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AVIAN RANGE DYNAMICS: TRAITS, BIOTIC INTERACTIONS AND NICHES IN CHANGING ENVIRONMENTS

Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften

vorgelegt beim Fachbereich Biologie der Johann Wolfgang Goethe- niversit!t in Frankfurt am "ain

von #rina $aube aus Bad %oden am &aunus

Frankfurt '()) *D +(,

vom Fachbereich Biologie der Johann Wolfgang Goethe- niversit!t als Dissertation angenommen-

Dekanin. /rof- Dr- 0nna %tarzinski-/owitz Erste Gutachterin. /rof- Dr- 1atrin B2hning-Gaese 3weite Gutacherin. /rof- Dr- 4atherine Graham

Datum der Dis5utation . '- "ai '()'

6When we try to pick out anything by itself, we find it hitched to everything else in the Universe.7

John "uir *)8)),

#llustrations b9 Wilhelm von Wright *):)(;)::<,- /ublic domain-

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1.1 Hintergrund und Zielsetzung.................................................................................................1 1.2 Zu einem mechanistischeren Verstndnis von Artmerkmalen und Verbreitungsgebietsgren ( a!itel "#..................................................................................$ 1." onkurrenz und Ausbreitungs%higkeit interagieren bei der &estimmung der geogra!hischen Verbreitung von Vgeln ( a!itel $#...........................................................'

1.$ (ischenver%)gbarkeit in Zeit und *aum+ Vogelzug der ,rasm)cken ( a!itel '#............' 1.' -chluss%olgerungen...................................................................................................................

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2.1 &ackground............................................................................................................................../ 2.2 -tructure and aims o% the thesis............................................................................................11

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".1 Abstract...................................................................................................................................1. ".2 0ntroduction............................................................................................................................1. "." 1ethods...................................................................................................................................12 +-+-) %tud9 s5ecies and areaA geogra5hic range sizes------------------------------------------------------------): +-+-' %5ecies traits------------------------------------------------------------------------------------------------------------------)8 +-+-+ %tatistical anal9ses---------------------------------------------------------------------------------------------------------'+ ".$ *esults.....................................................................................................................................2$ ".' 3iscussion...............................................................................................................................24

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$.1 Abstract..................................................................................................................................."2 $.2 0ntroduction............................................................................................................................"2 $." 1ethods..................................................................................................................................."4 B-+-) Data-------------------------------------------------------------------------------------------------------------------------------+< B-+-' %5ecies distribution models and range filling --------------------------------------------------------------+8 B-+-+ /otential determinants of range filling--------------------------------------------------------------------------B) $.$ *esults.....................................................................................................................................$$ $.' 3iscussion...............................................................................................................................$2

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'.1 Abstract...................................................................................................................................'$ '.2 0ntroduction............................................................................................................................'$ @-'-) 4once5tual framework for niche d9namics in s5ace and time--------------------------------------@E @-'-' 055l9ing the framework to animal migration---------------------------------------------------------------@: '." 1ethods....................................................................................................................................1 @-+-) %tud9 s5ecies-----------------------------------------------------------------------------------------------------------------E) @-+-' >anges---------------------------------------------------------------------------------------------------------------------------E) @-+-+ Environmental variables------------------------------------------------------------------------------------------------E) @-+-B Niche characteristics-----------------------------------------------------------------------------------------------------E' @-+-@ %tatistical 0nal9sis--------------------------------------------------------------------------------------------------------EB '.$ *esults....................................................................................................................................... '.' 3iscussion...............................................................................................................................41

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..1 5uture research !ers!ectives................................................................................................4. ..2 6oncluding remarks..............................................................................................................4/

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A!!endi7 1+ -tud8 s!ecies ........................................................................................................../2 A!!endi7 2+ -31 threshold sensitivit8 anal8sis.....................................................................192 A!!endi7 "+ Akaike model selection : %ull model regression results %or all habitat suitabilit8 levels....................................................................................................................19. A!!endi7 $+ *ange %illing estimates %or di%%erent -8lvia s!ecies and distribution modelling methods..................................................................................................................................11" A!!endi7 '+ ,ridding threshold sensitivit8 anal8sis..............................................................11$

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1 Zusammenfassung

1 Z;-A11<(5A--;(,
1.1 Hintergrund und Zielsetzung
Das geogra5hische Derbreitungsgebiet von 0rten ist ein fundamentales %truktur gebendes "erkmal der biologischen Welt- #n &eilen selbst fFr Gelegenheitsbeobachter zug!nglichA fFhrt das Wahrnehmen und Beschreiben der geogra5hischen Derbreitung von 0rten unvermeidlich zu Fragen Fber die treibenden FaktorenA die sie bestimmen- Warum 0rten so verteilt sind wie sie sindA ist seit langem eine der zentralen Fragen in GkologieA Biogeogra5hie und Evolution *WallaceA ):<EH "ac0rthurA )8<'H GastonA '((+,- Der 0nbruch des 0nthro5oz!ns stellt diese Frage mit neuer Dringlichkeit *4rutzenA '((',- Wurden geogra5hische 0rtverbreitungen zuvor durch die D9namik der natFrlichen Welt bestimmtA so sind menschliche 0ktivit!ten Ietzt der dominante FaktorA der sie auf Iedem "aJstab von der lokalen mgebung bis hin zum gesamten Erds9stem formt *$adle K WhittakerA '()),- Die menschliche F!higkeitA globalen Wandel auszul2sen und zu beschleunigenA Fbertrifft Iedoch unser Derst!ndnis der 0uswirkung auf 0rtverbreitungen und unsere F!higkeitA die 1onseLuenzen unserer gesellschaftlichen Entscheidungen fFr die Derteilung von Biodiversit!t vorherzusagen *WhiteA )8E<H 4ha5in et al.A '(((,- Gegenw!rtig ver!ndern sichA im Wesentlichen als unbeabsichtigtes Neben5rodukt menschlicher 2konomischer 0ktivit!ten und /o5ulationsd9namikA die geogra5hischen Derbreitungsgebiete von 0rten mit entscheidender Bedeutung in der $and- und ForstwirtschaftA als 1rankheitsvektoren oder als &eil der biologischen %9stemeA die Gkos9stemfunktionen bereitstellen *"E0A '((@aH /armesanA '((E,- Die steigende Besorgnis Fber invasive 0rten zeigt aufA dass der menschliche Einfluss auf 0rtverbreitungen oft unbeabsichtigt und unsere F!higkeit zur zielgerichteten 1ontrolle von Derbreitungen begrenzt ist *EltonA )8@:H /imentel et al., '((),- Daher ist es entscheidendA dass wir unser Derst!ndnis Fber die D9namikenA aus denen die geogra5hische Derbreitung von 0rten erwachsenA verbessern *Davis et al.A )88:,Die D9namik von 0rtverbreitungen zu verstehen ist eine ?erausforderungA da geogra5hische 0rtverbreitungen von einer Dielzahl interagierender Faktoren beeinflusst werden *"ac0rthurA )8<',- Mber die grundlegende #dee hinausA dass mweltbedingungen die Derbreitung von 0rten begrenzenA h!ngt das Dorkommen einer 0rt an einem bestimmten geogra5hischen /unkt auch davon abA ob dieser /unkt der 0rt 5rinzi5iell zug!nglich ist und )

1 Zusammenfassung welche biologische Gemeinschaft dort bereits besteht */ulliamA '(((H %oberonA '((<,- Es wurde vorgeschlagenA die relative Bedeutung dieser 0s5ekte sei skalenabh!ngig. abiotische Bedingungen wie z-B- 1lima und 0usbreitungsbeschr!nkungenA die sich aus der Geschichte einer 0rt ergebenA sind im Wesentlichen auf groJer r!umlicher %kala relevantA wohingegen biotische #nteraktionen zunehmend auf kleiner %kala an Bedeutung gewinnen */earson K DawsonA '((+H Guisan K >ahbekA '()),- #n Iedem Fall kann das geogra5hische Derbreitungsgebiet einer 0rt von ihren gegenw!rtigen "erkmalenA wie ?abitatwahl und 0usbreitungsf!higkeitA von ihrer evolution!ren und biogeogra5hischen GeschichteA von der raumzeitlichen D9namik ihrer bevorzugten mweltbedingungen sowie von den "erkmalen und der Geschichte anderer 0rten beeinflusst werden *NewtonA '((+H /rice K 1irk5atrickA '((8,- Noch haben wir kein koh!rentes Bild davonA wie diese Faktoren interagierenA von ihrer relativen Bedeutung oder davonA wie diese Beziehungen zwischen verschiedenen &aNa variierenW!hrend viele %tudien den Einfluss einzelner Faktoren auf einzelne 0rten dokumentierenA gab es erst im letzten Jahrzehnt Fortschritte in der BioinformatikA die es uns erm2glichen durch die #ntegration multi5ler Faktoren in derselben 0nal9se und das ntersuchen von "ustern auf groJen r!umlichen %kalen Fber viele 0rten hinweg die D9namik von 0rtverbreitungen besser zu verstehen *BrownA )88@H GastonA '((+H Brooker et al.A '((<,- ?eute haben Forscher nie dagewesenen 3ugang zu Daten Fber die DerbreitungA Gkologie und Evolution von 0rtenA zu DatenA die die mweltbedingungen der gesamten Erdoberfl!che beschreiben und zu den bioinformatischen WerkzeugenA um diese #nformationen zu organisierenA zu anal9sieren und zu integrieren *z- B- Graham et al.A '((BH >angel et al.A '((EH 1ozak et al.A '((:,- Dies hat zu einer raschen methodischen Entwicklung in Bereichen wie der 0rtverbreitungsmodellierung *Guisan K &huillerA '((@H Elith et al.A '((E, gefFhrtA die ihrerseits eine Neubetrachtung des klassischen 1onze5ts der 2kologischen Nische *GrinellA )8)<H EltonA )8'<H ?utchinsonA )8@<, und eine Diskussion Fber die #ntegration der durch die neuen "ethoden ins5irierten konze5tionellen #deen in die Nischentheorie */ulliamA '(((H %oberonA '((<H /earman et al., '((:H 4olwell K >angelA '((8H Wiens et al.A '()(, angeregt hat- 0rtverbreitungsmodelle sind weiterhin fFr das Derstehen der D9namik von 0rtverbreitungen von groJem WertA doch haben sieA wie Iedes wissenschaftliche WerkzeugA bisweilen wissenschaftliches Denken auch eingeschr!nkt. durch den Fokus auf groJe raumzeitliche %kalen und das 1lima als treibenden Faktor sowie durch

'

1 Zusammenfassung die Derwendung von 0lgorithmenA die 0rtverbreitung als 5r!zise definierteA statische Entit!ten behandeln und die ebenso statische Nischen berechnen *0raOIo K GuisanA '((EH Fisher et al., '()(H FranklinA '()(,"it dieser Doktorarbeit versuche ichA einen Beitrag zu unserem in Entwicklung begriffenen Derst!ndnis der multi5len FaktorenA die 0rtverbreitungsgebiete beeinflussenA zu leisten#ch verwende "ethoden aus BioinformatikA %tatistik und G#% *Geoinformationss9steme, und kombiniere Daten zu DerbreitungsgebietenA "erkmalenA GkologieA Evolution sowie der gegenw!rtigen und vergangenen Derst!ndnis der "echanismenA welche Gr2JeA /osition und mweltA um unser D9namik von

