seen at Jekyll Island

seen at St. Simons Island

seen (1) inland near Gainesville

seen (1) inland at Pendergrass

seen (1) near Pendergrass

seen inland at Eufaula NWR

seen off Jekyll Island


Burleigh 1958

Denton and Cham-
berlain 1950b

Tomkins 1958

Chamberlain 1952a

Tomkins 1958

Denton et al.
1977

Chamberlain 1959a,
Tomkins 1958

Chamberlain 1960a

Chamberlain 1960a

Parnell 1965a

Masters 1968

Teulings 1972c

Teulings 1976c

Denton et al. 1977

LeGrand 1976

LeGrand 1977b

LeGrand 1979b


Florida Atlantic Coast The Red-necked Phalarope is a rare migrant along
the coast and is occasionally found inland (Kale 1979 ms a). There are few
early reports of this species from the Atlantic coast of Florida; Howell (1932)

(1) These three 1976 records may be of the same birds.





Ked-necked Phalarope


listed one record, and another five were summarized by Sprunt (1954). Since
then there have been at least another 22 records from the coast, as well as
seven from inland localities. We list below only coastal records since
Sprunt's (1954) publication, but include all coastal observations in Table 7.
Numbers of Red-necked Phalaropes seen in recent years clearly indicate that
the species is much more abundant offshore than previous records suggested, and
is most abundant as a transient in April-May, and in September-October (Table
7). Numbers seen in December and January also suggest that some may winter
in offshore waters.

1958 8-10 May 1 seen at Canaveral Stevenson 1958c

1962 24 Apr. 1 seen at Fort Pierce Paulson and


1962 6, 8 May at least seen (some coll.) in small flocks E
32 of Palm Beach

1962 16 Dec. 16 counted 40 yds offshore between
Juno Beach and Jupiter

1963 24 Sep. 2 seen at Playalinda Beach

1965 10-11 Sep. 1 seen at Guano Lake

1967 21 Oct. 1 seen at Sawpit Sanctuary, Duval County


seen at Port Canaveral

seen in Indian Kiver at Rockledge

seen at Little Talbot Island

off Cocoa

seen off Boynton Inlet

seen in Indian River near Sebastion

seen 18 mi E Cape Canaveral

seen 24 mi off Mayport

seen daily at Port Canaveral

seen 20 mi off Cocoa

found dead near Stuart


Stevenson 1962

Paulson and
Stevenson 1962

Stevenson 1962b


Cunningham 1964a

Stevenson 1966a

Kobertson and
Ogden 1968

Robertson 1970

Stevenson 1972a

Ogden 1972

Stevenson 1973

Stevenson 1973

Woolfenden 1973

Kale 1973

Edscorn 1974

Edscorn 1974

Ogden 1974

Edscorn 1975


1969

1972

1972

1972

1972

1972

1973

1973

1973

1974

1974


9 Oct.

21 Mar.

28 May

26 Aug

28 Oct.

16 Dec.

4 May

9 Sep.

13-29 Oct.

29 June

25 Sep.


22

12

6

seen

4

1

65+

8

1-2

2





Red-necked Phalarope


1975 7 Sep. 19 seen off iayport

1977 23 Jan. 36 seen 10-15 mi off Canaveral

1978 29 Apr. up to 30 seen off Ponce Inlet

1978 30 Dec. 1 seen at Port Canaveral

Florida Keys We know of only five records from the Keys
of either fall migrants or wintering birds.

1954 15 Aug. 1 seen at Key West

1954 14-15 Nov. 1 seen at Little Duck Key

1954 23 Nov. 1 seen at Key Vaca

1967 15 Oct. 1 found dead at Norris Cut,
Virginia Key

1972 20 May 1 male seen off Duck Key, Florida
Straits


Edscorn 1976

Stevenson 1977

Kale 1978a

Stevenson 1979a

; all but one are


Stimson 1955

Stevenson 1955a

Stevenson 1955a

Robertson and
Ogden 1968

Kale 1972


Florida Gulf Coast Red-necked Phalaropes occur over the open waters of
the Gulf of Mexico but are seldom recorded from the Florida Gulf coast (Kale
1979 ms b). We found only 16 records for this area, predominantly of spring
migrants (14 March-25 May). More than half of all Red-necked Phalaropes report-
ed along the Florida Gulf coast have been seen between 13 and 25 May. Two sum-
mer records could have been either of early fall migrants or of birds that had
lingered in the south during the breeding season. Few Red-necked Phalaropes
have been recorded along this coast in fall and winter.

1918 14 Mar. 1 seen 175 mi W of Tampa Howell 1932

1929 14 May 1 coll. near Plant City Howell 1932

1954 29 Apr. 1 seen, coll. at Snake Bight (Cape Stevenson 1954a
Sable area)

1956 25 May 1 coll. at Lake Jackson Stevenson 1956c

1959 22 Nov. 2 seen at Cape San Blas Newman 1960a

1963 14 May 4 seen ca. 1 mi off Alligator Point Stevenson 1963b

1963 6 July 1 coll. near Tampa Stevenson 1963c

1965 27 June 1 seen at Sanibel Island Ogden and Stev-
enson 1965





Red-necked Phalarope


1969 19-20 May I seen on upper St. Marks River, Pantelidis and
Wakulla County Stevenson 1969

1970 12, 16 May 1,2 seen on freshwater pond, Tampa Edscorn 1971

1972 11-15 May 1 seen at McKay Bay Kale 1972

1972 13 May 1 seen at Navarre Beach Imhof 1972

1973 12 Oct. 1 seen at Tampa Edscorn 1974

1975 27 Nov. 1 found at Navarre Purrington 1976

1976 21 Jan. 1 seen at Sanibel Island Stevenson 1976

1977 25 May 1 ad. female seen at Toytown Dump Kale 1977

Alabama Red-necked Phalaropes occur only occasionally in Alabama, predom-
inantly during the fall migration (Imhof 1976b). We know of only 12 records
for the state; eight are from inland localities, the rest from the coast.
Spring records are from 29 April to 17 May. Fall records occur from 6 August
to 13 November, and there is one winter record. Imhof (1976b) listed nine
of the records; we list below those from the coast and those published subse-
quent to Imhof (1976b).

1964 27 Aug. seen at Dauphin Island Imhof 1976b

1973 10 Sep. 6 seen 9 mi S of Dauphin Island Imhof 1976b

1973 13-14 Oct. seen at Blakely Island, Mobile Imhof 1976b

1975 6 Aug. 1 seen at Wheeler NWR Purrington 1976

1977 29 Apr. 1 seen at Blakely Island, Mobile Imhof 1977

1977 27-29 Aug. 1(?) seen in Limestone Co. Purrington 1978

Mississippi Red-necked Phalaropes are rarely seen in Mississippi. There
are only three records, all of fall migrants. The first two phalaropes were
recorded inland at sewage lagoons in 1976. One was seen 11 September at Kos-
ciusko, Attala County (Sanders 1976) and another was seen 21 September at
Hattiesburg (Gates 1976). The third sighting was made along the coast at
Pascagoula on 10 September 1977 (Purrington 1978).

Louisiana Red-necked Phalaropes are rarely seen in Louisiana. We know
of only six records for the state, five of them listed by Lowery (1974). As
on the Texas coast, this species is evidently more frequent during spring
migration than in the fall.





Red-necked Phalarope


1966 8 May 1 ad. female coll. 10 mi W of Johnson's
Bayou, near Sabine Pass


1970 1 May


6 seen at Cameron


1971 30 Apr.- 4,3 seen at Cameron
1 May


1971 12 Sep.

1978 14 May


1 seen 10 mi W Johnson's Bayou Lowery 1974

1 seen at Holly Beach Oil Field, Cameron Imhof 1978


Texas Oberholser (1974) reported that Red-necked Phalaropes are regular
but scarce in the interior during fall migration; they are rare during spring
migration. Along the coast they are rare to casual in fall and (along the cen-
tral coast) casual in spring. Spring migration occurs mainly from late April
to late May, and most fall migration occurs from mid-August to mid-October. Ex-
treme dates of occurrence range from 4 April to 6 June, and from 12 July to 22
December (Oberholser 1974).

We know of approximately 55 records of Red-necked Phalaropes from Texas.
About three-quarters of these records are from inland localities; nearly two-
fifths of them were made at Austin, where the species is regularly recorded
during the fall migration. September is the month of peak occurrence in the
interior but along the coast most Red-necked Phalaropes have been seen from
the latter half of April through early May. We list below 13 records obtained
on or near the coast.


1938 12 July 1 seen at Jones Lake, Aransas NWR


1940 4 Apr. 2 seen at Aransas NWR


1940 25-29 Apr. 7 seen on flats W of Mullet Bay,
Aransas NWR


Oberholser
1963 ms

Oberholser
1963 ms

Oberholser
1963 ms


1952 26 Apr. 1 seen at Rockport


1959 4 Oct.


1959 15 Sep.

1961 18 July


1 seen at Baytown Tunnel


1 seen at Laguna Atascosa

2 seen at Aransas NWR


Watson and Gold-
man 1952, Ober-
holser 1963 ms

Webster 1960a,
Oberholser 1963 ms

Webster 1960a

Webster 1962a,
Oberholser 1963 ms


Lowery 1974


Lowery 1974

Lowery 1974





Ked-necked Phalarope


1962 17 Apr. ca. 35 seen eating insects on seaweed, Mustang Webster 1962b
Island beach

1963 9-10 May 2 seen on Uso Creek, W of Corpus Christi Webster 1963

1968 18 Oct. 1 seen 10 mi E of Brownsville Webster 1969a

1973 9-10 May 2 seen at Corpus Christi BlacklocK 1978 ms

1975 31 Oct. 1 photogr. at Padre Island Natl. Seashore BlacKlock 1978 ms

1979 29 Apr. 1 photogr. in Brazoria County Webster 1979c


SYNOPSIS OF PRESENT DISTRIBUTION AND ABUNDANCE

Breeding Red-necked Phalaropes breed circumpolarly in the northern Hol-
arctic, largely between 520 and 74 N. They are found mostly in the tundra,
but may also breed in the boreal zone.

Winter Red-necked Phalaropes winter at sea, primarily in tropical and
subtropical oceans. They may be found off the coasts of South America, north-
west Africa, Arabia, and southeast Asia.

Migration The Red-necKed Phalarope occurs in the southeastern United
States primarily as a transient in spring and fall (Tables 7, 8). Most occur
offshore, and their status off many states is largely unknown because these
waters have been poorly investigated. In North Carolina and Florida where off-
shore surveys have been conducted most frequently, these phalaropes are common
migrants. Although considerably less information is available for the northern
Gulf of Mexico, it seems evident that Red-necked Phalaropes are less abundant
there than off the Atlantic coast and are apparently relatively more common
during spring migration. Peak periods of occurrence in both the Gulf and At-
lantic areas are April-May and September-October (Table 7).

On the basis of ringing recoveries obtained in the Old World, Hohn (1966)
thought that the birds of Sweden, Finland, the Baltic area, and adjacent north-
ern Russia migrate overland toward the Caspian Sea and probably from there to
the Persian Gulf. Hohn also suggested that migrants off western Europe probab-
ly come from the east coast of Greenland, Iceland, the Faeroes, the northern
British Isles, and Norway. hilden and Voulanto (1972), after a more recent
examination of Finnish ringing recoveries, agreed with Hohn; they believed
that the migration from Finland takes place to the southeast over the European
continent. Additional information on migration in other areas is given by
Meinertzhagen (1925), Bent (1927), Hohn (1965), Schiemann (1972), and Glutz
von Blotzheim et al. (1977).





