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birdmigrationandglobalchange

BirdMigrationand GlobalChange

Copyright©2010GeorgeW.Cox

AllrightsreservedunderInternationalandPan-AmericanCopyrightConventions. Nopartofthisbookmaybereproducedinanyformorbyanymeanswithoutpermission inwritingfromthepublisher:IslandPress,Suite300,1718ConnecticutAvenueNW, Washington,DC20009.

ISLANDPRESSisatrademarkoftheCenterforResourceEconomics.

DesignandtypesettingbyKarenWenk PrintedusingGalliard

LibraryofCongressCataloging-in-PublicationData

Cox,GeorgeW.,1935–Birdmigrationandglobalchange/GeorgeW.Cox.—1sted. p.cm.

Includesbibliographicalreferencesandindex. ISBN-13:978-1-59726-687-1(cloth:alk.paper)

ISBN-10:1-59726-687-6(cloth:alk.paper)

ISBN-13:978-1-59726-688-8(pbk.:alk.paper)

ISBN-10:1-59726-688-4(pbk.:alk.paper)1.Migratorybirds—Adaptation. 2.Migratorybirds—Effectofhabitatmodificationon.3.Climaticchanges.4. Globalwarming.I.Title. QL698.9.C692010 598.156′8—dc22

2010004093

Printedonrecycled,acid-freepaper

ManufacturedintheUnitedStatesofAmerica 10987654321

Keywords:bird,migration,globalchange,climatechange,globalwarming,adaptation, evolutionarychange

Willthegreatmigrationsofbirdssurvive?Manybiologistsbelievethatmigratorybirdsareatgreateraverageriskofextinctionduetochangingclimatethanareresidentspecies.Thisideaisbasedonthefactthatmigratory birdsdependondifferenthabitatsandresourcesindifferentlocationsduringtheirannualcycleandonthefactthatthefailureofanyofthesecouldbe fatal.Asecondgeneralbeliefisthattheclimateisnowchangingatarate fasterthanbirdshaveexperiencedinthepastandthatevolutionaryadjustmentsbybirdsmaynotbeabletokeeppacewiththischange.

Thethesisofthisbookisthatmanymigratorybirdsexhibitahighdegreeofecologicalandevolutionaryadaptabilityandthatmanyarenow showingrapidadjustmenttoclimaticchanges.Asubsidiarythesis,however,isthatforotherspecies,climatechangecanseverelyconstrainorpreventfullecologicalandevolutionaryadjustment,puttingtheirsurvivalat risk.Theobjectiveofthisbookisthereforetoevaluatethecapacityofmigratorybirdstorespondtothechallengesofchangingclimate.

Thebookisareview,synthesis,andinterpretationofrecentscientific literatureonmigratorybirdsandtheirresponsestochangingclimate.We beginbycharacterizingthepatternsofmigrationshownbybirdsofdifferentecologicalandtaxonomicgroups,andthefrequencyofmigrationindifferentworldenvironments,frompolarregionstothetropics.Thenatureof climatechangethroughouttheworldisexamined,andtheimpactsofthis changeonbirdhabitatsareevaluated.Thespecialthreatsofchangetoparticularhabitattypesandresourcesutilizedbymigratorybirdsareexamined andtheconstraintsonresponsesbybirdsconsidered.Theparticularpatternsofinfluenceofclimatechangeonmigratorylandbirdsareconsidered fordifferentbiogeographicregionsandforspecialbirdgroups,including raptors,freshwaterbirds,andmarinebirds.Finally,thecapacityforecologicalandevolutionaryadjustmentisexamined,andtheadequacyofcurrent conservationprogramsisevaluated.Notreatmentofbirdmigrationhas pulledtogetherthesediversetopics,whichareofcriticalimportancetohumaneffortstoprotectbiodiversity.

Mostofthetopicsinthebookhavebeenaddressedforspecificgroups

ofbirdsorforbirdsinspecificgeographicalareas.Sometopics,suchasthe impactsofclimatechangeonmigratorytropicalbirds(Chapter10)and evolutionaryadaptabilityofwaterbirds(Chapter18),havereceivedonly spottyattentionotherthanthatdirectedatafewspecies.Othertopicsfor whichnorecentsyntheseshavebeenattemptedincludeworldwidealterationofmigratorybirdhabitats(Chapter3),migratorylandbirdsofthe temperateSouthernHemisphere(Chapter9),shorebirds(Chapter12), waterfowl(Chapter13),oceanicbirdsoftheNorthPacificandSouthern Hemisphere(Chapters15and16),evolutionaryabilityoflandbirds (Chapter17),andoverallcapacityforecologicalandevolutionaryadjustment(Chapter19).RecentsymposiahavetendedtofocusonlandbirdresponsestochangingclimateinNorthAmericaandEurope,butinterestis increasingrapidlyinAustralia.Wereallyknowlittleabouthowmigratory landbirdsofSouthAmerica,easternAsia,andAfricaarerespondingtoclimaticchange,butallindicationsarethatmajorchangesareoccurring.

Birdinghasbecomenotonlyahugelypopularoutdooractivityformillionsofpeoplebutalsoanactivitythatbringspeoplefacetofacewithbiodiversityandthethreatstoitssurvival.Ihavenotmetabirderwhoisnot concernedaboutprotectionofbirdhabitatsandabouttheimpactsof changingclimatesonbirds,whetherresidentormigratory.Ihopethatthis accountincreasesawarenessofthechangesthatmigratorybirdsareexperiencingandstimulatesreaderstoparticipateineffortstokeepthegreatmigrationsinmotion.Seetheappendix,aswellaswww.birdmigration.org, foralistofthecommonandscientificnamesofspeciesdiscussedinthe text.