Derbreitungsgebieten bestimmenA zu verbessern- #ch versuche ebenfallsA konze5tionell Fber die klassische 0rtverbreitungsmodellierung hinauszugehenA indem ich die raumzeitliche D9namik des verfFgbaren Nischenraums und die d9namische Natur der Nischenanforderungen berFcksichtige und somit die gegenw!rtige Diskussion Fber Nischentheorie um eine zus!tzliche /ers5ektive bereichere- &aNonomisch konzentriere ich mich auf die D9namik der Derbreitungsgebiete von D2gelnA s5ezifisch von euro5!ischen %ingv2geln in 1a5itel + und der Gattung Sylvia in 1a5itel B und @0ls ein "odells9stemA um die treibenden Faktoren von 0rtverbreitungen zu untersuchenA haben D2gel einige bedeutende Dorteile- D2gel haben schon immer die 0ufmerksamkeit von 0mateurenA Naturforschern und Gkologen auf sich gezogen und sind somit eine der am besten untersuchten =rganismengru55en- Der >eichtum an #nformationen zu DerbreitungA Gkologie und Evolution der D2gel erlaubt esA das /otential bioinformatischer "ethoden voll auszusch25fen- Euro5!ische %ingv2gel im Besonderen bieten uns die GelegenheitA entscheidende "erkmale wie die 0usbreitungsf!higkeit von mor5hologischen "essungen abzuleiten *Dawideit et al., '((8,- Die Gattung Sylvia *GrasmFcken, verbindet eine immense Dariation in Gr2Je und 1onfiguration von Derbreitungsgebieten mit einer langen &radition von %tudien zu biotischen #nteraktionen innerhalb der Gattung *z- B- 4od9 K WalterA )8<EH "artin K &hibaultA )88EH /ons et al., '((:, und zeigt das volle %5ektrum von "igrationsverhaltenA das bei D2geln generell zu beobachten ist *%hirihai et al., '((),- Die Gattung ist somit ideal geeignetA um die Beziehung zwischen biotischen #nteraktionenA 3ugverhalten und 0rtverbreitungsd9namik zu untersuchen- "eine drei eigenst!ndige 1a5itel gegliedert. ntersuchungen sind in

1 Zusammenfassung

1.2 Zu einem mechanistischeren Verstndnis von Artmerkmalen und Verbreitungsgebietsgren ( a!itel "#
Ein wichtigerA ungel2ster Fragenkom5leN in der "akro2kologie istA die immense inters5ezifische Dariation in der Gr2Je geogra5hischer Derbreitungsgebiete zu verstehenW!hrend man davon ausgehtA dass 0rtmerkmale wie Fekundit!t und 12r5ergr2Je einen Effekt auf Derbreitungsgebietsgr2Jen habenA fehlt ein allgemeines Derst!ndnis davonA wie Derbreitungsgebietsgr2Jen von mehreren "erkmalen gemeinsam beeinflusst werden- ?ier haben wir den Einfluss einer Dielzahl von 0rtmerkmalen auf die Gr2Je der globalen Derbreitungsgebiete euro5!ischer %ingv2gel getestetA um die m2glichen "echanismen hinter makro2kologischen 3usammenh!ngen besser zu verstehenWir haben den Effekt 2kologischen von $ebensgeschichtsmerkmalen *?abitatnischeA *Fekundit!tA 0usbreitungsf!higkeit,A "erkmalen NahrungsnischeA

3ugverhaltenA FleNibilit!t im 3ugverhalten, und mor5hologischen "erkmalen *12r5ergr2Je, auf die globale Derbreitungsgebietsgr2Je von )E@ euro5!ischen %ingv2geln beurteilt- Wir identifizierten ?95othesen zur Beziehung von 0rtmerkmalen und Derbreitungsgebietsgr2Jen aus der $iteratur und verwendeten die "ethodik der /fadanal9seA um sie zu testenFekundit!tA 0usbreitungsf!higkeitA ?abitatnischenbreite und Nahrungsnischen5osition hatten einen direkten 5ositiven Effekt auf die Derbreitungsgebietsgr2Je- 3ugverhalten hatte einen indirekten 5ositiven Effekt via 0usbreitungsf!higkeit- 12r5ergr2Je hatte einen starkenA direkten 5ositiven EffektA der durch indirekte negative Effekte Fber mehrere andere "erkmale reduziert wurdeDie Gr2Je der globalen geogra5hischen Derbreitungsgebiete euro5!ischer %ingv2gel wurde von $ebensgeschichtsmerkmalen *Fekundidt!t und 0usbreitungsf!higkeit,A 2kologischen "erkmalen *?abitatnischenbreiteA Nahrungsnischen5osition und 3ugverhalten, und von 12r5ergr2Je beeinflusst- 0rtmerkmale beeinflussten Derbreitungsgebietsgr2Jen auf direktem und indirektem Weg- #nsbesondere der Einfluss von 12r5ergr2Je war mit 5ositiven und negativen Effekten Fber verschiedene /fade sehr kom5leN- Die Gr2Je von Derbreitungsgebieten ist sehr wahrscheinlich auch von anderen Faktoren als von 0rtmerkmalen abh!ngig- Wir konnten zeigenA dass es notwendig istA den direkten und indirekten Einfluss einer Dielzahl von "erkmalen zu entwirrenA um die "echanismenA auf denen makro2kologische Beziehungen beruhenA aufzukl!renB

1 Zusammenfassung

1." onkurrenz und Ausbreitungs%higkeit interagieren bei der &estimmung der geogra!hischen Verbreitung von Vgeln ( a!itel $#
Es ist weiterhin eine ?erausforderung fFr Gkologie und EvolutionsbiologieA die Faktoren zu verstehenA welche die geogra5hische Derbreitung von 0rten beeinflussen- #nsbesondere besteht wenig 1onsens darFberA ob biotische #nteraktionen wie inters5ezifische 1onkurrenz Derbreitungsgebiete bestimmenWir untersuchen EinflFsse von 1onkurrenzA 0usbreitungsf!higkeitA das 0lter eines &aNons und ?abitatverschiebungen seit dem letzten glazialen "aNimum auf das 0usmaJA in dem 0rten der Dogelgattung Sylvia in allen >egionen mit geeigneten mweltbedingungen vorkommen *d- h- range filling,Wir haben range filling in der Dogelgattung Sylvia unter Derwendung von Boosted >egression &rees und >idge->egression Luantifiziert- "ittels multi5ler >egression haben wir fFr die Effekte von intragenerischer 1onkurrenzA 0usbreitungf!higkeitA 0lter des &aNons und ?abitatverschiebung seit dem letzten glazialen "aNimum auf range filling getestetm verschiedene ?95othesen widerzus5iegelnA wie lokale 1onkurrenz die D9namik von Derbreitungsgebieten auf groJer r!umlicher %kala beeinflussen k2nnteA haben wir unterschiedliche "ethoden verwendetA um 5otentielle %ignale von 1onkurrenz auf der %kala des Derbreitungsgebiets zu LuantifizierenGrasmFcken mit hoher 0usbreitungsf!higkeit zeigten h2heres range fillingA aber nur wenn 1onkurrenz in Gebieten mit weniger geeignetem ?abitat innerhalb ihres 5otentiellen Derbreitungsgebietes niedrig war- Das 0lter eines &aNons und ?abitatverschiebung seit dem letzten glazialen "aNimum hatten keinen konsistenten EffektWir konnten somit zeigenA dass die Derbreitungsgebiete von GrasmFcken mit hoher Wahrscheinlichkeit durch den simultanenA interaktiven Effekt von 1onkurrenz und 0usbreitungsf!higkeit geformt werden- Wenn biotische #nteraktionen wie 1onkurrenz generell die F!higkeit von 0rten beeinflussenA auf der kontinentalen %kala neue Gebiete zu kolonisierenA wird es in der &at eine ?erausforderung seinA den Effekt von 1limawandel auf Biodiversit!t vorherzusagen-

1 Zusammenfassung

1.$ (ischenver%)gbarkeit in Zeit und *aum+ Vogelzug der ,rasm)cken ( a!itel '#
#m 1onteNt neuer Fortschritte in der 2kologischen Nischenmodellierung sind sowohl die mwelt als auch die 2kologische Nische einer 0rt als statische Entit!ten behandelt und Luantifiziert worden- #n der >ealit!t sind aber sowohl die mwelt als auch die Nischenanforderungen einer 0rt auf einer Dielzahl von %kalen d9namisch- Wir schlagen ein konze5tionelles %9stem vorA das berFcksichtigtA wie die realisierte Nische und geogra5hische Derbreitung von 0rten durch die entko55elte raumzeitliche DerfFgbarkeit unterschiedlicher mweltbedingungen und durch Der!nderungen der Nischenanforderungen Fber die $ebenszeit eines =rganismus geformt werdenDas &esten der aus dem konze5tionellen %9stem abgeleiteten Dorhersagen am Beis5iel des Dogelzugs der GrasmFcken ergab neue Erkenntnisse. Das Derfolgen der 1limanische im geogra5hischen >aum war h2chstwahrscheinlich nicht die treibende 1raft fFr "igration in der Gattung und steht 5otentiell im 1onflikt mit dem Derfolgen der $andnutzungsnische- Die Nischen der GrasmFcken waren w!hrend der Brutsaison schmalerA was zeigtA dass Nischenanforderungen zeitlich d9namisch sein k2nnen- Wir legen naheA dass die BerFcksichtigung Bewegung von d9namischer =rganismen im mwelten >aum und und Nischenanforderungen der D9namik ihrer zu einer und entscheidenden Derbessserung unseres Derst!ndnisses der treibenden Faktoren hinter der Nischen Derbreitungsgebiete fFhrt-

1.' -chluss%olgerungen
#n der vorliegenden Doktorarbeit habe ich versuchtA unser gegenw!rtiges Derst!ndnis der D9namik von Dogelverbreitungsgebieten durch die "odellierung mutmaJlicher "echanismenA die #ntegration multi5ler Faktoren in einer einzigen 0nal9se und durch die Entwicklung neuer konze5tioneller #deen zu erweitern- 3u diesem 3weck habe ich Datens!tze aus GkologieA Evolution und den Erdwissenschaften kombiniert und moderne statistische Werkzeuge wie Geoinformationss9stemeA statistische /rogrammierumgebungenA /fadanal9seA Boosted >egression &reesA >idge->egressionA Bootstra55ingA 1erndichtesch!tzer und Nischenmetrik eingesetzt-