Red-necked Phalarope



Table 7. Approximate number of Red-necked Phalaropes recorded by month for
the coastal southeastern United States (a).


State/region


North Carolina
South Carolina
Georgia
Florida-Atlantic Coast

Subtotal-ATLANTIC COAST

Florida-Keys

Florida-Gulf Coast
Alabama
Mississippi
Louisiana
Texas-Coast

Subtotal-GULF COAST

TOTAL-ALL AREAS


JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC


1 171 7 4 7 144 130
S 8 1 2 1 -
3 6 2 2 5 16 1 -
36 12 31 104 2 2 83 44 2 32

36 4 12 31 289 12 6 14 245 176 2 32

- 1 1 1 2 -


1 1 1 12
- 1 1


1 1 1 3 -
- 3 6 4 2 1


3
- 4 11 1
- 46 4 4 1


3 -


1 1 52 28 1 5 3 11 8 5 1

37 4 13 83 318 13 11 18 256 185 9 33


(a) If the source did not make it clear whether one or more birds were seen,
we assumed that only one was seen. When birds were seen in more than
one month we counted them separately in each month.


HABITAT

Nesting Bent (1927) described the preferred nesting habitat of Red-necked
Phalaropes in the western Aleutian Islands as the wetter portions of flat allu-
vial plains. Nests were placed in small hollows in mounds or tussocks in wet
meadows near the edges of marshy ponds or near the mouths of small streams.
At Scammon Bay, Alaska, these phalaropes nested on sedge-grass marshlands, in
clearings in alder and willow scrub-covered slopes, and on heath-covered slopes
just above these slopes (Hohn 1968). Preferred nesting habitat was the lower
edge of the alder/willow scrub area. Kistchinski (1975) noted that Red-necked
Phalaropes in eastern Siberia nested more frequently along permanent bodies of
water than did the closely-related Red Phalarope.

In most Arctic areas these phalaropes have been reported to nest in marshes
with small ponds (Hilden and Vuolanto 1972 and authors cited therein). Nests
are made on moss in wet places or among sedges near the edge of the water;
these nests are often placed on small hummocks surrounded by water. Hilden and
Vuolanto (1972) found nest sites at the southernmost permanent breeding area
in Finland to be in considerably different habitat. Almost half (47.3%, n = 74)


---





Red-necked Phalarope


Table 8. Dates of occurrence for Red-necked Phalaropes in the coastal
southeastern United States (a).


Number
of Dates of occurrence
State occurrences Spring Fall


North Carolina 70+ 3 May-10 June
(10 July-4 Aug.


South Carolina
Georgia
Florida
-Atlantic coast
Florida-Keys

Florida
-Gulf coast
Alabama

Mississippi
Louisiana
Texas-coast


17 May-3 June
5 May-early June
21 Mar.-28 May


5 20 May

16 14 Mar.-25 May
12 29 Apr.-17 May
12 29 Apr.-17 May


30 Apr.-14 May
4 Apr.-10 May


:29 June)



:27 June-6 Jul






(2 July-18 Jul'


10 Aug.-25 Oct.
.) (5 Feb.)
11 Sep.-25 Oct.
24 July-3 Oct.
23 Aug.-2 Nov.
(20 Nov.-23 Jan.)
15 Aug.-15 Oct.
(14 Nov.-23 Nov.)


I:
y)


2 Oct.


(22 Nov.-21 Jan.)


6 Aug.-14 Oct.
(13 Nov.-6 Dec.)
10 Sep.-21 Sep.
12 Sep.
15 Sep.-31 Oct.
*)


(a) Kecords in parentheses may represent summering birds or early fall
migrants, or late fall migrants or wintering birds.


of the nests were 20 m (66 ft) or more from water, and most (78.6%, n = 70)
were on dry sites (sand, gravel, dry meadow) rather than on wet ones. All
nests found were in patches of low or sparse vegetation, usually grasses
(Festuca, Deschampsia, Puccinellia; 41.9%, n = 74), spike rushes (Eleocharis;
28.4%), or sedges (Carex; 21.6%). Hohn (1969) noted that this species tends
to nest farther inland than the Red Phalarope. Kistchinski (1975) reported a
density in eastern Siberia of 0.3-0.5 pairs/ha (0.7-1.2/ac) for birds nesting
in the Indigirka Delta, compared to 1-2 pairs/ha (2.5-4.9/ac) for the Red Phal-
arope. In an area south of the Kolymskaya Channel that Kitchinski regarded
as less desirable nesting habitat for Red Phalaropes, comparable figures were
0.5 pairs/ha (1.2/ac) for the Red-necked Phalarope and 0.9 pairs/ha (2.2/ac)
for the Red Phalarope.

All nests observed by Hilden and Voulanto (1972) were found in or near
colonies of Arctic Terns. These authors suggested that this association was
chosen by the phalaropes, pointing out that the phalaropes react to tern alarm
calls as do other species showing strong association with larids, and that the
highest nesting densities of these phalaropes in Finnish Lapland always occur
where there are nesting Arctic Terns.


Y





Red-necked Phalarope


Feeding Bianki (1967) remarked that Red-necked Phalaropes usually feed
in open water where bits of detritus collect. They are found less commonly
near the shoreline at high tide. Bent (1927) commented that both this species
and the Red Phalarope, when at sea, preferred to feed in tide rips, on or near
floating seaweed, and often near whales or schools of fish. Scott (1959) noted
migrant Red-necked Phalaropes feeding among rows of drifting seaweed off south-
western Nova Scotia. During fall migration along the Yukon coast, these phal-
aropes feed along beaches, in the lee of ice flows, and on the open ocean; they
preferred to feed in sheltered waters on windy days (Vermeer and Anweiler 1975).

Migrants inland are frequently seen on sewage ponds, rain pools, impound-
ments, and lakes. Cox (1973) suggested that Red-necked Phalaropes away from
pelagic habitats prefer shallow pools, in contrast to Red Phalaropes, which ap-
parently prefer deeper water.

Nonbreeding and Offshore During migration in the northern Chesapeake
Bight, just north of North Carolina, Red-necked Phalaropes occur somewhat closer
inshore than do Red Phalaropes; the former usually occur 20 km (12 mi) or more
offshore, the latter 70 km (44 mi) or more (Rowlett 1980). Migrants arriving
in Greenland and Iceland in the spring remain in large flocks in coastal bays
and deltas until inland nesting ponds are free of ice (authors cited by Hilden
and Voulanto 1972).


FOOD AND FEEDING BEHAVIOR

Red-necked Phalaropes feed primarily by surface-seizing (Ainley and Sanger
1979), i.e., sitting on the water and picking their prey from the surface with
their bills. Phalaropes may also turn rapidly in circles, a behavior referred
to as "spinning". This may stir up edible particles from the bottom in shal-
low water, but it also may serve to activate chilled invertebrates in deeper
water (Hohn 1971). Hohn suggested that this behavior occurs where there is a
dense concentration of food items, because the spinning motion allows the birds
to feed rapidly. Spinning may also play a role in courtship display (Hohn 1971,
Everett 1976).

Phalaropes occasionally "tip-up" like dabbling ducks, immerse their heads
and necks, and seize prey underwater (Hohn 1971). Diving has been recorded
once for this species (Selous 1915 in Hohn 1971), and some feed by using side-
to-side sweeps of the bill in shallow water (Hohn 1968). Benning (1971) noted
an instance in which this species hawked insects up to a foot or so above the
water.

Red-necked Phalaropes are gregarious and often form feeding flocks, par-
ticularly at the conclusion of spring migration when weather prevents entry to
the breeding grounds. This gregariousness extends into the breeding season.
During this period at Scammon Bay, Alaska, they often congregate into flocks
of about 20 birds (Hohn 1968).





Red-necked Phalarope


The only comprehensive study of the food habits of the Red-necked Phala-
rope in North America was conducted by Wetmore (1925), who examined 155 stom-
achs collected from May to October. He found that these shorebirds were almost
entirely carnivorous, with animal matter forming 97.2% by volume. Small insects
were the most important items of diet; flies (Diptera) constituted 32.8% of the
bulk, but true bugs (Hemiptera 31.8%) and beetles (Coleoptera 16.5%) were
also important. Of the flies, early developmental and adult stages of mosqui-
toes (Culicidae) and gnats (Chironomidae) were especially important; the larvae
of mosquitoes made up 6.3% of the sample. Most of the stomachs in which the
mosquito larvae were found evidently were collected on the breeding grounds in
Alaska. Among the other insect groups forming most of the diet a few taxa pre-
dominated. Water-boatmen (Corixidae) were the most significant food among the
Hemiptera eaten, and diving-beetles (Dytiscidae) and water-scavenger beetles
(Hydrophilidae) were most important among the coleopterids.

Crustaceans were eaten throughout the year but made up only a small pro-
portion (9.3%) of the diet; amphipods were frequently taken, and brine shrimps
(Artemia) were eaten extensively at Great Salt Lake, Utah, during migration.
Wetmore (1925) suggested that crustaceans probably form a considerably higher
proportion of the diet when these birds are at sea.

Other kinds of food that Wetmore (1925) recorded as occasionally or rarely
eaten by Red-necked Phalaropes included molluscs (snails or other gastropods),
other insects (largely those found in aquatic habitats), marine worms, spiders,
mites, and an unidentified fish. Plants comprised only a small portion of the
food (2.8%) but were regularly found in the stomachs; this material consisted
mostly of the seeds of plants characteristic of wet areas (e.g., Ruppia, Potamo-
geton, Scirpus).

Baker (1977) examined the contents of 24 stomachs collected at Churchill,
Manitoba, but did not report specific items of diet in detail, noting only that
these phalaropes preyed heavily upon adult chironomids. Stomachs of birds col-
lected on Kandalaksha Bay, U.S.S.R., contained only terrestrial insects (Bianki
1967) that evidently had been gleaned from the surface of the water. Dement'ev
and Gladkov (1951), in an earlier summary of food habits of Red-necked Phala-
ropes in the U.S.S.R., found that terrestrial insects were eaten only casually.
On the whole, prey taken in the U.S.S.R. was similar to that taken in North
America, i.e., aquatic insects and their larvae. Birds along the Caspian Sea
in fall mainly fed on amphipods, while those from the Murman Coast had eaten
molluscs, insects, vegetable matter, and pebbles (Dement'ev and Gladkov 1951).

Phalaropes on coastal islands in the Gulf of Bothnia fed primarily on larval
and adult Chironomidae in saline situations, but ate mostly Trichoptera larvae,
water fleas, tadpoles, water spiders, and collembolans in freshwater ponds (Hilden
and Vuolanto 1972).





Red-necked Phalarope


SUSCEPTIBILITY TO OIL POLLUTION

We found no reports of oiled Red-necked Phalaropes. Although they feed
on the surface in flocks, their offshore distribution, their short stay in most
southeastern waters, and their scarcity in parts of the southeast make them of
relatively low concern with regard to the development of petroleum resources.
This is probably not true in other areas. Their feeding habits and flocking
behavior on fall migration along the Yukon coast led Vermeer and Anweiler (1975)
to consider them perhaps the most vulnerable to oil pollution of all shorebirds
occurring there. King and Sanger (1979) agreed by giving the Red-necked Phala-
rope one of the highest ratings for vulnerability for any shorebird occurring
in the northeastern Pacific. They indicated that this species should be of
special concern vis a vis the potential effects of oil pollution.