Iamindebtedtomanyfriendsandcolleagueswhohavereviewedportionsofthebook,especiallyBenBecker,KeithBildstein,ChristianBoth, RichardBrewer,LyndaChambers,GlennConroy,HughDingle,George Divoky,PeterDunn,FredGehlbach,FrankGill,PhilHockey,SteveOberbauer,JanePhillips-Conroy,RobertRicklefs,KurtRiitters,TimSparks, NilsStenseth,NilsWarnock,andS.JosephWright.Manyothersgraciously tooktimetohelpmefindliteratureforparticularregionsortopics:Tatsuya Amano,KenChan,TimCoppack,VictorCueto,TomEbert,KenGreen, LouisHansen,StuartHurlbert,AlexJahn,LeoJoseph,BrianLinkhart, J.AlanPounds,MichaelScott,KimScribner,RobSimmons,Liliana Spescha,WayneTrivelpiece,RickWessels,EdWillis,andBenZuckerberg. IespeciallythankDarlaCox,mywife,forsharingmanybirdingexperiences andforreviewingmanyofthechaptersofthisbook.

GeorgeW.Cox SantaFe,NewMexico

PARTI Introduction

Globalclimatechangeisnowaffectingallmajorgroupsoforganismsand changingthedynamicsofecosystemsfromthetropicstopolarregions. Birdmigration,oneofthemostcomplexandintriguingpatternsofadaptationtoclimatesthatshowseasonalchanges,iscertaintobeaffectedby globalchange.Inthissectionweshallexamineworldwidepatternsofbird migrationandidentifythepotentialpatternsofresponsebymigratorybirds tothischallenge.

Chapter1 BirdMigrationandGlobalChange:

TheBirdsandtheIssues

Birdsarethemostmobileofvertebrates.Whethertheyfly,swim,orrun, theirabilitytocovergreatdistancesquicklyenablesmanyspeciestoexploit differentareasatdifferentseasonsintheannualcycleandatdifferentstages intheirlifehistory.Thisismostevidentinthosethatfly,butflightlesspenguinsroamthesouthernoceansfarfromtheislandsandcoaststhatsupporttheirbreedingcolonies.Evenafewflightlesslandbirdshavewelldevelopedmigrations.Emus1 inaridWesternAustraliamovehundredsof milesinresponsetorainfallandvegetationconditions.Intheforestsof NewGuinea,DwarfCassowariestrackseasonalfruitproductionwithaltitudinalmovementsofathousandormoremeters.

Changingclimateshavespecialimplicationsformigratoryspecies.Migratorybirdsdependonspecifichabitatsandresourcesindifferentgeographicalareasatdifferentphasesoftheirannualcycle.Howarethesecomplexpatternsofhabitatandresourceusebeingaffectedbyglobalclimatic change?Istheecologicalandevolutionaryadaptabilityofmigratorybirds adequatetokeeppacewithchangingclimatesandlandscapes?Arefuture changesinclimatelikelytocauseextinctionofmanyoftheworld’smostremarkablespecies?Thesearebasicquestionsthatweshalladdress.

1ScientificnamesforspeciesmentionedinthetextaregiveninAppendix1.

MigratoryBirds:EcologicalPatterns

Migratorybirdsoccupyalloftheworld’smajorenvironments,andtheir movementsareextraordinarilydiverse.Land,freshwater,andmarinebirds showwell-developedmigrations,asdobirdsofclimaticzonesranging frompolarregionstothetropics.

Forlandandfreshwaterbirds,migrantsvaryenormouslyinthedistancetheytravel,intheregularityoftheirschedulesfromyeartoyear,and inthefractionofindividualsthataremigratoryindifferentpartsoftheir ranges.Short-distancemigrantsrangefromthosethatshowlocalhabitat shiftsoraltitudinalmovementswithinasmallgeographicalareatothose thatmakeintracontinentalmovementsofuptoafewhundredkilometers. WhereIliveinnorthernNewMexico,forexample,thewinterDark-eyed JuncosatourfeederareamixoflatitudinalmigrantsfromthePacific NorthwestandnorthernRockyMountainsandgray-headedbirdsthatare altitudinalmigrantsfromnearbymountains.Weatherconditionsofthe particularyearinfluencemanyoftheseshort-distancepatterns.Frequently, onlyaportionofthepopulationofaspeciesinagivenlocalityshowsshortdistancemovements,apatterntermedpartialmigration.Overthegeographicalrangeofmanyspecies,localpopulationsrangefrombeingcompletelyresidenttopartiallyorfullymigratory.

Birdsofmanydifferenttaxaandspeciesatalllatitudesexhibitshortdistancemovements.Birdsofmountainousregions,fromhighlatitudesto thetropics,showaltitudinalshifts,withmanysuchmovementsbytropical speciesyettobedocumented.ManybirdsofsemiaridregionsinAfrica, Asia,andAustralia,characterizedas“nomadic,”showopportunisticmovementpatternsthatenablethemtoutilizeareasthathaverecentlyexperiencedfavorableweatherconditions.

Long-distancemigrationsoflandandfreshwaterbirdsinvolveflights acrossmajorclimaticzones,andoftenbetweencontinentsorhemispheres. Thesemovementsareoftenmoreregularintimingthanthoseofshortdistancemigrants.Strong-flyingraptorsandshorebirdsperformsomeof thelongestmigrations.Radiotrackinghasshown,forexample,thatBartailedGodwitsflynonstopfordistancesof10,000kilometersormorebetweenbreedingareasinAlaskaandwinteringareasinNewZealandand Australia.TheSwainson’sHawksthatoccasionallyflyovermyhomein NewMexicoinspringspendthewinterontheArgentineanpampas.Many smallsongbirds,however,makeintercontinentalmigratoryflightsofthousandsofkilometers.

Seabirdsshowdiversepatternsofmigration,aswell.Someareshortdistancemigrantsthatdispersefromcoastalorinsularbreedingcoloniesto

neighboringoceanicregionsuptoonlyafewhundredkilometersdistant. Others,suchastheArcticTern,arelong-distancemigrantsthatflythousandsofkilometersfrombreedingareastononbreedingrangesindistant oceans,sometimesintheoppositehemisphere.