1 Zusammenfassung Es ergeben sich folgende ?au5terkenntnisseA wobei es wichtig ist zu betonenA dass ihre Dalidit!t durch die notwendige Beschr!nkung auf %ingv2gel als "odells9stem auf diese Gru55e beschr!nkt bleibtA bis sie fFr andere &aNa best!tigt oder widerlegt werden k2nnen. *i, 0rtmerkmale k2nnen keinen GroJen 0nteil der Dariation in Derbreitungsgebietsgr2Jen erkl!renA aber sie s5ielen eine wichtige >olle- "ehrere 0rtmerkmale beeinflussen Derbreitungsgebietsgr2Je auf kom5leNe WeiseA sowohl direkt als auch indirekt Fber andere "erkmale- *ii, =b 0rten in der $age sindA geeignete 0reale auf groJen r!umlichen und zeitlichen %kalen zu kolonisierenA h!ngt von mehrerenA interagierenden Faktoren ab- Entgegen bestehender Dorstellungen */earson K DawsonA '((+H Guisan K >ahbekA '()), k2nnten biotische #nteraktionen Derbreitungsgebiete auf kontinentaler %kala beeinflussenA wobei ihre Effekte sehr wahrscheinlich von der ?abitatgFte modifiziert werden- *iii, Die Nischen und Derbreitungsgebiete von 0rten sind d9namische Entit!tenA die von der raumzeitlichen DerfFgbarkeit von mweltbedingungen abh!ngenDie DerfFgbarkeit solcher mweltbedingungen kann fFr verschiedene Nischendimensionen as9nchron seinA was 0rten vor kom5leNe =5timierungs5robleme stelltA wenn sie versuchenA mweltbedingungen im geogra5hischen >aum zu verfolgen- Die Nischenanforderungen von 0rten k2nnen Fber ihren $ebensz9klus hinweg variieren- 3usammenfassend kann die frFhe DorstellungA dass Derbreitungsgebiete nur von wenigen Faktoren bestimmt sind *z- B- &wome9A )8+E,A widerlegt werden- Die /rozesseA welche die Gr2JeA /osition und D9namik von Derbreitungsgebieten bestimmenA sind hochkom5leN und involvieren multi5leA interagierende &riebkr!fte- Wir stehen erst am Beginn der Entwicklung eines koh!rentenA umfassenden Derst!ndnis der D9namik von 0rtverbreitungsgebieten#m ?inblick auf zukFnftige Forschung gibt es einige BereicheA in denen die BerFcksichtigung zus!tzlicher 1om5leNit!t unser Derst!ndnis von 0rtverbreitungen voranbringen kann- Diese sollten insbesondere bei der Dorhersage der Derschiebung von 0rtverbreitungen durch globalen Wandel BerFcksichtigung finden. *i, Es gibt immer noch viele ungenutzte "2glichkeiten der #ntegration multi5ler &riebkr!fte der D9namik von 0rtverbreitungsgebieten in einer einzigen 0nal9se *Botkin et al.A '((<,A wofFr sich z- BBa9esische "ethoden anbieten *EllisonA '((BH 4ho9 et al.A '((8,- *ii, Welche Faktoren die Derbreitung von 0rten bestimmenA variiert im geogra5hischen >aum *z- B- BarnesA )8@<A Gross K /riceA '(((,- Die BerFcksichtigung solcher Dariation z- B- durch geogra5hisch gewichtete >egression erscheint vielvers5rechend *0ustinA '((<,- *iii, Die 0nnahmeA dass

<

1 Zusammenfassung 0rtverbreitungen auf groJer r!umlicher %kala nicht durch biotische #nteraktionen bestimmt werdenA sollte mit groJer Dorsicht betrachtet werden- Es erscheint ratsamA biotische #nteraktionen besser in bestehende "ethoden zu integrierenA was bedeuten k2nnteA dass die Derschiebung des Derbreitungsgebietes einer 0rt nicht in #solation von anderen 0rten modelliert werden kann *1eith et al., '((:H Baselga K 0raOIoA '((8,- *iv, Wenn Nischenanforderungen d9namisch sindA sollten die Nischen und Derbreitungsgebiete zu verschiedenen %tadien im $ebensz9klus einer 0rt getrennt betrachtet und modelliert werden *Doswald et al.A '((8H Jackson et al.A '((8,- *v, Wie 0rten mit der raumzeitlichen Des9nchronisation verschiedener Nischendimensionen umgehenA kann uns helfen zu verstehenA wie 0rten auf das vorhergesagte zukFnftige 0uftreten neuer 1limaregimes reagieren k2nnten *Williams et al.A '((<,- *vi, Das in 1a5itel @ vorgeschlagene konze5tionelle %9stem k2nnte auch zur Betrachtung der Der!nderung von Nischen und 0rtverbreitungen im Derlauf der EvolutionA z- B- im 1onteNt von Nischenkonservatismus verwendet werden *Wiens K GrahamA '((@H 4ris5 et al.A '((8,Die D9namik von 0rtverbreitungen ist kom5leN- #m >ahmen von Wissenstransfer zu betonenA was wir gegenw!rtig Fber die 0uswirkungen von globalem Wandel auf Biodiversit!t nicht wissen und nicht vorhersagen k2nnenA k2nnte den gesellschaftlichen Diskurs Fber >isikoakze5tanz und /lanung unter Einbeziehung von nsicherheit anregen *Dasgu5taA '((:H 44%/A '((8H Dawson et al.A '()),-

2 ntroduction

2 0(=*>3;6=0>(
2.1 &ackground
&he geogra5hic range of s5ecies is a fundamental 5ro5ert9 structuring the biological world0ccessible and observable in 5arts even to the casual naturalistA noticing and recording the geogra5hic distribution of s5ecies leads inevitabl9 to asking Luestions about the driving forces that sha5e it- Wh9 s5ecies are distributed in the wa9 the9 are has long been recognised as a central Luestion in ecolog9A biogeogra5h9 and evolution *WallaceA ):<EH "ac0rthurA )8<'H GastonA '((+,&he rise of the 0nthro5ocene 5oses this Luestion with new urgenc9 *4rutzenA '((',- #f geogra5hic ranges of s5ecies have 5reviousl9 been set b9 d9namics inherent to the natural worldA human activit9 is now the dominant agent sha5ing them at ever9 scale from the local 5atch to the whole Earth s9stem *$adle K WhittakerA '()),- ?oweverA the human abilit9 to trigger and accelerate global change eNceeds our understanding of how this will alter s5ecies ranges and our abilit9 to 5redict the conseLuences of societal decisions for the distribution of biodiversit9 *WhiteA )8E<H 4ha5in et al.A '(((,- $argel9 as an inadvertent b9-5roduct of human economic activities and 5o5ulation d9namicsA the geogra5hic ranges of s5ecies that are cruciall9 im5ortant to humans in agriculture and forestr9A as disease vectors or as 5art of the biological s9stems that maintain ecos9stem functionsA undergo far-reaching change *"E0A '((@aH /armesanA '((E,- &he increasing concern over invasive s5ecies highlights that the human im5act on s5eciesP ranges is often far from deliberate and that our abilit9 to 5ur5osefull9 control their distribution is limited *EltonA )8@:H /imentel et al., '((),&hereforeA it is im5erative that we enhance our understanding of the d9namics that give rise to s5ecies? geogra5hic distributions *Davis et al.A )88:,nderstanding range d9namics is challenging because geogra5hic ranges de5end on and are influenced b9 a multitude of interacting drivers *"ac0rthurA )8<',- Be9ond the fundamental idea that environmental conditions limit s5ecies distributionsA whether a s5ecies occurs in a 5articular geogra5hic location also de5ends on whether that area is in 5rinci5le accessible to the s5ecies and on the biological communit9 alread9 5resent */ulliamA '(((H %oberonA '((<,- #t has been suggested that the relative im5ortance of these as5ects is scalede5endentA such that abiotic conditions like climate and dis5ersal limitations resulting from a 8

2 ntroduction s5ecies? histor9 5rimaril9 govern distributions at large s5atial scales whereas biotic interactions gain increasing im5ortance at smaller scales */earson K DawsonA '((+H Guisan K >ahbekA '()),- #n an9 caseA a s5ecies? geogra5hic range can be influenced b9 its 5resent traitsA such as habitat 5reference and dis5ersal abilit9A b9 its evolutionar9 and biogeogra5hic histor9A b9 the s5atio-tem5oral d9namics of its 5referred environmental conditions and b9 the traits and histor9 of co-occurring s5ecies *NewtonA '((+H /rice K 1irk5atrickA '((8,- 0t 5resentA we do not have a coherent understanding of how these different drivers interactA of their relative im5ortance and of how these relationshi5s change across a broad range of taNaWhile man9 studies document the im5act of single factors on the range of individual s5eciesA the 5ast decade has seen bioinformatic advances that give us the o55ortunit9 to better understand range d9namics b9 integrating multi5le drivers in the same anal9sis and eNamining 5atterns across man9 s5ecies at large s5atial scales *BrownA )88@H GastonA '((+H Brooker et al.A '((<,- &oda9A researchers have un5recedented access to data on s5ecies distributionsA ecolog9 and evolutionA to data describing environmental conditions on the entire Earth?s surface and to the bioinformatic tools to manageA anal9se and integrate this information *e-g- Graham et al.A '((BH >angel et al.A '((EH 1ozak et al.A '((:,- &his has led to the ra5id methodological develo5ment of fields such as s5ecies distribution modelling *Guisan K &huillerA '((@H Elith et al.A '((E, which has in turn stimulated a re-eNamination of the seminal conce5t of the ecological niche *GrinellA )8)<H EltonA )8'<H ?utchinsonA )8@<, and an ongoing discussion about how to integrate conce5tual ideas ins5ired b9 the new a55roaches into niche theor9 */ulliamA '(((H %oberonA '((<H /earman et al., '((:H 4olwell K >angelA '((8H Wiens et al.A '()(,- While s5ecies distribution models continue to be of great value for understanding range d9namicsA the9 haveA like an9 scientific toolA sometimes also constrained scientific thought b9 focusing on large s5atio-tem5oral resolutions and on climate as a determinant of ranges and b9 using algorithms that treat geogra5hic ranges as 5recisel9 definedA static entities and that Luantif9 an eLuall9 static niche *0raOIo K GuisanA '((EH Fisher et al., '()(H FranklinA '()(,#n this thesisA # tr9 to contribute to our emerging understanding of the multi5le drivers that govern s5ecies distributions- # use bioinformaticsA statistics and G#% *geogra5hic information s9stems, methods and combine data on rangesA traitsA ecolog9A evolution and the 5ast and 5resent environment to inform our thinking about the mechanisms that determine the sizeA location and d9namics of s5ecies ranges- # also attem5t to conce5tuall9 go be9ond the classic

)(

2 ntroduction s5ecies distribution modelling a55roach b9 considering the s5atio-tem5oral d9namics of available niche s5ace and the d9namic nature of niche reLuirements and to thus add another 5ers5ective to the current debate about niche theor9&aNonomicall9A # focus on the range d9namics of birdsA s5ecificall9 Euro5ean 5asserines in cha5ter + and the genus Sylvia in cha5ter B and @- 0s a model s9stem to eN5lore the drivers of s5ecies distributionsA birds have several ke9 advantages- &he9 have alwa9s attracted the attention of amateursA naturalists and ecologists and are thus one of the best-studied grou5s of organisms- &he wealth of information on bird distributionsA traitsA ecolog9 and evolution allows us to use bioinformatic methods to their fullest advantage- Euro5ean 5asserinesA in 5articularA 5rovide us with an o55ortunit9 to gauge crucial traitsA such as dis5ersal abilit9A from mor5hological measurements *Dawideit et al., '((8,- &he genus Sylvia combines large variation in the size and configuration of ranges with a long tradition of studies investigating intrageneric biotic interactions *e-g- 4od9 K WalterA )8<EH "artin K &hibaultA )88EH /ons et al., '((:, and also eNhibits the full s5ectrum of migrator9 behaviours seen in birds generall9 *%hirihai et al., '((),- #t is thus ideall9 suited to investigate relationshi5s between biotic interactionsA migration and range d9namics-

2.2 -tructure and aims o% the thesis


# have organised the research carried out as 5art of this thesis into three maIor cha5ters- Each cha5ter is self-contained and structured in the st9le of a Iournal 5ublicationA with an abstract followed b9 the sections introductionA methodsA results and discussion- 0ll references and su55lementar9 information are given in a common reference list and a55endiN at the end of the thesis- &he main research cha5ters are followed b9 one final cha5ter containing a general s9nthesis and conclusions#n cha5ter +A # aim to im5rove our understanding of how s5ecies? traits interact to influence the size of geogra5hic ranges- # follow a macroecological a55roachA focusing on 5atterns across man9 taNa at large s5atial scales- ?ereA # consider the global range sizes of )E@ Euro5ean 5asserine s5ecies- # relate range size to multi5le traits of these bird s5ecies in a 5ath modelA which allows me to consider com5leN interactions among traits as well as the direct and indirect effects of traits on range size- &he aim here is to include a multitude of lifehistor9A ecological and mor5hological traits and link them to range size in a wa9 that reflects 5utative mechanistic relationshi5s re5orted in the literature- &his 5rovides us with an ))