BIBLIOGRAPHY

1982

Orr, C. D., R. M. P. Ward, N. A. Williams and R. G. B. Brown. 1982. Migration
patterns of Red and Northern Phalaropes in southwest Davis Strait and
in the northern Labrador Sea. Wilson Bull. 94: 303-312.

Ruttledge, R. F. 1982. Red-necked Phalarope--a hitherto unrecorded case of
breeding in Co. Mayo. Irish Birds 2: 196-197.

1981

Hohn, E. 0. 1981. Northern Phalarope flocks at Miquelon Lake, Alberta. Blue
Jay 39: 41-43.

Pigeon, P. 1981. Observation d'un Phalarope a bec etroit Phalaropus lobatus
et d'un Phalarope a bec large Phalaropus fulicarius a Aulnois-sous-Laon
(Aisne). Alauda 49: 232.

1980

Becker, P. 1980. Zum Vorkommen des Odinswassertreters (Phalaropus lobatus) an
den Kuste Sudwestafrikas und zur feldornithologischen Unterscheidung der
Wassertreter. Ornithol. Mitt. 32: 40-44.

Guex, C. 1980. Observation hivernale d'un Phalarope a bec etroit Phalaropus
lobatus sur le Leman. [Winter observations of the Red-necked Phalarope
Phalaropus lobatus on Lake Geneva.] Nos Oiseaux 35: 372. [In French.]

Hohn, E. and D. J. Mussell. 1980. Northern Phalarope breeding in Alberta.
Can. Field-Nat. 94: 189-190.

King, F. 1980. Red-necked Phalaropes breeding at Akeragh Lough in County
Kerry. Irish Birds 1: 540-541.





Red-necked Phalarope


Orsini, P. 1980. Premiere observation hivernale du Phalarope a bec etroit
Phalaropus lobatus dans le Moyen-Atlas (Maroc). [First winter observation
of Red-necked Phalarope Phalaropus lobatus in the Middle Atlas (Morocco).]
Alauda 48: 258-259.

1979

Holmes, G. 1979. First record of Red-necked Phalarope in New South Wales.
Austr. Birds 13: 75-76.

Roudybush, T. E., C. R. Grau, M. R. Petersen, D. G. Ainley, K. V. Hirsch, A. P.
Gilman and S. M. Patten. 1979. Yolk formation in some charadriiform birds.
Condor 81: 293-298.

Toups, J. A., M. B. Hodges and P. V. Donaldson. 1979. Northern Phalarope at
Pascagoula--first photographic record for Mississippi. Mississippi Kite
9: 9-10.

1978

Dunnett, G. M. and S. W. Kotogama. 1978. Red-necked Phalarope in Sri Lanka.
J. Bombay Nat. Hist. Soc. 75: 215-216.

Ruttledge, R. F. 1978. Red-necked Phalaropes breeding south of latitude 530
37' in Ireland. Irish Birds 1: 229-231.

Taylor, W. B. 1978. Seasonal status, population size, and habitat utilization
of Red and Northern Phalaropes in the Southern California Bight. (Abstract
only.) Pac. Seabird Group Bull. 5: 74.

1977

Ash, J. S. and 0. M. Ashford. 1977. Great Black-headed Gulls Larus ichthyaetus
and Red-necked Phalaropes Phalaropus lobatus inland in Ethiopia. J. East
Afr. Nat. Hist. Soc. 31: 1-3.

Baker, M. C. 1977. Shorebird food habits in the eastern Canadian Arctic.
Condor 79: 56-62.

Brichetti, P. 1977. II falaropo becco sottile, Phalaropus lobatus (L.) in
Italia. Uccelli d'ltalia 2: 129-131.

Gianella, C. 1977. Observazione di un falaropo becco sottile-Phalaropus
lobatus (L.)-nella Laguna di Ponente di Orbetello. Riv. Ital. Ornitol.
47: 293-294.

Muller, G. and H.-P. Keller. 1977. Beobachtungen von Weissflugelseeschwalben
(Chlidonias leucopterus), steinwalzer (Arenaria interpres) und Odinswasser-
treter (Phalaropus lobatus) in der Oberlausitz. Beitr. Vogelkd. 23: 60-61.





Red-necked Phalarope


Ruthke, P. 1977. Futtertragen und Dunenkleid des Odins-wassertreters (Phalar-
opus lobatus). [Food carrying and juvenile plumage of the Red-necked Phal-
arope Phalaropus lobatus.] Vogelwelt 98: 229-231. [In German with English
summary.]

Schiemann, H. 1977a. Uber das Vorkommen der Wassertreter (Phalaropodidae) in
den brandenburgischen sachsischen und thuringischen Bezirken, souie in Ber-
lin. Beitr. Vogelkd. 23: 49-56.

1977b. Beringungsergebnisse Nordeuropaischer Odinshunchen. [Banding
results of North European Red-necked Phalaropes.] Sterna 16: 73-80. [In
German with Norwegian summary.]

Sinclair, J. C. 1977. The phalaropes. Bokmakierie 29: 90-91.

1976

Crawford, P. and D. Crawford. 1976. Second record of Northern Phalarope in
Nashville area. Migrant 47: 94.

Everett, J. J. 1976. Mutual spinning by Red-necked Phalaropes. Brit. Birds
69: 219-220.

Gates, L. J. 1976. Northern Phalarope in Hattiesburg, Mississippi. Missis-
sippi Kite 6: 37.

McNamara, J. A. 1976. Two further records of the Red-necked Phalarope.
S. Austr. Ornithol. 27: 142.

Sanders, J. A. 1976. A Northern Phalarope in Attala County, Mississippi.
Mississippi Kite 6: 36.

Vanhercke, L. 1976. De grauwe franjepoot (Phalaropus lobatus). Wielewaal 42:
342-343.

1975

Dontschev, S. 1975. Neue Angaben uber das Auffinden von Phalaropus lobatus
(L.), Glareola nordmanni Fisch.-Waldh. und Arenaria interpres (L.) am der
bulgarischen Schwarzmeerkuste. Larus 26-28: 183-187.

Kistchinski, A. A. 1975. Breeding biology and behaviour of the Grey Phalarope
Phalaropus fulicarius in East Siberia. Ibis 117: 285-301.

Menon, S. S. 1975. Occurrence of the Red-necked Phalarope (Phalaropus lobatus)
on inland waters in Bangalore. J. Bombay Nat. Hist. Soc. 72: 846-847.

Nakamura, K. 1975. [On a Red-necked Phalarope encountered [in a] typhoon dur-
ing migration.] Tori 23: 95-96. [In Japanese with English summary.]





Red-necked Phalarope


Schmidt, G. 1975. Beobachtungen uber den Wegzug des Odinswassertreters aus
nordnorwegischen Brutgebieten an der Barents-See. Beitr. Vogelkd. 21: 233-
244. [In German.]

Vermeer, K. and G. G. Anweiler. 1975. Oil threat to aquatic birds along the
Yukon coast. Wilson Bull. 87: 467-480.

1974

Eller, G. D. 1974. Northern Phalarope in upper east Tennessee. Migrant 45: 96.

Kopke, G. 1974. Odinswassertreter (Phalaropus lobatus) im Juni auf Baltrum.
Vogelkd. Ber. Nieders. 6: 18.

Seeger, J.-J. 1974. Odinswassertreter am Gulper See. Falke 21: 284.

Steinbach, R. 1974. Vorkommer der wassertreter Phalaropus lobatus (L.) und
Phalaropus fulicarius (L.) im Gebiet der Haselbachar und Eschefelder Teiche
sowie am Speicherbechten Windischleuba. Abh. Ber. Naturkundemus. 'Maur-
itanium' 8: 333-338.

1973

Cox, J. B. 1973. A further occurrence of the Red-necked Phalarope in South
Australia. S. Austr. Ornithol. 26: 116-117.

Fevold, H. R., E. W. Pfeiffer and J. S. Rice. 1973. Steroid biosynthesis by
phalarope (Steganopus tricolor and Lobipes lobatus) adrenal tissue. Gen.
Comp. Endocrinol. 21: 353-357.

Smith, F. T. H. 1973a. Red-necked Phalarope at Altona saltworks (Phalaropus
lobatus). Bird Obs. 473: 12.

.1973b. Further to the Altona Red-necked Phalarope. Bird Obs. 477:
4-5.

Stelzer, M. 1973. Uber das Vorkommen der Wassertreter (Phalaropodidae) in
der Schweij und ihren Randgebieten. Ornithol. Beob. 70: 157-170.
[In German with French summary.]

van Avermaet, G. 1973. Franjepoten. Wielewaal 39: 216-220.

1972

Hilden, 0. and S. Vuolanto. 1972. Breeding biology of the Red-necked Phalarope
Phalaropus lobatus in Finland. Ornis Fenn. 49: 57-85.

Jovanovic, V. 1972. Novi podaci o seobi liskonoge tankokljune, Phalaropus
lobatus, u Vojvodini. Larus 24: 164.





Red-necked Phalarope


Raner, L. 1972. Forekommer polyandri hos smalnabod simsnappa och svartsnappa?
[Polyandry in the Northern Phalarope and the Spotted Redshank.] Fauna
Flora 67: 135-138. [In Swedish.]

Schiemann, H. 1972. Uber Winterquartiere nordeuropaischer Odinshuhnchen. [On
wintering areas of northern Red-necked Phalaropes (Phalaropus lobatus).]
Vogelwarte 26: 329-336. [In German with English summary.]

Vaisanen, R. A., 0. Hilden, M. Soikkeli and S. Vuolanto. 1972. Egg dimension
variation in five wader species: the role of heredity. Ornis Fenn. 49:
25-44.

1971

Benning, W. E. 1971. Phalaropes hawking insects over the water at Montezuma.
Kingbird 21: 11.

Edscorn, J. B. 1971. Northern Phalarope (Lobipes lobatus) occurrences in cen-
tral Florida. Fla. Nat. 44: 94.

Everett, M. J. 1971. Breeding status of Red-necked Phalaropes in Britain and
Ireland. Brit. Birds 64: 293-302.

Hohn, E. 0. 1971. Observations on the breeding behaviour of Grey and Red-
necked phalaropes. Ibis 113: 335-348.

McGregor, A. 1971. Rare birds: Red-necked Phalaropes. Bird Life 7: 2-5.

Pringle, J. S. and W. S. Pringle. 1971. Red-necked Phalarope Phalaropus lob-
atus at Strandfontein. Ostrich 42: 228-229.

Scholz, J. 1971. Odinswassertreter (Phalaropus lobatus) am Schlammteich bei
Artern. Beitr. Vogelkd. 17: 174-175.

Verheyen, J. A. 1971. Red-necked Phalarope Phalaropus lobatus in the Island
of Flores, Indonesia. Ardea 59: 64.

1970

Alsop, F. G., III. 1970. Northern Phalarope in the Great Smoky Mountains
National Park. Migrant 41: 39-40.

Coffey, J. W. 1970. Northern Phalarope in Washington County. Migrant 41:
66-67.

Haight, T. 1970. Northern Phalarope in Dutchess County. Kingbird 20: 179-180.