MigratoryBirds:TaxonomicPatterns

About9930speciesofbirdsexistworldwide,belongingto204families. Frommanysourcesintheliterature,Ihavecompiledapreliminarylistof migratorybirds,atasknotaseasyasitmightatfirstseem.Manyspecies thatarecommonlyregardedaspermanentresidentsarereallypartialmigrantsinsomepartsoftheirranges.InnorthernNewMexico,forexample, theSteller’sJay,consideredbymostreferencebookstobeapermanentresident,isapartialmigrant,withmanybirdsmovingtolowerelevationsin winter.Ourunderstandingoftheseasonalmovementsoftropicalspecies, especiallyaltitudinalmovementsofthoseofmountainousregions,isstill veryrudimentary.Mysurveyofmigratoryspeciesindicatesthatatleast 2600birdspeciesof141familiesshowsometypeofseasonalmigrationor substantialnomadism(Table1.1).Thiscorrespondstoabout26.2percent ofallbirdspecies,afigurethatissuretoincreaseaswelearnmoreabout birdmovementsinregionssuchaseasternAsia,muchofAfrica,andmountainareasthroughouttheworld.Myestimatealsosubstantiallyexceedsthe estimateof19percentofmigratoryspeciespresentedbyBirdLifeInternationalin2008.

Thefrequencyofmigrationvarieswidelyamongdifferentgroupsof birds.Membersofafewfamiliesoftropicalbirds,suchasthefamilyto whichtheantthrushesandantpittasbelong(Formicariidae),arealmost completelynonmigratory.Others,suchastheNewWorldandOldWorld warblerfamilies(ParulidaeandSylviidae,respectively),containsomespeciesthatarepermanentresidents,otherspartialmigrants,andstillothers long-distancetemperate–tropicalmigrants.Inmostfamiliesoftemperate zonesongbirds,suchasthetitmiceandchickadees(Paridae),somespecies areatleastpartialmigrants,thatis,withsomepopulationsthatconsist partlyorlargelyofmigratoryindividuals.Almostallseabirdsaremigratory,inthesenseofspendingnonbreedingperiodsatseaatfeedingareas substantiallydistantfromtheirnestingareas.Waterfowlthatbreedinthe temperatezonesortheArcticarealmostallmigratorytovaryingdegrees. Shorebirdsbreedingathighlatitudesarenearlyalllong-distancemigrants. FalconsandhawksthatbreedinthetemperatezonesortheArcticarealso mostlymigratory—someonlyshort-distancemigrantsbutothersshowing

Table1.1. Migratorybirdspeciessummarizedbyorder.

TotalMigratory

Orders FamiliesSpeciesSpecies

Tinamous(Tinamiformes) 1470

Rheas(Rheiformes) 120

Ostrich(Struthioniformes) 111

EmuandCassowaries(Casuariiformes) 242

Kiwis(Dinornithiformes) 150

Loons(Gaviiformes) 155

Grebes(Podicipediformes) 11915

Penguins(Sphenisciformes) 11715

PetrelsandAlbatrosses(Procellariiformes) 4114114

PelicansandRelatives(Pelecaniformes) 46748

DucksandGeese(Anseriformes) 2162124

Flamingos(Phoenicopteriformes) 166

WadingBirds(Ciconiiformes) 511754

Raptors(Falconiformes) 5313108

QuailandRelatives(Galliformes) 628413

CranesandRelatives(Gruiformes) 1220774

Shorebirds(Charadriiformes) 19356274

Sandgrouse(Pterocliformes) 11611

PigeonsandRelatives(Columbiformes) 130827

ParrotsandRelatives(Psittaciformes) 336837

Mousebirds(Coliiformes) 160

CuckoosandTuracos(Cuculiformes) 316450

Owls(Strigiformes) 221518

NightjarsandRelatives(Caprimulgiformes)512035

SwiftsandHummingbirds(Apodiformes) 344389

Trogons(Trogoniformes) 1405

MotmotsandRelatives(Coraciiformes) 1021948

Woodpeckers(Piciformes) 641227

PerchingBirds(Passeriformes) 9858931400

Total 20499302600

intercontinentalmovements.Owlsofthesesamelatitudes,ontheother hand,areinsomecasesmigrants,inothercasespermanentresidents.

MigratoryBirds:GeographicalPatterns

Therelativeabundanceofmigratoryspeciesalsodiffersmarkedlyamong majorgeographicalareas(Table1.2).ThelargelandmassesoftheNorth

Table1.2. Livingresidentandmigratorynonoceanicbirdspeciesinvariousworldregions.

PrincipalSources: Clements,J.F.2007. TheClementsChecklistofBirdsoftheWorld ,6thed.ComstockPublishingAssociates(CornellUniversityPress),Ithaca,NY.

AOUchecklistofNorthAmericanbirds.http://www.aou.org/checklist/north/index.php. Avibase.Birdchecklistsoftheworld.http://avibase.bsc-eoc.org/checklist.jsp?lang=EN. OrientialBirdClub.http://www.orientalbirdclub.org/publications/checklist/obcchecklist.txt. TheBirdsofAfrica .Vol.1–7.AcademicPress,NewYork.

TemperateandArcticzonescontainmanymigrantspecies,mostofwhich breedintheregion.Intropicalandsubtropicalregionsofthecontinents, winteringmigrantsarenumerous.Tropicalislandarchipelagossuchasthe EastandWestIndiesarealsowinteringareasformanymigrants,asare smallerislandareasthroughouttheAtlantic,Pacific,andIndianoceans. Manyofthemigrantstodistantoceanicareasarenonbreedingshorebirds andseabirds.SouthTemperateZoneregionsarealsohometomanybreedingmigrantspecies,andthebirdsthatbreedintheAntarcticarealmostall migratory.IntratropicalmigrationpatternsarewelldevelopedinAfrica, andtoalesserextentinAustraliaandtheNewWorld.Manybirdsofmountainousregions,regardlessoflatitude,showaltitudinalmovements.

Severalmajormigrationsystemscanberecognizedindifferentworld regions.IntheNewWorld,theseincludetheNearctic,Nearctic–Neotropical,Neotropical,andSouthAmericanmigrationsystems.The Nearcticmigrationsystemcompriseslandandfreshwaterbirdsthatbreed inCanadaandtheUnitedStatesandwinterprimarilynorthofcentralMexico.Thismigrationcomplexinvolvesabout423species,includingmany speciesofwaterfowlandotherfreshwaterbirds,hawksandowls,andsmall landbirds.About13oftheNearcticspeciesthatbreedathighlatitudesmigratetoAsiaratherthantomore-southernpartsofNorthAmerica.