2 ntroduction o55ortunit9 to assess the relative im5ortance of different traits and gain a better understanding of how the9 influence ranges- &his cha5ter builds 5artl9 u5on the di5loma thesis of ?eiko 1orntheuerA who took the mor5hological measurements of bird museum s5ecimensA conducted a 5reliminar9 anal9sis relating s5ecies? traits to their occu5anc9 in Euro5e and wrote this 5reliminar9 anal9sis into a manuscri5t draft- # have related the trait data to the global ranges of the s5eciesA refined the statistical methodolog9A calculated new 5ath modelsA incor5orated a new 5h9logen9 to test for 5otential bias due to relatedness and rewritten the manuscri5t- 1atrin B2hning-Gaese 5rovided data on s5ecies? traits other than dis5ersal abilit9 while 4arsten >ahbek 5rovided data on the s5ecies? global geogra5hic ranges- "onika %chwagerA %ven &rautmann and 1atrin B2hning-Gaese contributed to the stud9 design and manuscri5t#n cha5ter BA # aim to elucidate to what eNtent biotic interactionsA s5ecies? traitsA the evolutionar9 histor9 of s5ecies and the s5atio-tem5oral histor9 of the environment can 5revent s5ecies from colonising 5otentiall9 suitable habitat- ?ereA # narrow the taNonomic focus on the genus SylviaA which allows me to consider 5otential biotic interactions that have been re5orted for the genus from local studies- # use advanced G#% and s5ecies distribution modelling techniLues to estimate the 5otential ranges of the Sylvia warblers for the 5resent and the last glacial maNimum- # then relate 5otential intrageneric com5etitionA dis5ersal abilit9A taNon age and the amount of shift in the geogra5hic location of 5otential habitat since the last glacial maNimum to range fillingA i-e- the 5ercentage of the 5resent 5otential range that the s5ecies actuall9 occu59- For this cha5terA # have collated the data from different sourcesA conducted all G#%A statistical and s5ecies distribution modelling anal9ses and drafted the manuscri5t- 1atrin B2hning-Gaese and 4atherine ?- Graham contributed to the stud9 design and manuscri5t writing#n cha5ter @A # aim to enrich the current debate about new eNtensions of niche theor9 b9 a 5ers5ective that focuses on the highl9 d9namic nature of niches and ranges- # 5resent a conce5tual framework for how the s5atio-tem5oral d9namics in the environmental conditions available to a s5ecies ma9 affect its nicheA its distribution and its movements in geogra5hic s5ace- # also highlight the 5otentiall9 d9namic nature of niche reLuirements over a s5ecies? life-c9cle- From this frameworkA # derive 5redictions for the relationshi5 between niches and s5atio-tem5oral range d9namics and then test these 5redictions using migration in Sylvia warblers as a model s9stem- &o this endA # make use of new methods to Luantif9 the

)'

2 ntroduction characteristics of s5ecies? niches- # have develo5ed the frameworkA conducted all anal9ses and drafted the manuscri5t- 4atherine ?- Graham and 1atrin B2hning-Gaese contributed to the stud9 design and manuscri5t writing-

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%ubmitted to an international Iournal of ecolog9 as. $aubeA #-A 1orntheuerA ?-A %chwagerA "-A &rautmannA %-A >ahbekA 4- K B2hning-GaeseA 1*submitted, &owards a more mechanistic understanding of traits and range sizes-

)@

" #owards a more mechanistic understanding of traits and range si$es

".1 Abstract
0n im5ortantA unresolved Luestion in macroecolog9 is to understand the immense inters5ecific variation in geogra5hic range sizes- While s5eciesP traits such as fecundit9 or bod9 size are thought to affect range sizesA a general understanding on how multi5le traits Iointl9 influence them is missing- ?ereA we test the influence of a multitude of s5eciesP traits on range sizes of Euro5ean 5asserine birds in order to better understand 5ossible mechanisms behind macroecological relationshi5s- We evaluated the effect of life-histor9 traits *fecundit9A dis5ersal abilit9,A ecological traits *habitat nicheA diet nicheA migrator9 behaviourA migrator9 fleNibilit9, and a mor5hological trait *bod9 size, on global range sizes of )E@ Euro5ean 5asserines- We identified h95otheses from the literature relating traits to range size and used 5ath anal9sis to test them- Fecundit9A dis5ersal abilit9A habitat niche breadth and diet niche 5osition had a direct 5ositive effect on range size- ?abitat niche breadth also had an indirect 5ositive effect via fecundit9- "igrator9 behaviour had an indirect 5ositive effect via dis5ersal abilit9- Bod9 size had a strong 5ositive direct effect which was reduced b9 negative indirect effects via several other traits- Geogra5hic range sizes of Euro5ean 5asserines were influenced b9 life-histor9 traits *fecundit9 and dis5ersal abilit9,A ecological traits *habitat niche breadthA diet niche 5osition and migrator9 behaviour, and b9 bod9 size- &raits influenced range size both directl9 and indirectl9- Bod9 size effects were 5articularl9 com5leN with 5ositive and negative effects acting over different 5athwa9s- We show that it is necessar9 to disentangle the direct and indirect influence of multi5le traits on range size to better elucidate the mechanisms that generate macroecological relationshi5s-

".2 0ntroduction
=ne of the fundamental traits of a s5ecies is the size of its geogra5hic range *Brown et al.A )88EH Gaston K FullerA '((8,- >ange size influences 5atterns of s5ecies diversit9 *Jetz K >ahbekA '(('H %oberon K 4eballosA '()), and s5ecies with small ranges have a higher eNtinction 5robabilit9 *BrownA )88@H $ee K JetzA '()), making range size one of the most im5ortant criteria for classif9ing the threat status of a s5ecies *# 4N >ed $ist classificationA # 4NA '((),#nters5ecific range size variation can cover several orders of magnitudeA even between close relatives *Brown et al.A )88E,- CetA our understanding of the mechanisms that are )E

" #owards a more mechanistic understanding of traits and range si$es res5onsible for this immense variation is limited *$ester et al.A '((<,- 0mong the most im5ortant factors that influence range size are s5eciesP traits- $ife-histor9 traitsA such as birth rate and dis5ersal abilit9 *?olt et al.A )88<H B2hning-Gaese et al.A '((E,A ecological traitsA 5articularl9 habitat niche and diet niche *BrownA )8:BH Gregor9 K GastonA '(((, as well as migrator9 behaviour *Gaston K BlackburnA )88E, and mor5hological traits such as bod9 size *BrownA )88@, have been shown to influence range sizesBrown et al. *)88E, em5hasised that several traits might influence range size simultaneousl9 in a com5leN wa9A through direct as well as indirect effects- For eNam5leA large bod9 size in birds ma9 directl9 increase range size because of bod9 size-de5endent s5atial interactions with resources and the environment *BrownA )8:B,- =n the other handA large-bodied s5ecies have lower fecundit9 which might lead to reduced range size *Gaston et al.A )88<H B2hning-Gaese et al.A '(((,- Finall9A large-bodied birds are less likel9 to be migrator9 *?edenstr2mA '((:, and thus might have lower dis5ersal abilit9 and hence smaller ranges com5ared to small-bodied birds *?olt et al.A )88<H Dawideit et al.A '((8,- &he relative im5ortance of traits can onl9 be assessed b9 multi5le anal9ses- 0lsoA the a55arent statistical significance of traits in individual tests ma9 be caused b9 correlations with otherA non-tested traits *%hi5le9A '(((,- NonethelessA in most studiesA traits have been tested individuall9For a more mechanistic understanding of the relationshi5s between life-histor9 traitsA ecological traitsA mor5hological traits and range sizeA as man9 traits as 5ossible should be tested simultaneousl9 and interactions among traits should be considered- =ne o5tion for testing the direct and indirect effects of traits on a res5onse variable is structural eLuation modellingA in 5articular 5ath anal9sis *"itchellA )88'H %hi5le9A '(((,- %uch modelsA while based on eNamination of correlational 5atterns *%hi5le9A '(((,A have been used successfull9 to evaluate factors that directl9 or indirectl9 influence macroecological 5atterns such as s5ecies richness *1issling et al.A '((<H Qian K 1isslingA '()(, or eNtinction risk *$ee K JetzA '()),?ereA we tested the most com5rehensive set of traits to date for their direct and indirect effects on the geogra5hic range sizes of birds- We used birds in this anal9sis because traits and range sizes of birds are well documented and a number of studies have alread9 tested individual relationshi5s on which we can base a 5riori h95otheses *e-g- Gaston et al.A )88<H B2hning-Gaese et al.A '((EH ?urlbert K WhiteA '((<,- We incor5orated traits reflecting the life histor9 *annual fecundit9A dis5ersal abilit9,A ecolog9 *habitat niche breadthA diet niche breadth and 5ositionA migrator9 behaviour and fleNibilit9, and mor5holog9 *bod9 size, of

)<

" #owards a more mechanistic understanding of traits and range si$es birds into our anal9sisWe identified the following a 5riori h95otheses in the literature about the 5otential mechanistic relationshi5s between these traits and ranges sizes of birds *see methods for details,. ?igh annual fecundit9 and high dis5ersal abilit9 lead to larger range sizes *Blackburn et al.A '((EH B2hning-Gaese et al.A '((E,- Broader habitat niches and broader diet niches cause larger range sizes both directl9 and also indirectl9 via increasing annual fecundit9 *BrownA )8:BH ?urlbert K WhiteA '((<,- %5ecies with a diet niche 5osition at higher tro5hic levels have smaller ranges *GastonA )88B,- &here is a direct effect of migrator9 behaviour on range sizesA for which both negative and 5ositive relationshi5s have been 5ostulated in the literature *Gaston K BlackburnA )88EH BenschA )888,- "igrator9 behaviour also influences range size indirectl9 via dis5ersal abilit9 because migrants tend to be better dis5ersers which in turn increases range size *Baldwin et al.A '()(,- %5ecies with higher migrator9 fleNibilit9 have larger ranges *1eitt et al.A '((),- $arger bod9 size directl9 leads to larger range size *BrownA )8:B,- 0dditionall9A bod9 size is linked indirectl9 to range size via migrator9 behaviourA with large bodied birds being less freLuentl9 migrator9A and via annual fecundit9A which is lower in large-bodied bird s5ecies *B2hning-Gaese et al.A '(((H ?edenstr2mA '((:,We incor5orated these h95otheses into a 5ath model and estimated the strength of the direct and indirect effects of s5ecies? traits on range sizes-

"." 1ethods
".".1 -tud8 s!ecies and areaA geogra!hic range sizes
We anal9sed the relationshi5 between traits and global breeding range sizes of )E@ Euro5ean 5asserine bird s5ecies *see 055endiN ),- &he anal9sis was restricted to 5asserines because the9 share a similar bod9 5lan and because dis5ersal abilit9 can be Luantified com5arativel9 easil9 from mor5holog9 *Dawideit et al.A '((8,Global breeding range sizes of birds were calculated using data from a com5rehensive global geogra5hic bird range database at a resolution of )R S )R *version +(T(ET'((8,- &he geogra5hic breeding range of each s5ecies was ma55ed following the a55roach described in >ahbek and Graves *'(((A '((),- "a5s re5resent a conservative eNtent-of-occurrence based on museum s5ecimensA 5ublished sight records and s5atial distribution of habitatsA which have subseLuentl9 been validated b9 ornithological eN5erts- >ange size was Luantified as the sum

):

" #owards a more mechanistic understanding of traits and range si$es of the areas of all grid cells a s5ecies occu5ied- We considered onl9 the land surface area of grid cells in sLuare kilometres after a55l9ing a Behrmann global eLual-area 5roIection-

".".2 -!ecies traits


&he following traits and their 5otential relationshi5s with range size have been derived from the literature- We use the traits and their relationshi5s among each other and to range size to define a 5riori h95otheses on 5aths in the 5ath diagrams *Fig- +-),-

HabNb DietNb
0.0&'
0.2$&! 0.20&!!