Hohn, E. 0. 1970. Gonadal hormone concentrations in Northern Phalaropes in
relation to nuptial plumage. Can. J. Zool. 48: 400-401.






Red-necked Phalarope


Lowery, G. H., Jr. 1970. The Louisiana state list. La. Ornithol. Soc. News
56: 1-7.

1969

Bates, A. K. and M. L. Bierly. 1969. White-rumped Sandpipers, Northern Phal-
aropes, Ruddy Turnstones, Gull-billed Terns and Stilt Sandpipers at Marion
Fish Hatchery. Ala. Birdlife 17: 52-54.

Eckert, J. 1969. Red-necked Phalarope in south Australia. Emu 69: 184-186.

Eriksson, K. 1969. Weibchenschwarme des Odinswassertreters (Phalaropus loba-
tus) und die Datierung ihrer Gelege in Finnisch Lappland. [On the flock-
ing of female Red-necked Phalaropes (Phalaropus lobatus) during the breed-
ing season and on the time of egg-laying in Finnish Lappland.] Ornithol.
Mitt. 21: 157-160. [In German with English summary.]

Hohn, E. O. 1969. The Phalarope. Sci. Am. 220: 104-111.

Schoennagel, E. 1969. Gehauftes Auftreten des Odinswassertreters (Phalaropus
lobatus) auf der Insel Borkum (Nordsee). Ornithol. Mitt. 21: 18.

Smith, F. T. H. 1969. The Red-necked Phalarope near Melbourne. Austr. Bird
Watcher 3: 166-167.

1968

Gerasimov, N. N. 1968. [Breeding of the Red-necked Phalarope in Kamchatka.]
Ornitologiya 9: 344-345. [In Russian.]

Glayre, D. 1968. Un Phalarope a bec etroit sejourne dix jours a Chavornay.
Nos Oiseaux 29: 275-276.

Hohn, E. 0. 1968. Some observations on the breeding of Northern Phalaropes at
Scammon Bay, Alaska. Auk 85: 316-317.

Masters, G. 1968. Black-necked Stilts and Northern Phalaropes at Jekyll Island,
Ga. Oriole 33: 50-51.

Plucinski, A. 1968. Odinswassertreter (Phalaropus lobatus) bei Goslar. Orni-
thol. Mitt. 20: 239.

1967

Kumerloeve, H. and P. A. D. Hollum. 1967. Zum durchzug von Phalaropus lobatus
(L.) in Kleinasien. Vogelwarte 24: 64-65.






Red-necked Phalarope


1966

Hohn, E. 0. 1966. Ringing (banding) and recoveries of phalaropes--a summary of
presently available information. Bird-Banding 37: 197-200.

Warfield, R. W. 1966. Vagrant Northern Phalarope at Lilypons. Md. Birdlife
22: 14-15.

1965

Dittberner, H. and W. Dittberner. 1965. Sturmmowe, Larus canus, brutend und
Odinswassertreter, Phalaropus lobatus rastend am Grossen Schwerin (Muritz).
Beitr. Vogelkd. 11: 103-105.

Hohn, E. 0. 1965. Die Wassertreter. Neue Brehn-Bucheri 359. Ziemsen Verlag,
Wittenberg. 60 pp.

Turk, M. 1965. Northern Phalarope at Dauphin Island. Ala. Birdlife 13: 42.

1964

Clark, W. S. 1964. An unusual visitor (Northern Phalarope). Ala. Birdlife
12: 12.

Rittinghaus, H. 1964. Phalaropus lobatus (Phalaropodidae) Nahrungserwerb.
Pp. 309-311 in G. Wolf (Ed.) Encyclopedia Cinematographica, Inst. fur
den Wissenschaftlichen Film, Gottingen. [In German.]

1963

Smith, F. T. H. 1963. An Australian sight record of the Red-necked Phalarope
(Phalaropus lobatus). Austr. Bird Watcher 2: 1-4.

1962

Longstreet, R. J. 1962. Northern Phalarope (Lobipes lobatus) on Lake Monroe.
Fla. Nat. 35: 92.

Pocock, T. N. 1962. Red-necked Phalarope Phalaropus lobatus and other birds at
Iscor Dams, Vanderbijl Park. Ostrich 33: 41-44.

1961

Smith, E. D. 1961. Northern Phalarope on Lake Murray, South Carolina.
Chat 25: 68.

1959

Hall, G. A. 1959. A late record for Northern Phalarope in West Virginia.
Wilson Bull. 71: 194.





Red-necked Phalarope


Ogden, J. 1959. Northern Phalaropes at Nashville. Migrant 30: 55.

1958

Bellier, L. and R. Leveque. 1958. Phalarope a bec etroit en Camargue. Alauda
26: 230. [In French.]

1957

Haftorn, S. 1957. Om foreldrenes omsorg for egg og unger hos svommesnipe,
Phalaropus lobatus (L.) and temmincksnipe, Calidris temminckii (Leisl.).
[On parental care in Phalaropus lobatus (L.) and Calidris temminckii
(Leisl.).] K. Nor. Vidensk. Selsk. Mus. Arbok 1957: 12-144.

Harrisson, T. 1957. Red-necked Phalarope, Lobipes lobatus (Linnaeus) in south-
east Asia. J. Bombay Nat. Hist. Soc. 54: 947.

Morzer Bruijns, W. F. J. and M. F. Morzer Bruijns. 1957. Waarnemingen van de
Grauwe Franjepoot, Phalaropus lobatus (L.), in de Indischm Oceaan. [Obser-
vations of the Red-necked Phalarope, Phalaropus lobatus (L.), in the Indian
Ocean.] Ardea 45: 72-84. [In Dutch with English summary.]

1956

Christensen, N. H. 1956. Odinshanens (Phalaropus lobatus L.) og Thorshanens
(Phalaropus fulicarius L.) forekomst i Danmark. [The occurrence in Den-
mark of the Red-necked Phalarope (Phalaropus lobatus L.) and the Grey
Phalarope (Phalaropus fulicarius L.] Dan. Ornithol. Foren. Tidsskr. 50:
191-206. [In Danish with English summary.]

Mester, H. 1956. Odinshuhnchen (Phalaropus lobatus) bei Frondenberg/Ruhr.
Ornithol. Mitt. 8: 53.

1955

Geroudet, P. 1955. Le passage des phalaropes en Suisse romande. Nos Oiseaux
23: 42-47. [In French.]

Stimson, L. A. 1955. Northern Phalarope. Fla. Nat. 28: 29.

1954

Maluquer, S. M. 1954. Phalaropus lobatus en Espana. Ardeola 1: 121-122

1953

Alexander, H. G. 1953. Rednecked Phalarope near Delhi. J. Bombay Nat. Hist.
Soc. 51: 507.





Red-necked Phalarope


Berndt, R. and R. Reinecke. 1953. Erstnachweis des Odinswassertreters, Phal-
aropus lobatus (L.) fur das Braunschweiger Hugelland. Ornithol. Mitt.
5: 6.

Meier, H. 1953. Schmalschnabliger Wassertreter im Urner Reussdelta. Ornithol.
Beob. 50: 32.

Mouchet, E. 1953. Le Phalarope a bec etroit aux Grangettes. Nos Oiseaux 22:
72.

1952

Eberhardt, R. L. 1952. Northern Phalaropes and Xanthus Murrelet associated
with fishes. Condor 54: 314.

Simon, H. 1952. Observation d'un Phalarope hyperbore dans le Calvados. Oiseau
Rev. Fr. Ornithol. 22: 322.

1951

Hanstrom, R. 1951. Den smalnabbade simsnappen, en originell svensk smavadare.
Fauna Flora 1/2: 57-66.

[Wood, J. D.]. 1951. Studies of some species rarely photographed XXXI. The
Grey [and] Red-necked Phalarope. Brit. Birds 44: 235-236.

1950

Nevin, W. S. 1950. Red-necked Phalarope on the Kent-Susses border. Brit.
Birds 43: 191-192.

1949

Longstreet, R. J. 1949. Flock of Northern Phalaropes at Daytona Beach, Florida.
Auk 66: 204.

van Ijzendoorn, A. L. J. 1949. A preening phalarope (Lobipes lobatus). Auk
66: 89-90.

1948

Walkinshaw, L. H. 1948. Nestings of some shorebirds in western Alaska. Condor
50: 220-223.

1945

Vaughan, R. 1945. A rare visitor. Country Life 97: 1001.





Red-necked Phalarope


1943

Cole, L. J. 1943. Behavior of Northern Phalarope with young. Condor 45: 39.

1938

Michael, C. W. 1938. Behavior of Northern Phalaropes. Condor 40: 85.

1937

Low, G. C. 1937. Grey and Red-necked Phalaropes in the Arabian Sea. Ibis
(14th Ser.) 1: 866.

Meinertzhagen, R. 1937. Grey and Red-necked Phalaropes in Arabian Sea. Ibis
(14th Ser.) 1: 667.

Sits, E. 1937. Beobachtungen uber Phalaropus lobatus an der Matsulu Bucht.
Ann. Soc. Nat. Univ. Tartu 43: 16-24.

Wust, W. 1937. Schmalschnabliger Wassertreter, Phalaropus lobatus (L.) bei
Ismaning erlegt. Anz. Ornithol. Ges. Bayern 2: 449-450.

1935

Tinbergen, N. 1935. Field observations of East Greenland birds. I. The behav-
iour of the Red-necked Phalarope (Phalaropus lobatus L.) in spring. Ardea
24: 1-42.

1934

Bonami, L. 1934. Cattura di un Falaropo a becco sottile (Phalaropus lobatus
[L.]). Studi Trentini 15: 196-197.

Chamberlain, E. B. 1934. Northern Phalarope in South Carolina. Auk 51: 519-
520.

1933

Sprunt, A., Jr. 1933. Second occurrence of Northern Phalarope in South Caro-
lina. Auk 50: 358-359.

Stone, W. 1933. Northern Phalaropes on the New Jersey coast. Auk 50: 475-476.

Taning, A. V. 1933. The winter quarters of the phalaropes. Ibis (13th Ser.)
3: 132-133.

Young, C. G. 1933. The winter quarters of the phalaropes. Ibis (13th Ser.)
3: 548.





Red-necked Phalarope


1932

Urner, C. A. 1932. Phalaropes in New Jersey in spring. Auk 49: 475-476.

1931

Glegg, W. E. 1931. On the nesting of the Red-necked Phalarope in Shetland.
Oologists' Rec. 9: 10-14.

1929

Poole, E. L. 1929. Northern Phalarope (Lobipes lobatus) in Pennsylvania. Auk
46: 108.

Townsend, C. W. 1929. Breeding range of the Northern Phalarope (Lobipes lob-
atus). Auk 46: 108-109.

1928

Schorger, A. W. 1928. The wintering area of the Red and Northern Phalaropes.
Auk 45: 206.

1927

Momiyama, T. 1927. [On the specimens of Lobipes lobatus obtained at Prefect.
Saiitama, Hondo.] Tori 5: 267-271. [In Japanese.]

1926

Grote, H. 1926. Zur Verbreitung von Phalaropus lobatus (L.) und Phalaropus
fulicarius (L.). Ornithol. Monatsber. 34: 122.

Spingarn, E. D. W. 1926. Northern Phalarope in Dutchess County, N.Y. Auk
43: 90-91.

1925

Meinertzhagen, R. 1925. The distribution of the phalaropes. Ibis (12th Ser.)
1: 325-344.