TheNearctic–Neotropicalmigrationsystemincludesbirdspeciesthat breedinNorthAmericaandwinterinsouthernMexico,CentralAmerica, theWestIndies,andSouthAmerica.About184speciesofland,freshwater, andcoastalmarinebirdsareinvolved.About76ofthesespeciesspendthe nonbreedingseasoninMexico,theWestIndies,orCentralAmerica,but 108speciesextendtheirwinterrangesintoSouthAmerica.TheNeotropicalmigrationsystemcomprisesabout104species,mostofwhicharealtitudinalmigrants.About5speciesbreedintheNorthAmericantropicsand winterinSouthAmerica.WeknowmoreaboutpatternsofaltitudinalmigrationinMexicoandCentralAmericathananywhereelse,andwhatwe seeheresuggeststhatwehavemuchtolearnaboutaltitudinalmigrationin mountainousregionselsewhereintheworld.

TheSouthAmericanmigrationsystemcomprisesabout266land, freshwater,andcoastalmarinebirds.Mostofthesespeciesbreedinthe temperateregionofsouthernSouthAmericaandmigratenorthinthe australwinter,butatleast31speciesshowshort-distanceoraltitudinal movementsinthetropics.About77ofthespeciesbreedingintemperate SouthAmericaarefullymigratory.Some44species,mostlytyrannidflycatchersandswallows,winterinthehumidtropicalregioncenteredonthe Amazonbasin.Formorethantwo-thirdsofthesespecies,rangesofnonbreedingmigrantsoverlaprangesofresidentsofthesamespecies.

TheOldWorldexhibitsEuropean,European–African,andAsianmigrationsystems.InEurope,about116of524breedingspeciesofland, freshwater,andcoastalmarinebirdsareshort-distancemigrantsthatwinter withintheregion.TheEuropean–Africanmigrationsystemincludes185 speciesofland,freshwater,andshorebirdsofthirty-twofamiliesthatwinter inAfrica.Thesespeciesareprimarilyhawksandfalcons,waterfowl,shorebirds,OldWorldwarblers,thrushes,swallows,pipits,wagtails,andshrikes. Ofthese,62species,particularlymanyhawksandfalcons,shorebirds,Old Worldwarblers,thrushes,andshrikes,winterexclusivelyinsub-Saharan Africa.Oftheremainingspecies,somewinterinAfricaandsomeinareasof theMiddleEast.

TherichAsianmigrationsysteminvolvesbirdsthatmovefrombreedingareasineasternEurasiatowinteringareasinsouthernandsoutheastern Asia,thePhilippines,andtheEastIndies.About827speciesofforty-four familiesshowmigratorymovementswithinthisregion.About35species ofsandpipersandplovers,1tern,and10speciesoflandbirdscontinuefarthersouthtowinterintheAustralo–Papuanregion.InRussia,China,and otherpartsofeasternAsia,atleast66speciesofthrushes,flycatchers,Old Worldwarblers,finches,andotherpasserinesareshort-distanceoraltitudinalmigrantsthatdonotreachthetropics.

Africaexhibitstwowell-definedmigrationsystems.Thesouthern Africanmigrationsysteminvolves134fullyorpartiallymigratorylandand freshwaterbirdsthatbreedinsouthernAfricaandwinterfarthernorth.Waterfowl,rails,herons,kingfishers,cuckoos,swifts,swallows,andavariety ofpasserinebirdsareprominentinthissystem.ManybirdsbreedinginEuropeandAsiaappearinsouthernAfricaduringtheNorthernHemisphere winter.Theseincludemanyspeciesofsandpipersandplovers,gullsand terns,hawks,andOldWorldwarblers.

TheintratropicalAfricanmigrationsystemiscenteredontheequatorialregionandthesemiaridbeltstothenorthandsouth.About277species,rangingfromheronsandploverstolarksandfinches,moveprimarily northandsouthinthisregion,trackingfavorableconditionsrelatedtowet anddryseasonweatherpatterns.Somecuckoosbreedinthesemiaridzones bothnorthandsouthoftheequator,switchingplacesthroughmigration whennotbreeding.Inmore-aridregions,movementsofsomespeciesare bestdescribedasnomadic.AltitudinalmigrantsarefrequentinthemountainsofeasternAfrica.

TheAustralo–Papuanregion,comprisingAustraliaandNewGuinea, holdsalargelyself-containedmigrationsystem.Thismigrationsystemconsistsofabout272landandfreshwaterbirds.Somearemigrantsbetween AustraliaandNewGuinea,othersmovebetweenTasmaniaandmainland

Australia,andstillothersshowlatitudinalandaltitudinalmovements withinmainlandAustralia.Inaddition,manylandbirdsandwaterbirds performirregularmovementswithininteriormainlandAustralia.Some 169speciesshowpartialmigrationpatternsinAustraliaandTasmania. AmongTasmanianbirds,only20speciesaremigratory,andonly4arefully migratory,completelyleavingtheislandfortheAustralianmainland.

NorthernHemispheremigrantsalsoreachmanyPacificislands,includingNewZealand,althoughfewspeciesareinvolved.Long-distance migrantssuchasploversandsandpipersmostfrequentlyvisittheseisolated regions.

Seabirdmigrationsystemsrelatelesstolatitudinaltemperaturepatternsandmoretothelocationsofsuitablenestingareasandproductive oceanwaters.About234speciesofbirdsofeighteenfamiliesarelargelyor entirelypelagicintheirdistributionsinthenonbreedingseason(Table1.3). Onlyverygeneralmigrationpatternsareapparent.IntheeasternNorthAtlantic,about31speciesofloons,petrels,shearwaters,gannets,cormorants, seaducks,skuas,gulls,andalcidsbreedinArcticandsubarcticareasand winterinpelagicoroffshorecoastalareas.InthewesternAtlantic,about28 ofthespeciesofthesegroupsbreedinnorthernareasandwinterinwaters offthecoastsofCanadaandtheUnitedStates.TheNorthPacifichasa richerfauna.About53speciesoftheseseabirdswintersouthwardthrough theAleutianIslandsandalongtheeasternandwesterncoastsofthePacific. Tropicalandsubtropicaloceanshosttherichestandmostdiversefauna ofpelagicbirds,includingnumerouspetrels,shearwaters,terns,boobies, tropicbirds,cormorants,andfrigatebirds.