0.2&#!!

0.0#'

DietPos Body
0.29 !!! 0.$0%!!!

0.22'!!

Fe()nd

0.&90!

Range size

MigBeh
0."#9!!!

0.009

Dispers MigFlex

0."0#!!! 0.0' 0."&2!!!

5igure ".1+ /ath diagram for 5ath model relating avian traits to global range size *NF# U (-::A GF# U (-8@A n U )E@,- /ath coefficients and significance levels. V ! W (-(@H VV ! W (-()H VVV ! W (-(()- Bod9 U log*bod9 mass,A ?abNb U habitat niche breadthA DietNb U diet niche breadthA Diet/os U diet niche 5ositionA "igBeh U migrator9 behaviourA "igFleN U migrator9 fleNibilit9A Fecund U log*annual fecundit9,A Dis5ers U log*dis5ersal abilit9,-

%nnual fecundity
?igh annual fecundit9 *e-g- large clutchesA man9 broods 5er 9ear, ma9 cause large geogra5hic ranges as it could lead to high local abundances *Blackburn et al.A '((E, which are often correlated with large range sizes *BrownA )8:BH Blackburn et al.A )88EH Gaston et al.A )88<H

)8

" #owards a more mechanistic understanding of traits and range si$es Borregaard K >ahbekA '()(H direct 5ath from annual fecundit9 to range size in Fig- +-),- We Luantified annual fecundit9 as the 5roduct of clutch size times the number of clutches 5er 9ear using data from Ehrlich et al. *)88B,- For Sturnus unicolor the number of clutches 5er 9ear was taken from Birds of the Western /alearctic interactive *BW/iA '((E, because data were missing in Ehrlich et al. *)88B,- For anal9sisA fecundit9 was log)(-transformed-

&ispersal ability
>ange fillingA i-e- the ratio of realised to 5otential range sizeA can be limitedA amongst other factorsA b9 dis5ersal abilit9 *%venning K %kovA '((B,- 0ccordingl9A a 5ositive relationshi5 between dis5ersal abilit9 and geogra5hic range size has been shown in several studies *e-gDennis et al.A '(((H B2hning-Gaese et al.A '((EH direct 5ath in Fig- +-),We Luantified dis5ersal abilit9 as the Luotient of 1i55Ps distance *ti5 of the first 5rimar9 to ti5 of the wing, and bill de5th *measured at the 5roNimate edge of the nostrils,- &his measure has been shown to be the best mor5hological 5redictor of dis5ersal abilit9 in Euro5ean 5asserines *Dawideit et al.A '((8,- We aimed to measure 1i55Ps distance and bill de5th for at least eight museum s5ecimens 5er s5ecies- ?oweverA this was not alwa9s 5ossible *mean. <-'E s5ecimensH range. );)' s5ecimens,- We took care to select adultA non-moulting s5ecimens from localities as close as 5ossible to the centre of the Euro5ean geogra5hic range and whose time of death was between 05ril and Jul9 to avoid measuring wintering individuals- #f fewer than eight suitable individuals were availableA we relaNed the criteria on localit9 and time of death- For s5ecies that %vensson *)88', describes as seNuall9 dimor5hic we measuredA if 5ossibleA four individuals 5er seN- For s5ecies with more than one subs5ecies in Euro5eA we measured the nominate s5eciesA as it is usuall9 the most wides5read- For s5ecies where subs5ecies had geogra5hic ranges of similar sizeA we took measures of individuals from both and calculated the mean- 0ll measurements were taken b9 the same 5erson *?- 1orntheuer,- When calculating averages across s5ecimensA we first calculated the Luotient of log)(*1i55Ps distance, and log)(*bill de5th, for each individual and then averaged over individuals-

'abitat niche breadth


?abitat niche breadth ma9 be 5ositivel9 related to range size *e-g- ?urlbert K WhiteA '((<H 4arrascal et al.A '((:, as s5ecies that tolerate a wider range of conditions are able to colonise larger geogra5hic areas *BrownA )8:BH Gaston et al.A )88<H direct 5ath in Fig- +-),- ?abitat '(

" #owards a more mechanistic understanding of traits and range si$es niche breadth is also eN5ected to increase fecundit9 and hence indirectl9 increase range size *indirect 5ath from habitat niche breadth to fecundit9 in Fig- +-), as s5ecies that are able to live under a wide variet9 of conditions and use a broad range of resources should also be able to obtain more resources locall9 and raise more 9oung *BrownA )8:BH Gaston et al.A )88<,&o Luantif9 habitat niche breadthA the habitat use of a s5ecies was converted to a habitat gradient from closed forest to o5en countr9 with values of ) *closed forest,A ' *o5en forest,A + *forest edge,A B *orchardsA gardens,A @ *shrub land,A E *o5en countr9 with single trees or shrubsA e-g- agricultural land with hedgerows,A and < *o5en countr9 without trees or shrubsA e-g- structurall9 sim5le arable land, using data from Ehrlich et al. *)88B,- 0 s5ecies was assigned u5 to three different values along this habitat gradient *B2hning-Gaese K =berrathA '((+,- ?abitat niche breadth was calculated as the difference between the maNimum and minimum value-

&iet niche breadth


0nalogous to habitat niche breadthA s5ecies which use a broad range of food sources might be more wides5read than more s5ecialised s5ecies *direct 5ath in Fig- +-),- #n additionA a broad diet niche ma9 lead to increased fecundit9A causing an indirect 5ositive effect of diet niche breadth on range size *BrownA )8:BH indirect 5ath from diet niche breadth to annual fecundit9A Fig- +-),- Diet niche breadth was Luantified b9 taking into account the range of utilised food sources- We classified all s5ecies as herbivorousA insectivorous or omnivorous using data from Ehrlich et al. *)88BH B2hning-Gaese et al.A '(((,- We assigned s5ecies that were either herbivorous or insectivorous a diet niche breadth of ) *+: s5ecies,A s5ecies that were herbivorous and insectivorous a value of ' *)'( s5ecies,A and omnivorous s5ecies a value of + *< s5ecies,-

&iet niche position


We used the tro5hic level of a s5ecies as a measure of its diet niche 5osition- %5ecies at higher tro5hic levels are faced with lower food biomass andA conseLuentl9A might have lower local abundance and hence smaller range sizes than s5ecies at lower tro5hic levels *GastonA )88BH direct 5ath in Fig- +-),- We defined the diet niche 5osition of herbivorous s5ecies as ) *'8 s5ecies,A of s5ecies that were herbivorous and insectivorous or that were omnivorous as ' *+@ s5ecies,A and of insectivorous s5ecies as + *)() s5ecies,- Note that no true carnivores *vertebrate-eating s5ecies, were included in this stud9- #nclusion of these s5ecies might give ')

" #owards a more mechanistic understanding of traits and range si$es results that differ from the above h95othesisA as man9 carnivorous s5ecies *es5eciall9 birds of 5re9, a55ear to have rather large geogra5hic ranges *del ?o9o et al.A )88B,-

(igratory behaviour
ELuivocal results have been found for the effect of migrator9 behaviour on ranges size- =n the one handA migrator9 birds have been shown to have smaller geogra5hic ranges than nonmigrantsA 5otentiall9 because migrants are limited in eNtending their geogra5hic ranges along a longitudinal aNis within the ?olarctic due to constraints caused b9 their migrator9 behaviour *B2hning-Gaese et al.A )88:H BenschA )888,- =n the other handA long distance migrants have been shown to have larger geogra5hic ranges than sedentar9 birds in 0nseriformes *Gaston K BlackburnA )88E,- ?ere we tested for a 5otential direct effect of migrator9 behaviourA as well as for an indirect effect via dis5ersal abilit9 *Fig- +-), because migrator9 birds show ecomor5hological ada5tations to long-distance flight also resulting in better dis5ersal abilit9 *Winkler K $eislerA )88'H Dawideit et al.A '((8A Baldwin et al., '()(,- We classified the migrator9 behaviour of a s5ecies as ) *residentsA @) s5ecies,A ' *short-distance migrantsA with the centre of their non-breeding grounds south of the breeding grounds but north of the %aharaA @) s5ecies,A or + *long-distance migrants with the centre of their wintering grounds south of the %aharaA E+ s5eciesH B2hning-Gaese et al.A '(((,-

(igratory fle)ibility
%5ecies with fleNible migrator9 behaviour are more successful invaders than those with a fiNed migrator9 5rogramme *%ol K $efebvreA '(((,- ?igher invasion success might lead to larger geogra5hic ranges *1eitt et al.A '(()H direct 5ath from migrator9 fleNibilit9 to range size in Fig- +-),- For migrator9 fleNibilit9 we differentiated between s5ecies with an invariable migrator9 behaviour *value (A residents or long-distance migrantsA ))B s5ecies, and s5ecies with a fleNible migrator9 behaviour *value )A short-distance migrantsA @) s5ecies,- >esident birds and long-distance migrants were classified ver9 conservativel9 *B2hning-Gaese et al.A '(((, and included onl9 s5ecies with no intras5ecific variation in migrator9 behaviour within Euro5eA conseLuentl9 defining all s5ecies with intras5ecific variation in migrator9 behaviour as short-distance migrant-

''

" #owards a more mechanistic understanding of traits and range si$es

*ody si$e
$arger s5ecies interact with their environment at larger s5atial scales than smaller s5ecies&husA smaller organisms are able to attain higher densities in small rangesA while larger ones tend to have less denseA more widel9 distributed 5o5ulations *BrownA )8:BH )88@H direct 5ath in Fig- +-),- We also eN5ected s5ecies with large bod9 size to be less migrator9 *indirect 5ath to migrator9 behaviourH Fig- +-), because large birds ma9 need more time to raise their 9oung and to moult and hence have less time for migration and because bod9 size constrains flight s5eed during fla55ing flightA the most common flight st9le of 5asserines *?edenstr2mA '((:,FurthermoreA large bird s5ecies tend to have low fecundit9 *B2hning-Gaese et al.A '(((H indirect 5ath to fecundit9 in Fig- +-),- We used bod9 mass as a measure of bod9 size *4larkH )8<8,- Data were taken from BW/i *BW/iA '((E, and were log)(-transformed-

"."." -tatistical anal8ses !ath analyses


#n 5ath anal9sis *"itchellA )88'H %hi5le9A '(((,A su55osed mechanistic relationshi5s between variables are delineated in a 5ath diagram *Fig- +-),- Direct effects are measured b9 the standardised 5artial regression coefficient *in the following 5ath coefficient, for the direct link between a 5redictor variable and a res5onse variable- #ndirect effects are calculated b9 multi5l9ing the 5ath coefficients along a 5ath between a 5redictor and a res5onse variableA and then adding these 5roducts for all 5ossible 5aths between the twoA eNcluding the direct effect *"itchellA )88',- We used the a 5riori h95otheses described above to define 5aths between s5eciesP traits and geogra5hic range size *Fig- +-),- We allowed correlations between 5redictors if the9 were significantl9 correlated *XrX (-)@H ! W (-(@A n U )E@, and if there was no information in the literature on traits and range size regarding the 5otential direction and cause of the correlation *%hi5le9A '(((,- We thus fitted correlations between diet niche breadth and *i, dis5ersal abilit9A *ii, migrator9 behaviourA *iii, habitat niche breadthA *iv, diet niche 5osition and *v, bod9 weightA between migrator9 fleNibilit9 and *i, fecundit9A *ii, habitat niche breadthA between diet niche 5osition and *i, dis5ersal abilit9A *ii, migrator9 behaviourA as well as between habitat niche breadth and dis5ersal abilit9 *all X rX W (-@@,- For clarit9?s sakeA these correlations were omitted from Fig- +-)- #n additionA we eNamined generalised variance inflation factors from a linear model containing all 5redictors to assess the 5otential effect of