Rapine, J. 1925. Les Phalaropes. Rev. Fr. Ornithol. 17: 54-56.

Wetmore, A. 1925. Food of American phalaropes, avocets, and stilts. U.S. Dep.
Agric. Bull. 1359. 20 pp.

1923

Tschusi zu Schmidhoffen, V. 1923. Phalaropus fulicarius, P. lobatus und
Acanthis flavirostris in Oberosterreich. Ornithol. Monatsber. 31: 62.





Red-necked Phalarope


1921

Gordon, A. 1921. A note on the nesting of the Red-necked Phalarope. Brit.
Birds 15: 90-91.
1915

Selous, E. 1915. A diary of ornithological observations made in Iceland
during June and July, 1912. Zoologist 19 (Ser. 4): 54-56, 169-174,
303-307, 20: 54-68, 139-152, 267-272.

1914

Best, M. G. S. and M. D. Haviland. 1914. Notes on the Red-necked Phalarope
in the Outer Hebrides. Brit. Birds 8: 9-12.

1910

Brimley, H. H. 1910. Northern Phalarope in Bladen County, North Carolina.
Auk 27: 206.

Morris, R. O. 1910. Capture of the Northern Phalarope near Springfield,
Mass. Auk 27: 79.

1909

Smith, C. B. 1909. "Sundhani". Avic. Mag. 7: 309-311.

1908

Bahr, P. H. 1908. Some observations on the breeding habits of the Red-necked
Phalarope. Brit. Birds 1: 202-207.

Service, R. 1908. Red-necked Phalarope in Solway Area. Ann. Scott. Nat. Hist.
1908: 253.

1907

Gladstone, H. S. 1907. The Red-necked Phalarope in Ireland. Brit. Birds 1:
174-177.

1905

Wayne, A. T. 1905. Notes on certain birds taken or seen near Charleston,
South Carolina. Auk 22: 395-400.

1903

Williams, E. 1903. Breeding of the Red-necked Phalarope in Ireland. Irish
Nat. 12: 41-45.




Red-necked Phalarope


1901
Forrest, J. 1901. Red-neked Phalarope (Phalaropus hyperboreus) in North
Wales. Zoologist 1901: 428.
Gaal de yula, 1901. Der lobatus (L.) in der Vogelfauna des
alaon-See. Aquila 8: -

1880
Loom, L. 1880. The Northern Phalarope in Chester County, South Carolina.
Bull. Nuttall Ornthol. Club 5: 242.

1878
Coues 1878. The Northern Phalarope n North Carolina. Bull. Nuttall
Ornithol. Club 3: 40-41.


Red-necked Phalarope in winner plumage. Photograph by Clayton Taylor.





~s~r

~-C~h~c_ -1L~










RED PHALAROPE


(Phalaropus fulicaria)


[DA: Thorshane, DU: Rosse franjepoot, EN: Grey Phalarope, FI: Vesipaasky, FR:
Phalarope a bec large, GE: Thorshuhnchen, Thorswassertreter; IC: Thorshani,
IT: Falaropo a becco largo, JA: Hai-iro hire-ashi shigi, NW: Polarsvommesnipe,
PO: Platkonog plaskodzioby, PR: Falarodo, RU: (Flat-nosed Phalarope), SP: Fal-
aropo picogrueso, Chorlillo norteno; SW: Brednabbad simsnappa]


GENERAL DISTRIBUTION

North America The breeding range of the Red Phalarope in North America
overlaps that of the Red-necked Phalarope but extends considerably farther north.
This species breeds from western and northern Alaska east across Canadian Arctic
islands to Baffin Island and north to northern Ellesmere Island. Along the
Canadian mainland it breeds in northern Mackenzie, eastern Keewatin, northern
Quebec (AOU 1957, Godfrey 1966), and probably northern Labrador (Godfrey 1966).

North American populations migrate south along both coasts and in the Paci-
fic and Atlantic Oceans to winter off South America (AOU 1957).

World Distribution In the Old World, Red Phalaropes breed from Greenland
and Iceland east through the Arctic islands of Spitsbergen and southern Novaya
Zemlya to the Novosibirskiye Islands. Along the mainland they breed across
northern Siberia from the Taimyr Peninsula east to the Chukotski Peninsula and
the coast of Anadyrland (AOU 1957, Vaurie 1965).

Primary wintering grounds are at sea off both coasts of South America
south to Patagonia and the Falklands, off west Africa south to southern Arabia,
and probably in the Pacific off the Bonin Islands and Seven Islands of Izu
(Vaurie 1965) south to New Zealand (AOU 1957).


DISTRIBUTION AND ABUNDANCE IN THE COASTAL SOUTHEASTERN UNITED STATES

North Carolina Red Phalaropes occasionally occur along the coast of North
Carolina (Pearson et al. 1942) and are seen inland infrequently. Through the
early 1960's there were few records, but with the recent increase in offshore
observations, it has become clear that the Red Phalarope is a regular migrant
that is sometimes encountered in large numbers (Lee and Booth 1979). Most mi-
gration occurs between 30 September and 29 December, and again between 8 March
and 4 May according to Lee and Booth (1979). They suggested that the species
may winter in North Carolina waters because large numbers are seen in late
December and early April.

Lee and Booth (1979) implied there were at least 43 records of Red Phala-
ropes occurring in North Carolina. We found about 40 published records, but






Red Phalarope


list below only coastal records that involve five or more birds. Seven or
eight records are for inland localities where these phalaropes occur from
30 September through 21 March. There are records for all intervening months
except February; all other records are from the coast or offshore.


6 coll. at Bodie Island


1907 17 Apr. 6 coll. at Bodie Island


1937 1, 3 May 2,8 seen at Pea Island NWR


1956 14 Apr. 23 seen (3 coll.) at Pea Island NWR


1965 24 Apr.

1966 9 Apr.

1966 23 Apr.

1966 4 Sep.

1973 6 Dec.

1974 14 Nov.


1975 26 Jan.

1975 Aug.-Oct.

1977 29 Dec.


1978 4 Apr.


17 Apr.

14 Nov.

5 Dec.


72

149

1,000+

88

200

150


28

"a few"


seen 30 mi ESE Wrightsville Beach

seen S of Beaufort

seen 6 mi off Wrightsville Beach

seen off Hatteras Inlet

seen 5 mi off Charleston

seen in 3 flocks, 40-50 mi NE
Charleston

seen inshore, Wrightsville Beach

seen in Cape Hatteras area


100's-1,000's seen off North Carolina


100's-1,000's seen off North Carolina


1,100 seen in Gulf Stream off Oregon Inlet

8 seen off Oregon Inlet

100's-1,000's seen off North Carolina


Apr.

1907 8 Apr.


1896 2 or 3 ca. 12 found dead at Cape Lookout Lighthouse


Pearson et al.
1942

Pearson et al.
1942

Pearson et al.
1942

Pearson et al.
1942

Chamberlain
1956b

Jones 1965

Parnell 1966b

Parnell 1966b

Parnell 1967a

Teulings 1974b

Teulings 1975a


Teulings 1975b

Teulings 1976a

Lee and Booth
1979

Lee and Booth
1979

Teulings 1978a

LeGrand 1979a

Lee and Booth
1979










1978 30 Dec.

1979 23 Apr.


100's-1,000's seen off North Carolina


1,000+


seen off Oregon Inlet


Red Phalarope



Lee and Booth
1979
LeGrand 1979c


South Carolina Burton (1970) considered the Red Phalarope the rarest of
the three phalaropes in South Carolina. He added two records to the two cer-
tain ones reported by Sprunt and Chamberlain (1949). There are four additional
records, all from offshore. The numbers seen suggest that the status of this
species in South Carolina is similar to its status in North Carolina, i.e.,
common to abundant in late fall and early winter, as well as in early spring.
The birds seen in 1898 were not positively identified as Red Phalaropes, but
considering the difference in seasonal abundance between this species and the
Red-necked Phalarope off the Atlantic coast, this identification is the most
probable.


1898 17 Mar. "enormous seen ca. 50 mi off northern coast
flocks"


1900 4 Dec.


1934 22 Apr.

1960 11 June


1963 15 Sep.


1964 12 Mar.

1972 30 May

1973 6 Dec.

1974 14 Nov.


1 male caught at Mt. Pleasant


1 seen near Charleston Lightship

1 seen off Cape Island near McClellans-
ville

1 seen on pond near Wallace River,
Rantowles, Charleston Co.

21 seen 26-30 mi SE Charleston Harbor

2 seen offshore 35 mi ESE Charleston

ca. 200 seen 5 mi off Charleston

125 seen 40-50 mi NE Charleston


Sprunt and Cham-
berlain 1949

Sprunt and Cham-
berlain 1949

Sprunt 1934

Baldwin 1960


Shuler 1964


Sykes 1966


Teulings 1972c

Teulings 1974b

Teulings 1975a


Georgia Denton et al. (1977) considered the Red Phalarope accidental in
Georgia and listed only two records. One phalarope was collected 29 October
(Denton et al. 1977) or 5 November 1970 (Teulings 1971a) inland at Thomaston,
50 mi NE of Columbus. Another was seen 28 mi off St. Simons Island 8 March
1975 (Teulings 1975b). We know of no records since then but believe that
these phalaropes are almost certainly abundant offshore; they go unrecorded
for lack of observation.

Florida Atlantic Coast Kale (1979 ms a) reported that this species is
rare offshore during migration and in winter. The number of reports for the
Florida Atlantic coast are few (ca. 37), but the numbers seen offshore during
recent surveys make it clear that Red Phalaropes are common and at times abun-
dant. Too few data are available to delineate periods of peak abundance or to





Red Phalarope


estimate total populations occurring offshore.

Howell (1932) and Sprunt (1954) together listed eight observations for
the Atlantic coast of Florida; we found another 29, almost half of them made
since 1970. We list below only those published subsequent to Sprunt (1954)
and those known to involve more than one bird. Extreme dates of occurrence
range from 23 July to 27 April, but a preponderance (ca. 85%) of the records
are from 14 August to 27 March.

1956 2 Jan. small numbers seen 10-20 mi off Cocoa Beach Stevenson 1956b

1956 27 Mar. small numbers seen 10-20 mi off Cocoa Beach Stevenson 1956b


1959 7 Feb. 7 seen 10 mi off Miami


1960 Mar. ca. 500


1964

1967

1971


1971

1971

1972

1972

1973

1973

1973

1974

1975

1977

1977


July

Feb.

Apr.


Aug.

Sep.

Jan.

July

Jan.

Aug.

Sep.

July

Sep.

Jan.

Sep.