InthesouthernoceanssurroundingAntarctica,atleast94speciesof penguins,albatrosses,petrels,cormorants,skuas,andrelatedbirdsbreed andforagetoconsiderabledistancesfrombreedinglocalities.Intropical andwarmtemperateoceanareas,atleast97speciesofseabirdsofeleven familiesbreedandwanderwidelyoveroceanareasdistantfromtheirnestingislandsorcoastalrookeries.

TheChallengeofClimaticChange

Migratorybirdsfacemajorchallengesofsurvivalinthefaceofrapid,human-inducedglobalchange.Overthepastcentury,theearth’sclimatehas warmedby0.8°C,andbyAD2100,warmingwilllikelybebetween2.0°C and4.5°C.Becausetheydependonhabitatsandresourcesindifferentareas atdifferentstagesoftheannualcycle,populationsofseasonalmigrantsin

Table1.3. Worlddistributionofmigratoryoceanicbirds. 1

InterregionalMigrants

IntraregionalMigrants

Table1.3. Continued

InterregionalMigrants

IntraregionalMigrants TemperateWintertoWintertoTemperateWinterto

1 ZonalseparationsroughlycorrespondtoArcticandAntarcticcirclesandTropicsofCancerandCapricorn.Somespeciesarepartitionedbetweentwozones,giving risetothe0.5values.

everyworldregionwillbeaffectedbyclimaticchange.Theareasthatmigrantsuseseasonallyincludetheirbreedingranges,stagingandstopover locationsduringmigration,andareasoccupiedduringthenonbreedingperiod.Inonesense,becauseoftheirspecializationforuseofdifferenthabitatsorgeographicalareasatdifferenttimes,migratoryspeciesmightseem tobeatgreaterriskofextinctionthanpermanentresidents.Achangeinany oneoftheareasusedduringtheannualcyclemightcausetheirevolutionary strategytofail.Someornithologistsconcludethatsuchdependenceon multiplegeographicalareasplacesmigratoryspeciesatgreaterriskthanresidentspeciesinthefaceofglobalclimatechange.

Ontheotherhand,migratorycapabilityisanextensionofbasicphysiologicalandbehavioraladaptationsforlocalmovements,homing,andthe annualreproductivecycle,anditmustpossessadegreeofflexibility.Climaticchangesoverlonggeologicaltimehavetestedtheabilityofmigrants toadjustbreedingandnonbreedingrangesandaltermigrationroutes.Perhapsthisadaptabilityisadequatetorespondtotherapidenvironmental changesnowoccurring.Or,ofcourse,somemigratorybirdsmaybeableto adjustquickly,whileothersmaynot.

Severalfactorscouldconstraintheabilityofmigratorybirdstorespond tochangingclimate.Lackofgeneticvariabilityorphenotypicplasticity mightlimitthecapacityofspeciestorespondtochangingconditions.Limiteddispersalabilitymightalsoslowtheabilityofthespeciestoshifttheir geographicalrangesinresponsetochangingconditions.Anincreaseinfrequencyofextremeweatherevents,overalllossordegradationofoneor moreofthehabitatsonwhichtheydependthroughouttheannualcycle, andchangedinfluencesofcompetitorspecies,predators,ordiseasesmay alsoimpactmigratoryspeciesnegatively.

Isuspectthatpopulationlimitationofmigratoryspeciesisadynamic relationshipinvolvingbothbreedingandnonbreedingareas.Ifconditions inthewinteringrangefavorincreasedsurvival,expansionofthebreeding rangeintoregionswithlowerreproductivesuccesswilltendtooccur.If conditionsinthebreedingrangefavorincreasedreproductivesuccess,expansionofthenonbreedingrangeintoregionswithlowersurvivalwill likelyresult.Inasmuchasbreedingsuccessandnonbreedingsurvivalvary fromyeartoyear,theoutcomeofthisrelationshipchangesthroughtime. Bothhumanmodificationofthelandscapeandglobalclimaticchangeobviouslyinfluencethisprocess,aswell.

Issuesrelatingtomigratorybirdsattractenormousscientificconcern. TheSmithsonianInstitutionsponsoredsymposiaonmigratorybirdsinthe NewWorldin1977and1989.In2002,anotherSmithsoniansymposium,

entitledBirdsofTwoWorlds:AdvancesintheEcologyandEvolutionof Temperate–TropicalMigrationSystems,consideredmigrationpatterns worldwide.Thefirstorganizedefforttofocusdirectlyonclimateissueswas asymposium,BirdMigrationinRelationtoGlobalChange,heldatthe UniversityofConstance,Germany,inMarch2003.Mostrecently,acongressonBirdMigrationandGlobalChangetookplaceatAlgeciras,Spain, inMarch2007.

Since2006,theUnitedNationsEnvironmentProgrammehassupportedWorldMigratoryBirdDay(WMBD),aglobalinitiativefocusedon migratorybirdsandtheirconservation.ThesecretariatoftheAfrican–EurasianMigratoryWaterbirdAgreementinitiatedWMBD,whichnowinvolvesseveralothercooperatingorganizations.Actuallya2-dayeventon thesecondweekendinMay,eachannualprogramemphasizesaparticular issuerelatingtomigratorybirds.In2009WMBDfocusedonman-made structuresthatposedangerstomigratingbirds.

PatternsofResponsetoClimateChange

Changingclimatesarealteringmigratorybehaviorinseveralways.These include(1)changeintherelativenumbersofmigrantandresidentindividualsinlocalpopulations,(2)changeinthedistancebetweenareasoccupied atdifferentstagesintheannualcycle,(3)changeinthedirectionofmigratorymovements,and(4)changeintimingandspeedofmigratorymovements.Someoftheseresponsesappeartobesimplephenotypicresponses ofindividualstoalteredenvironmentalconditions,whereasothersare genotypicormicroevolutionaryresponses.