'+

" #owards a more mechanistic understanding of traits and range si$es multicollinearit9 on 5arameter estimates- &he 5ath model was evaluated using the normed fit indeN *NF#, and the goodness-of-fit indeN *GF#, *Bentler K Bonett )8:(A 0rbuckleA '((:,/ath anal9ses were calculated using 0"=% *0rbuckleA '((:,-

!hylogenetic relatedness
#ndividual s5ecies do not necessaril9 re5resent inde5endent data 5ointsA as closel9 related s5ecies tend to have more similar traits than distantl9 related s5ecies *?arve9 K /agelA )88),&o check for 5otential statistical issues arising from 5h9logenetic non-inde5endenceA we tested the residuals from a multi5le regression of range size against all s5ecies trait variables *corres5onding to the direct effects in the 5ath model in Fig- +-), for 5h9logenetic autocorrelation- We used a 5ublished su5ertree for Euro5ean birds *&huiller et al.A '()), which contains all of our stud9 s5ecies eNce5t for Sitta whiteheadii and %nthus petrosus- 0ll anal9ses were conducted in > '-)'-' *> Develo5ment 4ore &eamA '()),- We tested for 5h9logenetic signal in the residuals using the 0bouheif test *0bouheifA )888, with 888 randomisations as im5lemented in the 5ackage ade5h9lo *Jombart et al.A '()(, and b9 calculating /agelPs YA a maNimum-likelihood based measure of 5h9logenetic signal */agelA )88<,A and testing for a significant difference to a lambda of zero *no 5h9logenetic structure,A as im5lemented in the 5ackage 40#4> *FreckletonA '((8,-

".$ *esults
&he 5ath model *n U )E@ s5ecies, adeLuatel9 described the data structure *NF# U (-::A GF# U (-8@,A 9et the variables included in the model eN5lained onl9 > ' U (-'@ of the inters5ecific variation in global range size- %5ecies with higher fecundit9A better dis5ersal abilit9A broader habitat nichesA lower tro5hic level and larger bod9 size had larger ranges *Fig- +-'ae,?abitat niche breadth had a 5ositive effect on annual fecundit9 while bod9 size had a negative effect- Bod9 size had a negative effect on migrator9 behaviour and migrator9 behaviour 5ositivel9 affected dis5ersal abilit9 *Fig- +-),- Generalised variance inflation factors for all 5redictors were smaller than '-<A indicating that 5arameter estimates were not affected b9 multicollinearit9&he standardised total effect size of each trait on range size could be s5lit into direct and indirect effects *&able +-),- &he strong total effect of habitat niche breadth on range size was mostl9 caused b9 a direct 5ositive effect on range size and onl9 a weak indirect 5ositive effect

'B

" #owards a more mechanistic understanding of traits and range si$es through annual fecundit9- #n contrastA the total effect of migrator9 behaviour on range size was driven b9 a stronger indirect effect through dis5ersal abilit9A and a weak direct effect- &he total effect of bod9 size on range size was com5leNH its strong 5ositive direct effect was counteracted slightl9 b9 two indirect negative effectsA one via fecundit9 and the other via migrator9 behaviour and dis5ersal abilit9 *Fig- +-),A but still resulted in a significant 5ositive total effectBoth tests on the 5otential influence of 5h9logenetic relatedness confirmed that there were no significant 5h9logenetic signals in the multi5le regression residuals *0bouheif test. ! U (-(@<H $ikelihood ratio test for lambda U (. ! U ),A indicating anal9ses of the data with non5h9logenetic methods were a55ro5riate- ?enceA our results were not affected b9 the 5h9logenetic relatedness of the s5ecies=able ".1+ %tandardised total effectsA direct effects and indirect effects of bird traits on global range sizes of )E@ Euro5ean 5asserine s5ecies- &he correlation between 5redictor and res5onse variableA the total effectA can be s5lit u5 into direct effects and indirect effects via other de5endent variables- Direct effects are measured b9 the standardised 5artial regression coefficients between a 5redictor variable and a res5onse variable *i-e- the direct link,- #ndirect effects are calculated b9 adding the 5roducts of all 5ath coefficients over all 5aths between a 5redictor and a res5onse variableA eNcluding the direct effect *"itchellA )88',Bird traits Fecundit9 Dis5ersal abilit9 ?abitat niche breadth Diet niche breadth Diet niche 5osition "igrator9 behaviour "igrator9 fleNibilit9 Bod9 size &otal effect (-)8( (-B(@ (-'@+ -(-(E( -(-''< (-)8@ -(-(<: (-'8@ Direct effect (-)8( (-B(@ (-')@ -(-(@< -(-''< (-((8 -(-(<: (-B)' #ndirect effect N0 N0 (-(+: -(-((+ N0 (-):E N0 -(-))<

'@

" #owards a more mechanistic understanding of traits and range si$es

5igure ".2+ $everage 5lots after %all *)88(, of bird traits with a significant direct effect on global range size. *a, log*annual fecundit9,A *b, dis5ersal abilit9A *c, habitat niche breadthA *d, diet niche 5ositionA *e, log*bod9 mass,A calculated from a multi5le regression-

'E

" #owards a more mechanistic understanding of traits and range si$es

".' 3iscussion
We tested the direct and indirect effects of a multitude of traits on the global breeding range sizes of Euro5ean 5asserine birds- /ath anal9ses revealed direct effects of fecundit9A dis5ersal abilit9A habitat niche breadthA diet niche 5osition and bod9 sizeA as well as indirect effects of habitat niche breadthA migrator9 behaviour and bod9 size on global range sizes%5ecies which raised more offs5ring 5er 9ear had larger geogra5hic ranges *&able +-)A Fig- +-),- &hese results confirmed earlier studies that found 5ositive relationshi5s between fecundit9 and range size *e-g- Blackburn et al.A )88EA Gaston et al.A )88<,- ?igh fecundit9 might be linked to large range sizes through higher local abundance *BrownA )8:BH Blackburn et al.A '((E,- 0s a conseLuenceA 5o5ulations in sink habitats might be 6rescued7 through regular immigration from source habitats with the result thatA on averageA a larger 5ro5ortion of habitat 5atches might be occu5ied *GastonA '((+,Better dis5ersers had larger geogra5hic ranges- /oor dis5ersal abilit9 ma9 lead to a larger 5ro5ortion of 5otentiall9 suitable habitat remaining unoccu5ied *$ester et al., '((<,- 0lsoA good dis5ersers should be able to sustain sink 5o5ulations at longer distances to source 5o5ulations than 5oor dis5ersers- Even for mobile s5ecies such as birds and when multi5le traits are tested simultaneousl9A dis5ersal abilit9 has an influence on range size *B2hningGaese et al.A '((E,- &his suggests that not onl9 treesA am5hibiansA and re5tiles *%venning K %kovA '((BH 0raOIo et al.A '((:, but also birds might not have full9 recolonised their 5otential geogra5hic range since the last glacial 5eriod '(A((( 9ears ago- #t a55ears thatA in the face of anthro5ogenic climate changeA at least some bird s5ecies might not be mobile enough to track s5atial shifts in their climate niche *Devictor et al. '((:,0s shown b9 other studies *?urlbert K WhiteA '((<H 4arrascal et al.A '((:,A habitat niche breadth had a 5ositive direct effect on range sizeA reflecting that the habitat niche directl9 constrains the area which can be colonised b9 a s5ecies- %5ecies with a broad habitat niche also had higher fecundit9A resulting in an additional 5ositive indirect effect of habitat niche breadth on range size- %5ecies with broader habitat niches should find the o5timal conditions for re5roduction more freLuentl9A achieving on average higher fecundit9 in a given area *BrownA )8:BH Gaston et al.A )88<,4ontrar9 to habitat niche breadthA diet niche breadth did not have an effect on range sizeDifferent food sources can occur side b9 side in the same siteA while habitat t95es cannot&hereforeA it is not sur5rising that diet niche breadth is less limiting for a s5ecies ? range size '<

" #owards a more mechanistic understanding of traits and range si$es than habitat niche breadth- FurthermoreA in regions intensivel9 used and modified b9 humansA the abilit9 to use anthro5ogenic food sources might be more im5ortant in determining range size than diet niche breadth *B2hning-Gaese K =berrathA '((),- Finall9A our classification of diet niche breadth was rather broad and data taking the relative consum5tion of finer classified diet items into account might give different resultsBird s5ecies of higher tro5hic level had smaller geogra5hic ranges- &his suggests that herbivorous birds indeed have more food biomass available than insectivores and are hence able to reach higher abundances and find enough food to sustain their 5o5ulations in more 5laces than insectivores- We detected a 5ositive indirect effect of migrator9 behaviour on range sizeA mediated via dis5ersal abilit9 while migrator9 fleNibilit9 had no effect- &his link between migrator9 behaviour and dis5ersal abilit9 has been described 5reviousl9 for /asserines *Winkler and $eislerA )88'H Dawideit et al.A '((8,B9 combining the traits in a 5ath model it was 5ossible to assess direct and indirect effects on range sizes- &he benefit of such an anal9sis was best illustrated for habitat niche breadthA which had both direct and indirect effects on range sizeA b9 the 5resence of an indirect effect of migrator9 behaviour on range size in the absence of a significant direct effect and b9 bod9 size showing direct and indirect effects influencing range size in o55osite directions- &he strong 5ositive direct effect of bod9 size on range size was moderated b9 two indirect negative effectsA one via fecundit9 and the other via migrator9 behaviour and dis5ersal abilit9&he relationshi5 between bod9 size and range size has alwa9s been a matter of debate with 5ublished 5ositive *4arrascalA '((:,A negative *GlazierA )8:(,A triangular *Brown K "aurerA )8:<, and non-significant relationshi5s *DirkkalaA )88+,- &he 5resent stud9 demonstrates that a 5otential reason for these com5leN 5atterns might be the heterogeneit9 in mechanisms b9 which bod9 size affects range size- De5ending on the s5atial scale of the anal9sisA the set of s5ecies anal9sed and other traits included in the stud9A this might result in 5ositiveA negative or no total effect of bod9 size on range sizeBoth life-histor9 traitsA three out of five ecological traits and one mor5hological trait showed significant direct or indirect effects on range size- >ange size thus de5ended on the life histor9A ecolog9 and mor5holog9 of s5ecies and ecological and mor5hological traits acted both via direct and indirect 5athwa9s- &his underlines that range size is concurrentl9 influenced b9 several traits via a number of differentA simultaneousl9 acting mechanismsGiven that our 5ath model was able to account for onl9 a fraction of the total variabilit9 in

':