20

22

2


31

500

20-100

2

19

8

120+

4

67

400+


seen off Canaveral


seen 18 mi off Cocoa

seen at Jacksonville Beach

seen 10-30 mi E Port
Canaveral

seen 15-28 mi off Cocoa

seen off Port Canaveral

seen off Jacksonville

seen off Cocoa

seen off Mayport

seen off Canaveral

seen 24 mi off Mayport

seen 20 mi off Cocoa

seen on peak count off Mayport

seen 10-15 mi off Canaveral


20 seen off Boynton Inlet


Stevenson 1959b

Stevenson 1960b,
1961

Stevenson 1964b

Stevenson 1967b

Kale 1971


Ogden 1971

Robertson 1972

Stevenson 1972a

Ogden 1972

Woolfenden 1973

Edscorn 1974

Edscorn 1974

Ogden 1974

Edscorn 1976

Stevenson 1977

Edscorn 1978


Florida Gulf Coast Kale (1979 ms b) considered this species extremely
rare on the Florida Gulf coast and inland. Sprunt (1954) listed three records
from the coast, and we found only seven since, listed below. The temporal span






Red Phalarope


of these records, all but one involving single birds, is 29 October to 8 May;
half of these sightings occurred in November and December. The status of this
species offshore may be quite different. Red Phararopes were regularly reported
off Pensacola (mostly by commercial fishermen) from 1946 through 1954, in large
numbers. Reports published in American Birds and by Weston (1953) reveal that
birds were seen on at least 26 individual dates during this period. Most of
these were summarized by Weston (1953), who concluded that the Red Phalarope
could be "considered a regular, and sometimes common, winter resident of the
middle northern Gulf of Mexico." He added that all but a few observations were
made more than 30 miles out in the Gulf and that only one observation was made
as near as 5 miles. These observations spanned a period from 13 October (Weston
1953) through 11 April (Newman 1954); the largest numbers of birds were seen
from late December through late February. We do not list these offshore records
or the three records given by Sprunt (1954) here, but include them in Table 9.

1957 21 Apr. 1 seen on inland pond at Pensacola Newman 1957b

1959 7 May 3 seen 5 mi off Manasota Key Stevenson 1959c

1962 30 Nov. 1 found inland at WCTV Tower Stevenson 1967a

1965 8 May 1 seen near Alligator Point Cunningham 1965b

1966 21 Nov. 1 found inland at WCTV Tower Stevenson 1967a

1967 6 Dec. 1 seen inland at Lake Talquin Stevenson 1968a

1975 27 Nov. 1 found injured at Navarre Purrington 1976

Alabama Imhof (1976b) considered the Red Phalarope regular and sometimes
common in winter off the Alabama coast, primarily on the basis of observations
made out of Pensacola Bay. Imhof (1976b) listed five records off Pensacola
from 29 October 1946 (Weston 1947) through 23 December 1954 (Newman 1955).
He pointed out that these observations could have been made off either the Ala-
bama or Florida coasts. We have chosen to include these observations with the
Florida Gulf coast records for the purposes of Table 9. The only other observa-
tions available to Imhof (1976b) were four from inland; we know of only two sub-
sequent reports, one from inland, one from offshore. Numbers present in Alabama
waters remain a mystery. Duncan and Havard (1979 ms) saw only one bird off Ala-
bama in several years of observation, but they mentioned that commerical fisher-
men in Alabama report large numbers of phalarope-like birds in winter.

1924 last half 1 coll. inland at Pickett Springs Holt 1924
Jan.

1968 20 Oct. 1 seen inland in Shelby Co. Purrington 1969


1972 5-11 Jan. 1 seen inland at Lake Purdy Imhof 1976b,
James 1972






Red Phalarope


1972 10-15 Sep. 1


1976 May


photogr. inland at Lake Purdy


Imhof 1976b,
1973


1 female seen 100 km off Alabama coast Duncan and
Havard 1979 ms


1978 5 Nov.


1 seen inland at Eufaula Refuge


Mississippi We know of only three records for the Red Phalarope from Mis-
sissippi. One was seen inland on the Hattiesburg sewage lagoons on 9 October
1977 (Gates and Runzo 1978) and another was seen there 12 to 15 October 1980
(Moore et al. 1981). The only record from the coast is of one seen off Biloxi
on 30 September 1978 (Purrington 1979). It is likely that the species is
more common offshore.

Louisiana We know of only three records of Red Phalaropes in Louisiana:
one was collected 12 October 1950 on the Baton Rouge Campus of Louisiana State
University; another was collected 16 September 1961 1 mi W of Holly Beach,
Cameron Parish; and one was seen 29 November-10 December 1970 at the Natchi-
toches Fish Hatchery (Lowery 1974). Lowery commented that these phalaropes
should occur in offshore waters in winter.

Texas According to Oberholser (1974), Red Phalaropes are casual in Texas
in fall and accidental in spring. We know of only 18 records of this species
in Texas, 11 of which are listed by Oberholser (1974). We list below only re-
cords from coastal areas and those published subsequent to Oberholser (1974).
Two-thirds of the records were made inland and all but four records were made
in the fall or early winter (8 September-24 November). There are three spring
records (1 April-31 May), and one summer record (15 July); the latter was pro-
bably an early fall migrant. These four records are all from the coast, sug-
gesting that Red Phalaropes are more common there than inland in spring and
summer.


1935 1 Apr.

1952 5-6 Oct.

1952 9 Oct.

1973 19-28 Sep.

1975 13 Sep.-
1 Nov.

1975 15 Nov.


seen at Cove


I

1


seen at Rockport


1 seen on W shore of Laguna Atascosa

1 seen inland at Austin sewage pond

2 seen inland at Austin sewage ponds


1 seen inland at Mitchell Lake, San
Antonio


1 female seen at
Galveston


Oberholser 1974

Oberholser 1974

Oberholser 1974

Webster 1974a

Webster 1976a


Webster 1976a


Webster 1976c


Purrington
1979


1976 31 May


Bolivar Flats,






Red Phalarope


1977 8 Mar. 11 seen (incl. 4 females) off Port Webster 1977c
Aransas

1978 15 July 1 photogr. at Laguna Atascosa NWR Webster 1979a

1978 4 Nov. 1 seen inland at Austin sewage ponds Webster 1979a


SYNOPSIS OF PRESENT DISTRIBUTION AND ABUNDANCE

Breeding The Red Phalarope has the most northerly distribution of the
three phalaropes, breeding circumpolarly in the northern Holarctic between 60*
and 82 N latitude (BOU 1971). They breed from northern and western Alaska
through northern Quebec, Greenland, and Iceland, along the Arctic Eurasian
coast, and on islands from Spitsbergen east to Siberia.

Winter The winter range of the Red Phalarope is at sea and believed to
be primarily off the coasts of southern South America, western Africa, and
southern Arabia.

Migration Red Phalaropes migrate along continental coastlines and over
the broad expanses of the Atlantic, Pacific, and Indian Oceans. They are
common migrants off the southeastern Atlantic coast of the United States that
occasionally winter. They occur later in the fall and earlier in the spring
along the Atlantic coast than do Red-necked Phalaropes (Lee and Booth 1979,
Tables 9, 10). Along the Pacific coast of the United States Red Phalaropes mi-
grate about one month later than do Red-necked Phalaropes. The spring migra-
tion there is short and usually near the coast; fall migation is longer and
birds are found over a wider area and farther out to sea (Taylor 1978).

Migrants usually occur singly or in small flocks. Rowlett (1980) report-
ed that Red Phalaropes in the northern Chesapeake Bight are seen in flocks of
20 to 80 birds in April and December, Scott (1959) observed that migrants in
the Bay of Fundy in August generally occur in flocks of 15-30 birds, but noted
two flocks that contained at least 500 birds.

The status of the Red Phalarope in the northern Gulf of Mexico is largely
unresolved and there are few recent records from any state, leading Duncan and
Havard (1979 ms) to regard it as casual in occurrence. On the other hand, scat-
tered observations (Weston 1953, Duncan and Havard 1979 ms) suggest that it may
be periodically, perhaps regularly, common in some offshore areas. More exten-
sive pelagic surveys are needed to define its temporal, geographical, and numer-
ical distribution in the northern Gulf. Although Red Phalaropes occur commonly
and are at times numerically abundant off the southeastern coast, the proportion
of the world or continental population of the species that occurs there must be
very small.






Red Phalarope _


Table 9. Approximate number of Red Phalaropes recorded by month for the
coastal southeastern United States (a).


State/region JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC


North Carolina 28 1 14 3456 10 4 96 5 160 503
South Carolina 21 1 2 1 1 125 200
Georgia 1 -
Florida-Atlantic Coast 447 30 503 41 26 41 689 1 1 5

Subtotal-ATLANTIC COAST 475 31 539 3498 12 1 26 45 787 6 286 708

Florida-Gulf Coast 227 667 87 36 4 24 35 1033
Alabama 2 1 1 1 1 -
Mississippi 1 2 -
Louisiana 1 1 1 1
Texas-Coast 11 1 1 1 2 -

Subtotal-GULF COAST 229 667 98 37 6 1 3 30 37 1034

TOTAL-ALL AREAS 704 698 637 3535 18 1 27 45 790 36 323 1742

(a) Birds found dead in the first 10 days of a month are arbitrarily assigned
to the previous month. We assumed only one was seen if the source did
not specify whether one or more was seen. If an indefinite number was
seen (e.g., "a few", "numbers", "hundreds to thousands"), or if a range
was given, we have assumed the smallest number implied by the statement.


Table 10. Dates of occurrence for Red Phalaropes in the coastal southeast-
ern United States.


Approximate
number of
State occurrences Dates of occurrence (a)


North Carolina 43+ 4 September 4 May (29 May, 19-30 August)
South Carolina 9 15 September 22 April (30 May-11 June)
Georgia 2 29 October 8 March
Florida-Atlantic Coast 37 14 August? 27 April (28 July-2 August)
Florida-Gulf Coast 36 13 October 21 April (7-8 May)
Alabama 6 10 September 15+ January (May)
Mississippi 2 30 September 9 October
Louisiana 3 16 September 10 December
Texas 6 5 October 1 April (31 May, 15 July)

(a) Exceptional dates of occurrence are listed in parentheses.






Red Phalarope


HABITAT

Nesting Red Phalaropes nest primarily in low coastal Arctic tundra. May-
field (1979) described the nesting habitat at Bathurst Island in the Canadian
Arctic as sedge-moss tundra interspersed with freshwater ponds. Similar habi-
tat in eastern Siberia was described by Kistchinski (1975) as polygonal and
tussocky moss-sedge tundra rich in swamps, lakes, and temporary ponds. These
birds usually nest near water, but the "ponds" may be very small, sometimes
only a few square meters in extent (Portenko 1921). Hohn (1969) noted that Red
Phalarope nests are never found more than a few miles inland, pointing out that
this species is more coastal in its nesting distribution than is the Red-necked
Phalarope.

Red Phalaropes on Bathurst Island in the Canadian Arctic usually place
their nests among sedges tall enough to cover them but not thick enough to
completely conceal the eggs (Mayfield 1979). Mayfield discerned no relation-
ship between nest-site and prominent features of the habitat (e.g., ponds,
marsh edge, boulders) except that nests were always on the wetter portions of
flats. In eastern Siberia, Red Phalarope nests were found mostly (84.9%, n =
93) among sedges near the edges of temporary ponds or on tussocks in flooded
swamps. They nested less frequently along permanent lakes (7.5%), on islets
in small lakes (7.5%) or on dry tussocks and ridges in the polygonal tundra
(7.5%) (Kistchinski 1975). Sites reported elsewhere include the mossy rim of
an Ivory Gull (Pagophila eburnea) nest on bare Seymour Island near Bathurst
Island (MacDonald in Mayfield 1979), piles of seaweed in the open in Spitsber-
gen (Munsterhjelm and LeRoi in Lovenskiold 1964), and shingle (Lovenskiold
1964).