Changesintheproportionofmigrantsandresidentscanoccurifamelioratedclimaticconditionsduringthenonbreedingseasoninaparticularlocationfavormoreindividuals’becomingpermanentresidents.Or,if weatherconditionsandresourceavailabilitypatternsbecomemorestrongly seasonal,anincreaseinthemigratoryfractionofthepopulationcanoccur. Sincepatternsofglobalclimaticchangevaryfromregiontoregion,bothof thesechangesareexpected.

Withclimaticwarming,favorablehabitatsforbreedingbymanyspeciesmayappearathigherlatitudes.Thismayleadtolongermigrationsby individualscolonizingsuchareas,iftheareassuitableduringthenonbreedingseasonremainthesame.Or,asinthecaseofleapfrogmigration patterns,individualsbreedingfartherpolewardmaytendtomovetononbreedingareasevenfartherfromtheirbreedingareas.Lengthenedmigra-

tionscouldbeaseverechallengeifstopoversitesalsobecamelessfrequent orlessfavorable.Ontheotherhand,ameliorationofclimateinnonbreedingareasmightallownonbreedingbirdstoremainclosertotheirbreeding ranges.

Sinceclimaticchangesareunevenacrosslargecontinentalmasses, changesinthedirectionofmigratorymovementsarelikelytooccur.Regionssuitableforoccupationduringbreedingornonbreedingperiods mightappearathigherlatitudesbutatlongitudesdifferentfrompreviously suitableregions.Likewise,previouslyfavorablebreedingornonbreeding areasmightshiftinlongitude.

Withclimaticchange,theseasonsfavorableforoccupationofthe breedingrangeoroccupationofthenonbreedingareasmaylengthenor shorten,leadingtoaneedforchangesinthemigrationschedule.Opposing selectionpressuresatdifferentseasonsmaysloworpreventadjustmentsin timingofmigration,andlifehistoryfeaturesmayfalloutofsynchronywith seasonalprogression.Especiallyforlong-distancemigrants,suchasEurasianbirdswinteringinsub-SaharanAfricaorNearcticmigrantswintering inthetropics,correlationsbetweenhabitatconditionsonbreedingand nonbreedingrangesmaybeveryweakorevenlacking.Endogenousfactors andphotoperiodicresponsesthatarenotdirectlytiedtotemperatureare whatcontroltheannualcyclesofmostofthesespecies.Asaresult,many birdsmayfailtoadjustmigrationschedulestotakeadvantageofoptimal conditionsforbreeding.

Acentralissueofclimaticchangeforallmigratoryspeciesistheirdegreeofvulnerabilitytoextinction.Migratorybirdshavedealtsuccessfully withmajorclimaticshiftsduringperiodsofPleistoceneglaciation,althoughsomeextinctionsalmostcertainlydidoccur.Now,however,climate ischangingmorerapidlythanduringthePleistocene.Theextensivetransformationandfragmentationofhabitatsbyhumanactivitiesmayconstrain theabilityofmanyspeciestorespondtogeographicalshiftsintheiroptimal habitat.Forbirdsandotherlandanimalsthatoccupyhabitatsdefinedby plantcommunitystructure,theabilityofplantspeciestomoveinresponse tochangingclimatemaybecritical.Inaddition,eventhoughbirdsare highlymobile,strongsitefidelityandthemechanismbywhichsitefidelity isestablishedinjuvenilesmightalsolimitthecapacityofmigratorybirdsto occupynewregionsofsuitablehabitat.

Short-andlong-distancemigrantsmightalsobedifferentiallyvulnerabletoextinctioninthefaceofchangingclimates.Short-distancemigrants tendtoresponddirectlytoimmediateweatherconditionswithinthelocal environment.Inasmuchastheweatherconditionsontheirnonbreeding

rangesarelikelytobecorrelatedwiththoseontheirnearbybreeding ranges,appropriatechangesinmigratoryschedulesarequitelikely.Longdistancemigrantsmustrespondtoenvironmentalcuesthatarecorrelated withchangesontheirdistantbreedingornonbreedingareasandalong theirmigrationroutes.Howquicklytheycanrespondisuncertain.

Inthefollowingchaptersweshallfirstexplorethenatureoftheclimaticandenvironmentalchangesthatareoccurring.Weshallthenexamine responsesbybirdsofvariousworldregionsandecologicalgroups.Finally, weshallevaluatethecapacityofmigratorybirdstoadapttothechangesin climatethatareforeseen.

Summary

Migratorybirdsthroughouttheworldarefacingclimaticchange.Over26 percentofallbirdspeciesshowsomepatternofmigration.Majorsystems ofmigrationbylandandfreshwaterbirdsexistinNorthAmerica,Eurasia, andthecontinentsoftheSouthernHemisphere.IntheNorthernHemispheretheseincludepatternsofshort-distance,intracontinentalmovement andlong-distancetemperate–tropicalorintercontinentalmovements. Seabirdsatalllatitudesshowlatitudinalmovementsbetweenbreedingand nonbreedingareasornonbreedingdispersaltodistantoceanareas.Thecapacityofthesediversemigrantstoadjusttochangingclimatesandfood supplieswilldeterminewhetherornotmigrationwillremainamajorphenomenonofavianecology.

keyreferences

Alerstam,T.,A.Hedenstron,andSusanneÄkesson.2003.“Long-distancemigration:Evolutionanddeterminants.” Oikos 103:247–260.

Chesser,R.T.1994.“MigrationinSouthAmerica:Anoverviewoftheaustralsystem.” BirdConservationInternational 4:91–107. Cox,G.W.1985.“Theevolutionofavianmigrationsystemsbetweentemperate andtropicalregionsoftheNewWorld.” AmericanNaturalist 126:451–474. Dingle,H.2004.“TheAustralo–Papuanbirdmigrationsystem:AnotherconsequenceofWallace’sline.” Emu 104:95–108. Greenberg,R.andP.Marra(Eds.).2005. BirdsofTwoWorlds:TheEcologyandEvolutionofMigration.JohnsHopkinsUniversityPress,Baltimore. Hockey,P.A.R.2000.“Patternsandcorrelatesofbirdmigrationinsub-Saharan Africa.” Emu 100:401–417.