" #owards a more mechanistic understanding of traits and range si$es range sizesA it is clear that im5ortant 5redictors were lacking from the model- We could not test three 5otentiall9 im5ortant s5ecies? traits in the model due to insufficient data. ?igh relative brain size can influence the success of a s5ecies in a novel environment and the 5robabilit9 of eN5loiting novel food sources *%ol et al.A '((@, and ma9 hence lead to large geogra5hic ranges- 0nother trait that might influence range size is the 5osition of a s5ecies ? habitat niche. %5ecies that 5refer wides5read habitats have larger geogra5hic ranges than s5ecies 5referring rare habitat t95es *Gregor9 K GastonA '(((H ?urlbert K WhiteA '((<,Evolutionar9 age ma9 also affect avian range sizeA with ranges increasing ra5idl9 after s5eciation and then graduall9 declining again *Webb K GastonA '(((,- FurthermoreA since our 5ath model focused onl9 on s5ecies? traitsA it does not incor5orate a number of im5ortant factors which might also influence geogra5hic range size. the climatic and geologic histor9 of a s5ecies? habitatA the histor9 of a s5ecies? distribution in s5ace or biotic interactions with other s5ecies such as mutualism and 5athogens *=rme et al.A '((EH %oberon K 4eballosA '()),#n this stud9A we demonstrated how multi5leA interacting traits have direct and indirect effects on range size- While our results a55l9 to 5asserinesA other bird grou5s ma9 show different relationshi5s between s5ecies? traits and range size- Birds of 5re9A for eNam5leA have a high tro5hic level but freLuentl9 ver9 large ranges- $ooking be9ond birdsA it might be worthwhile to carr9 out similar studies with other grou5s of organisms for which similarl9 good data on traits and range sizes eNistA e-g- mammalsA am5hibiansA re5tilesA butterflies or 5lants- For eNam5leA it has long been noted thatA on averageA birds generall9 have larger geogra5hic ranges than mammals *0ndersonA )8:B,A which might be eN5lained b9 different direct and indirect effects of traits on the range sizes of the two grou5s- We eN5ect that for other grou5s of organismsA other traits might 5rove to be im5ortant- For less mobile s5eciesA e-g- re5tiles or 5lantsA one ma9 eN5ect dis5ersal abilit9 to have an even stronger effect than for birdsA whereas for butterfliesA diet niche breadth *of the larval stages, might 5otentiall9 5rove to be essential- While our 5ath model re5resents a good h95othesis for how the s5ecies ? traits we measured influence range sizesA it is clear that those traits cannot full9 eN5lain inters5ecific range size variation- We suggest that it is necessar9 to disentangle the direct and indirect influence of multi5le other s5ecies traits and of factors related to the biogeogra5hical and evolutionar9 histor9 of s5ecies in order to better elucidate the mechanisms that generate macroecological range size 5atterns-

'8

4?0/&E> B

4="/E&#&#=N 0ND D#%/E>%0$ 0B#$#&C


#N&E>04& &= DE&E>"#NE GE=G>0/?#4 >0NGE% =F B#>D%
$ 6>1B<=0=0>( A(3 30-B<*-AC A&0C0=D 0(=<*A6=

Sylvia curruca

=> 3<=<*10(< ,<>,*ABH06 *A(,<- >5 &0*3-

%ubmitted to an international Iournal of ecolog9 as. $aubeA #-A GrahamA 4- ?- K B2hning-GaeseA 1- *submitted, 4om5etition and dis5ersal abilit9 interact to determine geogra5hic ranges of birds-

+)

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds

$.1 Abstract
nderstanding the factors that influence the geogra5hic ranges of s5ecies remains a challenge in ecolog9 and evolutionar9 biolog9- #n 5articularA little consensus eNists as to whether geogra5hic ranges of s5ecies are determined b9 biotic interactions such as inters5ecific com5etition- We evaluated how com5etitionA dis5ersal abilit9A taNon age and habitat shift since the last glacial maNimum influenced the eNtent to which s5ecies in the bird genus Sylvia occur in all areas 5redicted as environmentall9 suitable *i-e- range filling,We Luantified range filling in the bird genus Sylvia using boosted regression trees and ridge regression- We tested for effects of intrageneric com5etitionA dis5ersal abilit9A taNon age and habitat shift since the last glacial maNimum on range filling using multi5le regression- We eN5lore several wa9s to Luantif9 5otential signals of com5etition at the range scale to reflect different h95otheses about how local com5etition might scale u5 to influence large-scale range d9namicsSylvia warblers with higher dis5ersal abilit9 showed higher range fillingA but onl9 if com5etition in less suitable habitats within their 5otential range was low- &aNon age and habitat shift since the last glacial maNimum had no consistent effectWe show that Sylvia ranges are likel9 sha5ed b9 the simultaneousA interactive effect of both com5etition and dis5ersal abilit9- #f biotic interactionsA like com5etitionA generall9 influence the abilit9 of s5ecies to colonise and occu59 habitat at the continental scaleA 5redicting the im5act of climate change on biodiversit9 will be challenging-

$.2 0ntroduction
#dentification of the factors that determine s5eciesP geogra5hic ranges has long fascinated ecologistsA biogeogra5hers and evolutionar9 biologists *Dobzhansk9A )8@(H "ac0rthurA )8<'H GastonA '((+,- While a series of abiotic and biotic factors acting across s5atial and tem5oral scales can influence rangesA knowledge of the s5ecific mechanisms that sha5e ranges has long remained elusive- &his is 5artl9 because research has generall9 focused on onl9 one or two factors at a time- #lluminating the relative im5ortance of and the interactions among different determinants reLuires integrative anal9sis *Brooker et al.A '((<H "unguia et al.A '((:,- ?ere we evaluate in the bird genus Sylvia how biotic interactionsA dis5ersal abilit9A taNon age and current and historic climate conditions influence the eNtent to which a s5ecies occurs in all +'

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds environmentall9 suitable habitat- We show that ranges are likel9 sha5ed b9 the simultaneousA interactive effect of both com5etition and dis5ersal abilit9 and that biotic interactions do influence biogeogra5hic 5atterns at the continental scaleDetermining if all environmentall9 suitable habitat is occu5ied b9 a s5ecies and wh9 such habitat is not occu5ied can 9ield new insights into the 5rocesses that sha5e geogra5hic ranges */ulliamA '(((H %oberonA '((<,- #n the conteNt of this stud9A we define environmentall9 suitable habitat as areas with environmental conditions *such as tem5eratureA 5reci5itation and vegetation structure, that are similar to the area where the s5ecies is currentl9 5resent- &he sum of all such environmentall9 suitable areas is the 5otential range *GastonA '((+H %oberonA '((<,A which ma9 or ma9 not be occu5ied b9 the focal s5ecies- &he ratio of actual range size to the size of this 5otential range has been 5reviousl9 defined as range filling *GastonA '((+H %venning K %kovA '((B,- %5ecies distribution models have been used to estimate 5otential ranges and eNamine 5atterns of range filling in wood9 5lants *%venning K %kovA '((BH %churrA '((<H /aul et al.A '((8,A birds *Graham et al.A '()(, and mammals *"unguia et al.A '((:,&he main drivers of range filling at large s5atial scalesA i-e- across whole continentsA are thought to be biotic interactionsA dis5ersal abilit9A taNon age and historic climate change *Brooker et al.A '((<H "unguia et al.A '((:,Biotic interactions have long been considered a 5otential force in setting range limits *Dobzhansk9A )8@(H JaegerA )8<)H Bullock et al.A '(((H 4ase et al.A '((@H /rice K 1irk5atrickA '((8,- 4om5etition results in loss of energ9 through direct antagonistic interactions and restricted access to s5ace and food and thusA ma9 negativel9 affect individual re5roductive out5ut- &he aggregate effect of com5etition over man9 individuals ma9 limit 5o5ulation size and successful establishment in a given areaA 5otentiall9 to the eNtent where 5o5ulation growth is negative and the s5ecies is eNcluded b9 its com5etitors- 0s a resultA com5etition ma9 affect range filling through com5etitive eNclusion or inhibitor9 5riorit9 effects *Fukami et al.A '((@H /hil5ottA '()(,- "oreoverA the influence of com5etition on s5eciesP ranges is contingent on abiotic conditions *Dunson K &ravisA )88)H GZmez-"estre K &eIedoA '((',A with the im5act of com5etition t95icall9 being stronger at range edges where habitat is less suitable *4unningham et al.A '((8H "ooreA '((8,0ssessing the influence of com5etitionA and biotic interactions generall9A over large geogra5hic eNtents is fraught with 5ractical difficulties- Biotic interactions are events between individuals andA conseLuentl9A have mainl9 been documented on local to regional scales over

++

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds short time 5eriods *e-g- 4atch5oleA )8<:H >obinson K &erborghA )88@H $ovette K ?ochachkaA '((EA Jankowski et al., '()(,- #t remains a crucial Luestion if local-scale interactions translate to broader scale distributions and if their effects can be detected at a large scale *4onnor K BowersA )8:<A ?eikkinen et al.A '((<,- For instanceA Gotelli et al. *'()(, identified a broad scale signature of com5etition b9 showing large-scale s5atial segregation of congenerics and foraging guilds of Danish avifauna even when controlling for habitat availabilit9- &he9 suggested that this 5attern might be due to a combination of com5etitive interactionsA such as inters5ecific territorialit9A and cons5ecific attractionDis5ersal has long been regarded as a crucial 5rocess determining the colonization of environmentall9 suitable habitat *Bullock et al.A '((',- %5ecies with greater dis5ersal abilit9 should show higher range filling- When considering large s5atial eNtentsA such as continentsA range filling can also be influenced b9 constraints on the time available for dis5ersalA such that evolutionar9 9ounger s5ecies should show lower range filling than older s5ecies of similar dis5ersal abilit9 *GastonA '((+H B2hning-Gaese et al.A '((E,- "iNed su55ort has been obtained for taNon age and dis5ersal h95otheses *%venning K %kovA '((BH %churrA '((<H "unguia et al.A '((:H /aul et al.A '((8,- Finall9A variation in range filling ma9 be a result of historical 5rocesses related to climate and geogra5h9- %5ecies that were forced to track their 5referred habitat across continents due to climate fluctuations associated with glacial c9cles should have low range filling *%venning K %kovA '((BH "unguia et al.A '((:,- #nteractions among the four 5otential drivers of range filling are ecologicall9 5lausible- For instanceA dis5ersal abilit9A taNon age and habitat shift due to 5ast climate changeA ma9 have little effect if range filling is constrained b9 com5etition */rice K 1irk5atrickA '((8,A but such interactions have never been evaluated?ere we evaluate all four 5otential drivers of range filling and their interactions to understand which factors sha5e geogra5hic ranges- We also eN5lore different wa9s to ca5ture the 5otential large-scale signal of com5etition in order to understand how the influence of com5etition on range d9namics is contingent on habitat suitabilit9- &he grou5 that we use to illustrate the advantage of such a com5rehensive a55roach is the bird genus Sylvia- &he Sylvia warblers *sensu B2hning-Gaese et al.A '((+,A a genus of twent9-siN s5ecies of 5rimaril9 insectivorous 5asserinesA are ideall9 suited for studies of range filling due to the eNtensive evidence for local com5etitive interactions within the genus *e-g- 4od9 K WalterA )8<EH 4od9A )8<:H GarciaA )8:+H "artin K &hibaultA )88EH ElleA '((+H /ons et al.A '((:,-