Red Phalaropes, like other phalaropes, tend to nest near one another.
Some observers (Kistchinski 1975, Mayfield 1979) have referred to this as clus-
tering of nests; others consider the species to be colonial (Lovenskiold 1964,
Kozlova 1961 in Mayfield 1979). Hohn (1965 in Mayfield 1979) stated that all
three phalaropes nested "...mostly in loose colonies of four to eight pairs;
solitary pairs are rarer." Mayfield (1979) found groups of three to five
nests at Bathurst Island, while the largest colony found by Lovenskiold (1964)
in Spitsbergen contained 25 pairs.

Nest density, in what Mayfield (1979) considered optimum habitat from 1970
to 1976 at Bathurst Island, averaged 4.9 nests/sq km (12.9 nests/sq mi), but the
number of nests present in the 1 sq km (0.39 sq mi) study plot varied from 0 to
14 (0-36.3 nests/sq mi) from year to year. Mayfield pointed out that nest den-
sities reported in other areas were much greater. Citing other authors, he
mentioned densities of 7 and 15 pairs/sq km (18.1 and 38.9 pairs/sq mi) on
southwestern Baffin Island, and estimates of 20 and 12 pairs/sq km (51.8 and
31.2 pairs/sq mi) for Cambridge Bay, Victoria Island, and Jenny Lind Island,
respectively. At a 0.67 sq km (0.26 sq mi) study area near Barrow, Alaska, in
1974 and 1975, densities of 9.0 and 24.0 nests/sq km (23.3 and 62.2 nests/sq
mi) were found by Schamel and Tracy (1977). One particularly favorable marsh
held 90% of all nests at a density of 44.6 nests/sq km (115.5 nests/sq mi).
Mayfield (1979) also noted earlier papers that suggested concentrations of 50-





Red Phalarope


100 males/sq km (130-260 males/sq mi) in Siberian nesting areas, but he remark-
ed that "the units of area may be misleading if the best nesting areas are
attenuated and discontinous."

Feeding On the breeding grounds, Red Phalaropes feed in or near their
nesting habitats. Phalaropes near Barrow, Alaska, fed extensively at the edges
of shallow (1-2 cm) ponds (Schamel and Tracy 1977). In eastern Siberia, incu-
bating males fed both on temporary ponds and permanent lakes; the latter are
used later in the season, after the temporary ponds have dried up, and may be
far from the nests (Kistchinski 1975). Red Phalaropes breeding near lagoons
and the sea on the Chukchii Peninsula fed near the surf, in the estuaries, and
"wherever else they can find much food." When the sea was rough these birds
fed between the surf and the shore run-off, but preferred calmer waters such
as bays and lagoons (Portenko 1972).


Nonbreeding and Offshore Prebreeding phalaropes arriving at the breed-
ing grounds on the Chukchii Peninsula remain around lagoons where the ice opens
sooner than on the sea (Portenko 1972). Postlaying females and nonbreeding
males in eastern Siberia depart from the nesting grounds and move to lakes and
coastal lagoons; by mid-July virtually all phalaropes except incubating males
are found along the seashore (Kistchinski 1975). Some females occur well off-
shore, particularly among ice floes (Portenko 1972). During migration, the Red
Phalarope is the most pelagic of the phalaropes (Mayfield 1979) and is seldom
seen inshore. Migrant Red Phalaropes in the northern Chesapeake Bight, north
of North Carolina, occasionally fed around small clumps of sargasso weed (Sar-
gassum sp.) and along rip tides containing detritus (Rowlett 1980). Red Phal-
aropes seen here were usually more than 70 km (44 mi) from shore.


FOOD AND FEEDING BEHAVIOR

Red Phalaropes, like Red-necked Phalaropes, feed primarily by surface-
seizing, and like that species often exhibit the characteristic "spinning" be-
havior. Ridley (1980) recently described differences in feeding techniques
for three situations: (1) birds on the water and feeding on prey below the sur-
face may immerse the head and neck with the bill pointing vertically downward
while tipping up (much in the manner of a dabbling duck); (2) birds feeding
on land and at the edge of a pool probe and peck vertically downward; and (3)
birds feeding on organisms on the surface or on vegetation seize their prey
with the beak held horizontally. Ridley termed these three techniques "deep-
feeding", "edge-feeding" and "surface-feeding", and concluded that females
on the breeding grounds in Svalbard consistently prefer deep-feeding and that
incubating males prefer surface-feeding. Ridley also pointed out that 80% of
all pecks were successful in obtaining prey and noted that high rates of peck-
ing were observed in surface-feeding birds while considerably lower ones were
seen in edge- and deep-feeders.

Like Red-necked Phalaropes, Red Phalaropes are gregarious throughout the
year. They often gather in feeding flocks of four or five birds during the





Red Phalarope


breeding season, but incubating males may also feed solitarily (Kistchinski
1975). Most feeding probably occurs by day, but these phalaropes have also
been seen feeding on bright nights (Portenko 1972).

Wetmore (1925) provided the only detailed account of the diet of North
American Red Phalaropes, on the basis of 36 stomachs collected from May to Nov-
ember, mostly from the Pribilof Islands. These phalaropes consumed almost en-
tirely animal food. Insects made up most of the diet; the most important food
items were beetles (Coleoptera), which made up 27.3% of the bulk, and flies,
which accounted for 22.7%. A considerable variety of species was eaten, but
only dung-flies (Scatophaga) were found in a large proportion (38.8%) of the
stomachs. Crustacea were also frequently taken and represented 33.5% of the
bulk eaten. Amphipods were found in 14 stomachs and water-fleas (Daphniidae)
were found in three. A number of very small fishes were also eaten; those
that could be identified were sculpins (Cottidae). A few spiders, molluscs
and other insects were eaten infrequently.

Little recent information from North America is available. Hohn (1971)
examined three stomachs from birds collected in June in western Hudson Bay.
One contained larval chironomids and small eumelibranchs; the others contained
seeds or willow catkins and flowers. Wander (1981) observed a phalarope feed-
ing on the carcasses of a Red Knot (Calidris canutus) and a Herring Gull at
Jamaica Bay Wildlife Refuge in New York City.

LeRoi (in Lovenskiold 1964) remarked on the diet of Red Phalaropes on
Svalbard, basing his comments on an examination of the contents of 58 stomachs.
Of these, 67.2% contained Crustacea, primarily either gammarids or ostracods,
depending on where the birds were collected. Also found in a significant pro-
portion of the stomachs were molluscs (46.6%), dipterid larvae (15.5%), and
algae (22.4%); a few arachnids, annelids, and one beetle were also taken. Rid-
ley (1980) based a more recent report of food eaten on Svalbard on an analysis
of 21 faecal samples obtained during the 1978 breeding season. The diet con-
sisted primarily of small flies; chironomids were most important, but myceto-
philids also were eaten commonly. These and a few other flying insects were
present in 95% of the samples and were abundant (i.e., more than 10 individual
remains) in nearly half of them. Spiders were also important and were found
in 63.6% of the samples; the remains of unidentified Crustacea were found in
22%. Ridley (1980) noted that different foods were taken in different habitats.
Phalaropes feeding below the surface of the water took fly larvae and Crustacea,
surface-feeding birds took flies, and those feeding on land or at the edge of
the water took spiders, flies, and (perhaps) Collembola.

In experimental studies, Dodson and Egger (1980) found that Red Phalaropes
prefer to feed on larger zooplankton (particularly those more than about 1 mm
long). They chose cladocerans over copepods and chose Daphnia middendorffiana,
D. pulex, Eurycercus lamellatus, and Diaptomus bacillifer most frequently. Two
species of small calanoid copepods (Diaptomus bacillifer and Eurytemora canaden-
sis) were taken more frequently than was predicted on the basis of their size.
Dodson and Egger suggested that this may have occurred (1) because phalaropes
are somehow specialized for catching copepods; (2) because the smaller, slower






Red Phalarope


food items are more easily caught; and (3) because the smaller copepods are
more visible to the phalaropes than the larger, more transparent cladocerans.


SUSCEPTIBILITY TO OIL POLLUTION

We found little record of Red Phalaropes being affected by oil. They are
a gregarious, surface-feeding bird that occurs in large numbers off the Atlan-
tic coast, however, and are potentially moderately susceptible to oiling, at
least during cold weather. The strongly pelagic distribution of this species
suggests that it would be vulnerable only to large offshore spills and not to
inshore development.

King and Sanger (1979) gave the Red Phalarope a high oil vulnerability in-
dex value indicating that they are among the most vulnerable shorebirds occur-
ring in the northeastern Pacific region. In general, phalaropes are much less
susceptible to oiling than other marine birds. Connors and Gelman (1979) re-
ported that naive young Red Phalaropes initially were equally likely to choose
pans of food that did or did not have a thin film of oil on the surface of the
water. Experienced birds preferred clear water and fed for longer periods. As
a result of these observations, Connors and Gelman suggested that phalaropes
can easily learn to avoid oiled surfaces.


BIBLIOGRAPHY
1982

Conners, P. G. and K. G. Smith. 1982. Oceanic plastic particle pollution:
suspected effect on fat deposition in Red Phalaropes. Mar. Pollut.
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Orr, C. D., R. M. P. Ward, N. A. Williams and R. G. B. Brown. 1982. Migration
patterns of Red and Northern Phalaropes in southwest Davis Strait and in
the northern Labrador Sea. Wilson Bull. 94: 303-312.

1981

Green, J. C. 1981. First Red Phalarope for Minnesota Point. Loon 53: 52-53.

Moore, J. R., T. Fairley, D. Hamilton and P. Rodriguez. 1981. Observations of
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Pigeon, P. 1981. Observation d'un Phalarope a bec etroit Phalaropus lobatus et
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(Aisne). Alauda 49: 432.

Wander, W. 1981. Red Phalarope eating carrion. Wilson Bull. 93: 557.






Red Phalarope


1980

Chappell, M. A. 1980. Thermal energetic of chicks of Arctic breeding shore-
birds. Comp. Biochem. Physiol. A. Comp. Physiol. 65: 311-317.

Dodson, S. I. and D. L. Egger. 1980. Selective feeding of Red Phalaropes on
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Ridley, M. W. 1980. The breeding behavior and feeding ecology of Grey Phal-
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1979

Connors, P. G. and S. Gelman. 1979. Red Phalarope responds to thin oil films
in foraging experiments. (Abstract only.) Pac. Seabird Group Bull. 6: 43.

Gustafson, D. 1979. A Red Phalarope in Milwaukee. Passenger Pigeon 41: 136.

Maurizio, R. 1979. Thorshuehnchen auf dem Silsersee bei Maloja. [Grey Phal-
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Mayfield, H. F. 1979. Red Phalaropes breeding on Bathurst Island. Living
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Roudybush, T. E., C. R. Grau, M. R. Petersen, D. G. Ainley, K. V. Hirsch, A. P.
Gilman and S. M. Patten. 1979. Yolk formation in some charadriiform birds.
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1978

Brown, A. 1978. Reaction of Grey Phalarope to Sparrowhawk. Scott. Birds 10:
22.

Brown, B. and P. C. M. Latham. 1978. Grey Phalarope in the Bay of Plenty.
Notornis 25: 198-202.

Gates, L. and J. Runzo. 1978. A Red Phalarope in Hattiesburg, Mississippi.
Mississippi Kite 8: 2.

Huber, H. 1978. Red Phalarope--Mille Lacs Lake. Loon 50: 45-46.

Mayfield, H. F. 1978. Undependable breeding conditions in the Red Phalarope.
Auk 95: 590-592.