Huntley,B.,Y.C.Collingham,R.E.Green,G.M.Hilton,C.Rahbek,andS.G. Willis.2006.“Potentialimpactsofclimaticchangeupongeographicaldistributionsofbirds.” Ibis 148:8–28.

Joseph,L.1996.“PreliminaryclimaticoverviewofmigrationpatternsinSouth Americanaustralmigrantpasserines.” Ecotropica 2:185–193.

Moreau,R.E.1972. ThePalearctic–AfricanBirdMigrationSystem.AcademicPress, NewYork.

Rappole,J.H.1995. TheEcologyofMigrantBirds:ANeotropicalPerspective.SmithsonianInstitutionPress,Washington,DC.

PARTII

TheChangingEnvironment

Humanactivitiesaremodifyingthechemistryandphysicsoftheearth’satmosphere,aswellascharacteristicsofthelandsurface,freshwaterlakesand streams,andtheoceans.Thesechangesarealteringtheearth’senergybalance,itsbasicpatternsofatmosphericandoceancirculation,andclimatic conditionsofalloftheearth’secosystems.Inturn,theseeffectsarealtering thehabitatsandresourcesthatarecriticalformigratorybirds.Directhumanimpactsontheearth’secosystemsareinteractingwiththeserelationships,creatingacomplexpatternofglobalenvironmentalchange.

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This name is said to be applied to the Bella Coola by the Tsimshian. Perhaps the inlet so called included Milbank sound, Seaforth channel, and Dean inlet. ↑

Perhaps this name means “floating.” ↑

The word seems to mean “precious” or “valuable.” ↑

The landing party found itself unable to withdraw without great danger. ↑

See the story of “Sacred-one-standing-and-moving, Stone-ribs, and Upward,” note 23. ↑ [Contents]

RAID BY THE N

INSTINTS

HAIDA ON THOSE OF THE WEST COAST

[Told by Richard of the Middle-gîtî′ns.]

The Pebble-town people warred with the people of Ninstints. The East Coast people1 were also at war with them. They were all at Gētgā′ñ.2

From that place the father of Qꜝaolga′s went to hunt. Then they saw a war canoe pass a place named Gia′gēs lying toward the south. It came along close to the shore and passed behind an islet. And they did not know those who were in it.

And when it was far off he started straight out to sea. And, when the rocky shore had nearly passed out of sight, he turned about. At evening they heard the sound of his guns at Gētgā′ñ. He had seen the enemy. Then they went for two shamans who were there. And they whipped the souls of the enemy. At that time they said that a white raven flew into the inlet. After they had drunk salt water for two nights all the warriors went out to meet the enemy.

After they had crept along close to the shore for a while they feared to round ʟuqā′łgᴀłdᴀs.3 So they stopped there. Some of them said that Kaisun could be seen from there. By and by, however, they went thither. They then saw some persons walking on the beach at Kaisun.

After some time had passed the canoe came in front of them (those at Kaisun). They got into it and went seaward. In the middle was a shaman whipping the souls of his enemies. Then one [Ninstints man] in it saw a strange sight. “Look at the cormorant flying about.

It has no head.” And, when they looked at it, its head was lacking. Kꜝadja′-i alone4 did not see this.

After they had gone on for a while a shot was fired at them. At once [the guns shooting] downward resounded everywhere. It (the canoe) turned bottom up. And as they came alongside they shot at them. And after they had destroyed them and had turned the canoe over one person was in it. He alone they saved. When the first gun sounded, the war chief said: “Sqas, take the gun away. It is not time for that.”

They then went away. Now they sang war songs. And the next day they went to get the heads. They then cut them off and dried them in the sunshine. The shaman who had whipped the souls of his enemies had his hair bunched together.5 He had told them to go back. On the way they (the Ninstints people) saw portents. They heard the [374]sound of drying frames dropping from above. And Tia6 also called near them. He called, and blood spurted out of his neck.

This encounter was referred to by others. See Memoirs of the American Museum of Natural History, volume V, part 1, page 31. [375]

The usual word applied to the east coast of the Queen Charlotte islands means “The coast where canoes can land easily.” ↑

A camping place about 1½ miles from Tcꜝā′ał. ↑

Probably means “Canoe-going-about.” ↑

The one who was saved. ↑

A shaman might not touch his hair with his fingers, and in consequence it became long and matted. ↑

The supernatural power that presided over slaughter and made his presence known at a time when it was about to take place. See Memoirs of the American Museum of Natural History, v, pt. 1, p. 31. ↑

F

IGHT

BETWEEN THE KAIGANI AND WEST COAST HAIDA

[Told by Richard of the Middle-gîtî′ns.]

They had occupied Thin-fort. There were many black cod there. Then they saw portents. The black cod, the day after they were brought in, moved their mouths. And at one time a headless cormorant came there.1 Some time after that, they say that [the children] who had a little fire in a cave below them and were picnicking there, ran out of it. Some small being with disheveled hair and a yellow-cedar-bark blanket over its shoulder came out of the cave. It was Supernatural-slave2 who was among them, they say. Its belly was big, they say. Then they feared to play in the cave.

After that the rotten gills lying about groaned. Another day Tia flew over to the fort from the opposite side. He said “Tia, tia,” and blood spurted out of his neck.1

One day, while they were away fishing for black cod, they (the enemy) came upon them. These were the Kaigani, Sta′stas, and Middle-town people. Then they shot up at the fort. My grandmother was born among them. That was why they did not touch her people.3

One man then shot from the fort. When his ammunition failed they went up to the fort. They enslaved all of the Pebble-town people. And, going out to those who were fishing, they destroyed half of them also. Some escaped to Kaisun.

When the warriors started off they were told about a child of the Middle-gîtî′ns,4 whose cousin was carrying her on her back. They would have taken her back, but were afraid. When they had

recrossed her friends adopted her.5 At the end of the following autumn they brought her back.

After that they sent word by canoe that they wanted to make peace. They (the West Coast people) then went thither. When they arrived at Tꜝē6 no attention was paid to them. And, since their food was gone, they wanted to buy food from one who lived opposite. He was stingy, and they laid hold of him. Although he was a chief they enslaved him. They also fought the people of Tꜝē. And they killed many of them, and those they enslaved were many. The Pebbletown people made matters even.

And, after they had talked over where they should have a fort, they made one on the west coast. Two were staying at Kꜝiū′stᴀ. After some time had passed the Kaigani people came to make peace. When they stopped in front they began to shoot at them. When they fled they [376]met the two persons7 in a canoe. When they were going to enslave them they jumped into the water. And they swam over to an islet. They now fled in terror.

Some time after that they went to the Kaigani country to fight again. Then they destroyed some people there. They enslaved ten and killed many. And, while they were on the way back, the wind was strong, and they threw some slaves overboard. They did this to four. The Kꜝiū′stᴀ people then came to them. And they took the slaves away from them and split up their canoe. They then started home around by the West Coast shore. They went to Tcꜝā′ał.8 They came to Lagî′nda,9 where people were catching salmon. A single slave was with them. The chief finished sending food through the fire.10 [377]

Compare the preceding story. ↑ a b

A being who appeared to persons that were about to be enslaved. ↑

Since his grandmother belonged to one of these families or to a closely related family they let her people alone. ↑

See the story of “A raid on the Tlingit,” notes. ↑

That is, members of her family in the Kaigani country. ↑

A town on the northwest coast of Graham island formerly owned by the Sandtown people and later probably by a branch of the Rear-town people. Richard pronounced the name Tꜝī′xī. ↑

The two persons just referred to as having remained at Kꜝiū′stᴀ. ↑

See the story of “Sacred-one-standing-and-moving, Stone-ribs, and Upward,” note 31. ↑

A creek on Graham island, running into the channel between it and Moresby. ↑

To the souls of those who had been slain. ↑

[Contents]

WARS

BETWEEN THE STIKINE AND SITKA TLINGIT

[Told by Richard of the Middle-gîtî′ns]

The nephew of Cēks lived at Sitka for the sake of some woman. He was killed there. Then all the Stikine people went to Sitka to fight. At that time they had a pitched battle there, and they destroyed many Sitka people. After that the Sitka people also started out to war.

At that time the eulachon were running into the Stikine. After they had filled the ground with holes they went out to get stones. They did not have the right kind of stones there. They had only whetstones. And, when they came from it, a man passed out by them to get some. His name was Daoł.1 Then he gave them the following prediction: “They will kill me [and my family] when the tide is on the ebb, and I shall never come back.”

As soon as the tide was out they went out to fight. The warriors came upon him at the place where he was gathering stones. Then they took him into the war chief’s canoe. He had left his gun behind. They then made fun of him by telling him to do various things.

While they were talking to him the warriors (his friends) came into the bay. They asked him then: “One-who-is-always-mentioned, are you still alive?” “Yes,” he said. He also asked: “Did you bring my gun? Did you bring my knife?” “Yes.” “Give them to me.” They then got his things to him.

When he put on his cartridge box some one shouted: “Ixiâ′⁺ī,2 Onewho-is-always-mentioned has his weapons in his hands.” Then, forgetting himself, he shot. Straightway they all shot. And he also said that he pulled out his knife and kept striking them as he moved

forward. They then destroyed the Sitka people. They stabbed those who there escaped to the woods. They said that two young fellows then came from watching for their enemies. They took them into Cēks’s canoe.

They then went away. On the next day they collected heads. Some of them got twelve. Others got ten. The heads were drying in the sunshine, looking like clothes drying on a line and bellying in the wind. Then Cēks, having called the people together, told them to stop fighting. He told them that they had destroyed the Sitka people.

By and by they let the two youths that they had enslaved go. They then ripped open the seams in a little, old, 3-fathom canoe. In it they started off. They also gave them a small paddle. They thought then that they would drown. In the summer after that they heard that they had escaped. [378]

Some time afterward the Sitka people were coming to make peace. And they got ready for them. They thought it well to make peace [they said]. Then Cēks again called the people together, and they agreed to destroy them. They then talked over how they should preserve themselves from injury.

By and by the Sitka people came in many canoes. After they had danced for a while in front of the town they came ashore, and the dancers entered the house and danced there. Then property was given to them. They gave them four or two slaves apiece as blood money. After that they also danced in Cēks’s house. When nearly all were in they shut the door, and they killed those outside and threw their bodies over the cliff in front.

After they had killed all of those they crowded against each other near the door. They then quietly pulled out one after another, stabbed each one, and threw out the bodies. At that time a woman

looked in through the smoke hole. She held a knife. She made the motion of cutting off heads. She said that they were going to destroy them. Those who were dancing paid no attention to her. After they had killed on for a while they began to discover it for themselves. But still they kept dancing. Although only ten were left they kept dancing. Presently they killed all. Six that they saved they let go home.

Some time afterward they began to visit back and forth. Once a great many went to [Sitka], and Qała′x3 paid a great sum to the Russians. Then many canoes came there and, when all the Stikine people were inside the stockade, Qała′x began to fight them. And they destroyed the Stikine people.

Some time after that they became good to each other. They began visiting back and forth again. Then ten canoes came to the Stikine, and Yaqoa′n began to kill them. And they destroyed all of them. At that time they stopped visiting each other.

Some time after that Qała′x’s nephew was in love with the daughter of a Russian. For that the Russians killed him. They then killed the Russian’s son. They said that the Russians were going to fight them with Qała′x, and they fortified themselves. They built the walls out of big cedars. And they built the houses inside. They put flat rocks along the fronts and sides of the houses.4 And, after they had lived there for a while, the Iron people5 came in a vessel to destroy them. After they had shot at them ten times they called for Qała′x. When he answered they shot at him still more.

After they had done this for a while they came off to fight them in three boats. All had guns with bayonets. They came on land at once. And, after they had prepared for them in the house, they went out. They then shot at them. While the Russians were shooting by command they shot into them. They also threw out their cartridges

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