Taylor, W. B. 1978. Seasonal status, population size, and habitat utilization
of Red and Northern phalaropes in the Southern California Bight. (Abstract
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Red Phalarope


1977

Anderson, E. 1977. Red Phalarope at Moorhead. Loon 49: 173.

Savaloja, T. 1977. Minnesota's second Red Phalarope. Loon 49: 44-45.

Schamel, D. and D. Tracy. 1977. Polyandry, replacement clutches and site
tenacity in the Red Phalarope (Phalaropus fulicarius) at Barrow, Alaska.
Bird-Banding 48: 314-324.

Schiemann, H. 1977a. Uber das Vorkommen der Wassertreter (Phalaropodidae) in
den brandenburgischen sachsischen und thuringischen Bezirken, souie in
Berlin. Beitr. Vogelkd. 23: 49-56.

Sinclair, J. C. 1977. The phalaropes. Bokmakierie 29: 90-91.

Wachtler, D. and G. Wachtler. 1977. Red Phalarope at Moorhead. Loon 49:
172-173.

Watson, G. E. 1977. A second specimen of Red Phalarope from Delaware.
Delmarva Ornithol. 12: 49.

1976

Brinkschroder, W. 1976. Beobachtung eines Thorswassertreters (Phalaropus
fulicarius) im Reg.-Bez. Osnabruck. Vogelkd. Ber. Nieders. 8: 24.

Risebrough, R. W., G. E. Watson and J. P. Angle. 1976. A Red Phalarope
(Phalaropus fulicarius) in breeding plumage on Anvers Island. Antarct.
J. U.S. 11: 226.

Smith, F. T. H. 1976. An Australian record of the Grey Phalarope. Austr. Bird
Watcher 6: 292-299.

Wheeler, J. 1976. A new wader for Australia--Grey Phalarope Phalaropus fuli-
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1975

Buhler, P. 1975. Thorshuhnchen (Phalaropus fulicarius) im Prachtkleid, in Sud-
deutschland. Jahresh. Ges. Naturkd. Wuerttemb. 129: 78-80.

Heise, G. 1975. Thorwassertreter (Phalaropus fulicarius) in der Uckermark.
Beitr. Vogelkd. 21: 362-364.

Kazama, T. 1975. [A new record of Phalaropus fulicarius from Japan Sea.]
Tori 23: 29-30. [In Japanese with English summary.]

Kistchinski, A. A. 1975. Breeding biology and behaviour of the Grey Phalarope
Phalaropus fulicarius in East Siberia. Ibis 117: 285-301.





Red Phalarope


van Swelm, N. D. 1975. Foraging behaviour of Grey Phalarope. Bird Study 22:
52.

1974

Adams, R. J., Jr. and D. E. Wood. 1974. First spring record of a Red Phalarope
in Michigan. Jack-Pine Warbler 52: 44.

Boyle, G. L. 1974. Grey Phalarope feeding on earthworms. Brit. Birds 67: 352.

Brown, J. F. 1974. Grey Phalaropes (Phalaropus fulicarius) near Lagos. Bull.
Nigerian Ornithol. Soc. 10: 53.

Gillandt, L. 1974. Beobachtungen an einer Thorshuhnchen-Population (Phalaropus
fulicarius) in Sudwest-Island (Aves: Charadriiformes: Phalaropodidae). Abh.
Verh. Naturwiss. 17: 55-83. [In German.]

Lauermann, H. 1974. Ein Thorshuhnchen (Phalaropus fulicarius) am Meiseldorfer
Teich. Egretta 17: 23-27.

Marsh, C. and R. J. Hoder. 1974. Red Phalarope in Wake County, N.C. Chat 38:
71-72.

Noer, H. and B. M. Sorensen. 1974. Forekomen af stormfugle Procellariae, Thor-
shane Phalaropus fulicarius og Sabinemage, Xema sabini ved Blavandshuk
1963-71. Dan. Ornithol. Foren. Tidsskr. 68: 15-24.

Steinbach, R. 1974. Vorkommer der wassertreter Phalaropus lobatus (L.) und
Phalaropus fulicarius (L.) im Gebiet der Haselbachar und Eschefelder Teiche
sowie am Speicherbechten Windischleuba. Abh. Ber. Naturkundemus. 'Mauri-
tanium' 8: 333-338.

1973

But'ev, V. T. 1973. [On the nesting of the Grey Phalarope in the north of
Novaya Zemla Is.] Ornitologiya 10: 329-330. [In Russian.]

Fitzpatrick, J. 1973. Great Crested Grebes attacking Grey Phalarope and Storm
Petrel. Brit. Birds 66: 365.

Kistchinskii, A. A. and Y. I. Chernov. 1973. Notes on the foods of the Grey
Phalarope in the tundra of east Siberia. Moscow Society of Naturalists.
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Kocian, L. 1973. Novy vyskyt lyskonoha ploskozobeho Phalaropus fulicarius
(L., 1758) na Slovenska. [New record of Phalaropus fulicarius (L., 1758)
in Slovakia.] Zb. Slov. Nar. Muz. Prir. Vedy 19: 197. [In Czech with
German summary.]





Red Phalarope


Lok, C. M. 1973. Some observations on the Grey Phalarope. Sterna 12: 284-285.

Rettig, K. 1973. Thorswassertreter (Phalaropus fulicarius) am Rysumer Nacken
[Red Phalarope at Rysum Neck.] Ornithol. Mitt. 25: 222. [In German.]

Sannazzaro, A. 1973. II Phalaropus fulicarius in Piemonte. [Phalaropus fuli-
carius in Piedmont.] Riv. Ital. Ornitol. 43: 462-466. [In Italian.]

Schmitz, J.-P. and J. Weis. 1973. Premiere observation et capture du phalarope
a bec large (Phalaropus fulicarius) dans le Grand-Duche de Luxembourg.
Regulus 11: 3-5.

Stelzer, H. 1973. Uber das Vorkommen der Wassertreter (Phalaropodidae) in der
Schweiz und ihren Randgebieten. Ornithol. Beob. 70: 157-170. [In German
with French summary.]

van Avermaet, G. 1973. Franjepoten. Wielwaal 39: 216-220.

Webster, R. E. 1973. Short notes. Grey Phalarope. Distribution. Lammergeyer
15: 76.

1972

Portenko, L. A. 1972. Ptisy Chukotskogo poluostrova i ostrova Vrangelya.
[Birds of the Chukchii Peninsula and Wrangel Island.] Volume 1. xv and
446 pp. [1981 translation. Amerind Publ. Co., New Delhi.]

Prigge, R. 1972. Beobachtung eines Thorwassertreters (Phalaropus fulicarius)
am Treuel/Elbe im Bezirk Magdeburg. Beitr. Vogelkd. 18: 434.

Sehl, R. H. 1972. Red Phalarope in Bucks County. Cassina 53: 46-47.

Summerfield, D. 1972. Red Phalarope in December. Kentucky Warbler 48: 18.

1971

Bond, S. I. 1971. Red Phalarope mortality in southern California. Calif.
Birds 2: 97.

Danks, H. V. 1971. A note on the early season food of Arctic migrants.
Can. Field-Nat. 85: 71-72.

Fantin, G. 1971. Notizie dal Veneto: il Falaropo becco largo. [Reports
from Venice: the Grey Phalarope.] Riv. Ital. Ornitol. 41: 17-24. [In
Italian.]

Hohn, E. 0. 1971. Observations on the breeding behaviour of Grey and Red-necked
phalaropes. Ibis 113: 335-348.





Red Phalarope


King, B. 1971. Grey Phalarope on autumn passage plunging into waves. Brit.
Birds 64: 29-30.

1970

Rettig, K. 1970. Thorswassertreter bei Emden. Ornithol. Mitt. 22: 47.

1969

Hohn, E. 0. 1969. The phalarope. Sci. Am. 220: 104-111.

Kittinger, H. H. and J. N. Carusos. 1969. Second inland Red Phalarope record
in Alabama. Ala. Birdlife 17: 50-51.

Schiemann, H. 1969. Uber das Vorkommen den Wassertreter (Phalaropodidae) in
Suddeutschland. Vogelwelt 90: 184-188.

Schmidt, G. D. 1969. Polymorphus petrochenkoi sp. n. (Acanthocephala) from
the Red Phalarope, Phalaropus fulicarius L., in Alaska. J. Parasitol. 55:
335-336.

Sutton, G. M. 1969. The Red Phalarope in Oklahoma. Bull. Okla. Ornithol.
Soc. 2: 26-28.

1968

Bengtson, S.-A. 1968. Breeding behaviour of the Grey Phalarope in West Spits-
bergen. Var Fagelvarld 27: 1-13.

Ruppell, G. 1968. Phalaropus fulicarius (Phalaropodidae) Bruten. Encyclopaedia
Cinematographica. E1381. Angaben zum Film and Filmenhalt, pp. 1-6. [In
German.]

1967

Smith, W. and R. Klauke. 1967. A sight record of the Red Phalarope in Alberta.
Blue Jay 25: 25.

Spasskij, A. A. and J. N. Konovalov. 1967. Anomotaenia reticulata sp. n.
Cestoidea, Dilepididae, a parasite of Phalaropus fulicarius. Vestn. Zool.
1967: 43-48.

1966

Hohn, E. 0. 1966. Ringing (banding) and recoveries of phalaropes--a summary
of presently available information. Bird-Banding 37: 197-200.






Red Phalarope


1965

Cooch, F. G. 1965. An example of sinistralism in Red Phalaropes (Phalaropus
fulicarius). Auk 82: 276-277.

Hohn, E. 0. 1965. Die Wassertreter. Neue Brehm-Bucheri 359. Ziemsen Verlag,
Wittenberg. 60 pp.

Woodward, I. D. 1965. The status and distribution of the Grey Phalarope in
Hertfordshire. Birds Illust. 11: 150.

1964

Acton, J. R. 1964. Red Phalarope recorded from eastern Washington. Murrelet
45: 11.

Lovenskiold, H. L. 1964. Avifauna svalbardensis, with a discussion on the
geographical distribution of the birds in Spitsbergen and adjacent islands.
Skr. Nor. Polarinst. No. 129. 460 pp.

Prozesky, 0. P. M. 1964. Notes on the Grey Phalarope (Phalaropus fulicarius
(Linnaeus)) found at Voortrekkerhoogte, Pretoria, Transvaal. Ostrich 35:
70.

Shuler, J. B. 1964. Sight record of a Red Phalarope in Charleston County,
South Carolina. Chat 28: 30.

1963

Lakeman, M. 1963. Red Phalarope at Hunting Creek? Atl. Nat. 18: 250.

1962

Breiding, G. H. 1962. Red Phalarope in West Virginia. Wilson Bull. 74: 288.

1961

Alderson, G. 1961. Inland occurrences of the Red Phalarope in Oregon. Condor
63: 97-98.

Cohen, E. and J. H. Taverner. 1961. Colouration of soft parts of Grey Phal-
aropes. Brit. Birds 54: 116-117.

1960

Baldwin, W. P. 1960. Red Phalarope at Cape Romain, S.C. Chat 24: 76.

Cowcill, L. and A. Smith. 1960. Grey Phalarope blowing bubbles. Brit. Birds
53: 574.

Milne, B. S. 1960. Huge flocks of Grey Phalaropes in the Isles of Scilly.
Brit. Birds 53: 403.

82