+B

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds &he Sylvia warblers eNhibit great intrageneric ecological similarit9H all members of the genus are 5rimaril9 insectivorous foliage gleaners with a 5reference for deciduousA wood9 vegetationA in which the9 build sim5leA cu5-sha5ed nests *%hirihai et al-A '((),- &his shared ecolog9 is reflected in the intrageneric similarit9 of mor5hological traits such as the size and sha5e of billA feet and bod9 size *%hirihai et al.A '((),- 0ccordingl9A investigations of biotic interactions and habitat selection along habitat gradients find eNtensive overla5 in fine-scale habitat utilization and foraging nicheA leading to local-scale intrageneric com5etition that ma9 influence range filling- Field observations and removal eN5eriments demonstrate cooccurrence and interactions at the scale of individual territories *e-g- 4od9 K Walter )8<EA ElleA '((+A /ons et al. '((:,A inters5ecific territorialit9 *4od9 K WalterA )8<EH 4od9A )8<:A GarciaA )8:+,A shifts in habitat utilization in the 5resence of congenerics *GarciaA )8:+A "artin K &hibault )88E, and 5riorit9 effects with regard to timing of migration *GarciaA )8:+,- &husA Sylvia warblers 5rovide the o55ortunit9 to test if local-scale com5etition scales u5 to influence range filling over larger areas0nother advantage of using the genus Sylvia as a stud9 s9stem is the wealth of geogra5hicA ecologicalA mor5hologicalA and 5h9logenetic information available *%hirihai et al.A '(()H B2hning-Gaese et al.A '((+H '((E,- &he9 eNhibit large variation in range size *BAB((;'A8<@A((( km', and in the number of overla55ing ranges *one to nine s5eciesA FigB-)a,- %tudies on how Sylvia s5ecies var9 mor5hologicall9 across their range have resulted in detailed knowledge about range boundariesA 5articularl9 for the Eurasian breeding ranges *%hirihai et al.A '((),- Sylvia wing mor5holog9 gives an indication of the intrageneric variation in dis5ersal abilit9 *B2hning-Gaese et al.A '((EA Dawideit et al.A '((8,- Finall9A h9bridization seems not to be im5ortant for limiting range eN5ansion in the genus as there are no maIor h9brid zones *%hirihai et al.A '((),-

+@

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds

5igure $.1+ *a, %5ecies richness of Sylvia warblers based on their breeding ranges- =nl9 the '+ s5ecies included in the range filling anal9sis are shown- *b, >ange filling of selected Sylvia warblers- 0rrows indicate direction of increasing dis5ersal abilit9 and increasing intrageneric com5etitive 5ressure- Dark blue. observed rangeA light blue. 5otential range estimate based on )([ range ma5 conversion threshold and boosted regression trees- >F U range filling+E

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds Given the local-scale evidence for com5etition among members of the genusA we eN5ect to see lower range filling in Sylvia s5ecies where a large amount of 5otentiall9 suitable habitat is occu5ied b9 man9 congenerics- #f the effect of com5etition is inde5endent of habitat suitabilit9A we might eN5ect the 5resence of congeners throughout a s5eciesP range to im5act negativel9 on s5ecies 5ersistenceA which might result in a decreased ca5acit9 of that s5ecies to eN5and its range- When this is the caseA the mean number of congenerics in the total 5otential range should affect range filling- 0lternativel9A we might assume that the effect of com5etition on large-scale range d9namics is eNacerbated when habitat suitabilit9 for the focal s5ecies is low *4ase K &a5erA '(((,- $ess suitable habitat might increase resource limitation which in turn could limit 5o5ulation sizes and might also force s5ecies to use a broader range of resourcesA leading to increased niche overla5 and com5etition with congeners- #n this caseA the number of congenerics in highl9 suitable habitat could be irrelevant for testing the 5otential effects of com5etitionA since such habitat might allow coeNistence- &hereforeA we eN5ect the 5resence of congenerics to influence range d9namics at larger scales 5articularl9 where habitat is less suitable- Given that some Sylvia s5ecies successfull9 colonised large eNtents of suitable habitat in Northern Eurasia that have onl9 become available after the last glacial maNimumA we do not eN5ect a strong effect of taNon age or habitat shift since the last glacial maNimum on range filling- Finall9A we eN5ect greater range filling in s5ecies with higher dis5ersal abilit9- &his stud9 is the first to use a LuantitativeA com5arative a55roach to determine how biotic interactionsA dis5ersal abilit9A taNon ageA habitat shift since the last glacial maNimum *$G", and their interactions influence range filling-

$." 1ethods
$.".1 3ata
For data on the distribution of the Sylvia s5ecies we used breeding range ma5s from a monogra5h *%hirihai et al.A '((),A eNce5t for S. abyssinica who was identified as a member of the genus more recentl9 *B2hning-Gaese et al.A '((+, and whose breeding range was taken from *Fr9 et al.A '(((,- &he range ma5s in %hirihai et al. *'((), are based on a combination of eN5ert knowledge with an eNtensive collation of 5oint records and re5resent the most com5rehensive com5ilation of knowledge about global Sylvia warbler distributions available to date- We restricted our anal9sis to the breeding ranges because the breeding season is a +<

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds critical life-c9cle stage for 5o5ulation 5ersistence where habitat reLuirements of the s5ecies are likel9 to be most eNacting */ulliamA '(((,- &he breeding range ma5s show a high amount of s5atial detail and were thus gridded at a resolution of '@ km S '@ km and converted into 5resences and absences- Given that the decision on how much of a range must be 5resent in a grid cell for a s5ecies to be considered 5resent is somewhat arbitrar9A we conducted all anal9ses for two range ma5 conversion thresholds. )([ and @([ 5resent- We eNcluded S. melanothora) and S. balearica from the anal9ses since their ver9 low 5revalenceA i-e- the 5ro5ortion of grid cells in the stud9 region the9 occu59 *"anel et al.A '((),A rendered them unsuitable for modelling at the continental scale and no reliable range filling values could be obtained for themWe used environmental data from all biogeogra5hic realms currentl9 inhabited b9 Sylvia warblers. /alearcticA 0frotro5ic and #ndo-mala9 *=lson et al.A '((),- We included the #ndomala9 region because three Sylvia s5ecies have eNtensive wintering ranges there- ?enceA this realm is accessible and suitable for Sylvia warblers and should not be eNcluded a 5riori as 5otential habitat- 0ll environmental data were resam5led to the same '@ S '@ km grid as the gridded range ma5s- =ur choice of environmental data was informed as much as 5ossible b9 Sylvia ecolog9 and behaviour- We used mean tem5erature and total 5reci5itation to re5resent abiotic constraints *time 5eriod )8E);8(A 4> 4$ '-(H htt5.TTwww-cru-uea-ac-ukTcruTdataTtmc-htmA New et al.A '(((, and the normalised difference vegetation indeN *NDD#, to reflect 5lant 5roductivit9 *time 5eriod )8:';)888H Global $and 4over Facilit9H htt5.TTglcf-umiacs-umd-eduTdataTgimmsTH &ucker et al.A '((@,- NDD# correlates with green biomass and net 5rimar9 5lant 5roductivit9 *e-g- 4hong et al., )88+, and ma9 thus be linked to the availabilit9 of bird food resourcesA in 5articular insectsA in the breeding season *?urlbertA '((B,- We used the mean values of these variables for the three 5eak breeding months for each s5ecies for modelling *?eikkinen et al.A '((E,- &he breeding season for each s5ecies was defined based on information collated from the literature * rban et al.A )88<H Fr9 et al.A '(((H %hirihai et al.A '(()H Bauer et al.A '((@,- FurtherA since habitat choice of Sylvia warblers is strongl9 affected b9 vegetation t95e and structure *%hirihai et al.A '((),A we also used data on vegetation cover using the "D $and 4over 4lassification data *htt5.TTglcf-umiacs-umd-eduTdataTlandcoverH ) km 5iNel resolutionA ?ansen et al.A '(((,- We distinguished between o5en shrub-landA closed shrub-landA wooded grasslandA woodland and non-needleleaf forest and calculated the 5ro5ortion of each of these classes for each grid cell-

+:

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds

$.".2 -!ecies distribution models and range %illing


Quantif9ing range filling reLuires an estimate of a s5eciesP 5otential range- For each Sylvia s5eciesA we fitted s5ecies distribution models combining the climateA remote-sensing and classified land-use data for each grid cell with the range ma5s and 5roIected them onto the geogra5hic realms where Sylvia warblers 5resentl9 occurA i-e- the /alearcticA 0frotro5ic and #ndo-mala9- &he sum of the grid cells 5redicted as suitable for the s5ecies b9 the distribution model re5resents an estimate of the 5otential range- &o assess the sensitivit9 of our results to the modelling methodA we used two different s5ecies distribution modelling algorithms. boosted regression treesA which can fit ver9 com5leN relationshi5s in a data-drivenA iterative a55roach and ridge regressionA where fitted relationshi5s are eN5licitl9 s5ecified and t95icall9 sim5lerBoosted regression tree models are built in an iterative 5rocedureA where multi5le regression trees *i-e- models that relate the 5robabilit9 of a s5eciesP 5resence to environmental conditions b9 recursive binar9 s5litsH ?astie et al.A '((),A are combined in a linear fashion and subseLuent regression trees focus on the residuals of the 5revious model so as to minimise a loss function such as deviance *Elith et al.A '((:,- &he 5rocess of building and combining the collection of regression trees is called 6boosting7 *Friedman et al.A '(((,- 0s a resultA boosted regression trees fit com5leN non-linear effects and interactions in a data-driven fashion- We fitted models using a bag fraction *the 5ro5ortion of data drawn randoml9 at each iterative ste5, of (-@ and a tree com5leNit9 of seven- $earning rate *the contribution of each tree to the final model, was adIusted according to the number of 5resences for each s5ecies *W )(( 5resences. (-(()A W )((( 5resences. (-()A W )(A((( 5resences. (-('A \ )(A((( 5resences. (-(@,- &he o5timal number of trees was estimated using )(-fold cross validation to calculate 5redictive deviance on models of increasing com5leNit9A 9ielding final models with '(((; :((( trees- &o evaluate the final modelsA we used )(-fold cross validation with each of ten data subsets having the same 5revalence as the original data and re5ort cross-validated 0 4 and 5ercentage of deviance eN5lained>idge regression is a logistic regression techniLue where model com5leNit9 is constrained through a 5enalt9 term to avoid over-fitting *>eineking K %chr2derA '((E,- #n this method the generalization abilit9 of the logistic regression model is o5timised to enhance the fit on the training data b9 increasing model com5leNit9 onl9 when the resulting decrease in variance outweighs the increase in bias- &he ridge or 5enalised maNimum likelihood method uses the

+8

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds sum of the sLuared values of the 5arameter estimates to Luantif9 model com5leNit9 *?arrellA '((),- >idge regressions were fitted using restricted cubic s5lines with three nodes for tem5eratureA 5reci5itation and NDD#- &he five land cover variables were fitted as linear terms- For each modelA we estimated the best 5enalt9 value b9 o5timizing a modified 0#4 *?arrellA '((),- &o evaluate the final modelsA we used a bootstra5 resam5ling 5rocedure *EfronA )8:+, with one thousand re5licates and re5ort validated >'Nagelkerke and 0 4 as 5erformance measures- Bootstra5 sam5les were generated b9 randoml9 selecting grid cells with re5lacement while kee5ing the same 5revalence and total number of grid cells as the original dataset- For each resam5ling runA we refitted the model on the bootstra5 sam5le and calculated the difference in 5erformance measures between the bootstra5 sam5le and the original data- &his difference is an estimate of statistical o5timism i-e- the tendenc9 of a model to have better 5redictive accurac9 when evaluated using the training data as o55osed to new data *%te9erbergA '((8,- %ubtracting the average o5timism over all resam5ling runs from the 5erformance of the model fitted and evaluated on the original data then gives the final internall9 validated 5erformance value *?arrellA '((),We acknowledge recent criticism of 0 4 as a measure of evaluation for s5ecies distribution models *$obo et al.A '((:,- &he 5revalence of our s5ecies in the stud9 area is low *based on @([ range ma5 conversion threshold. min U (-(((BA '@[ Luartile U (-(((EH median U (-()BA <@[ Luartile U (-(+<A maN U (-'+),- &his might lead to an overestimation of 0 4 values- CetA we note that 0 4 values are not significantl9 lower for s5ecies with higher 5revalence *r \ ](-BA ! \ (-(E for all algorithms and range ma5 conversion thresholds,- 0lso none of our h95otheses and tests is based on 0 4 values andA thusA we do not eN5ect that 5otentiall9 overestimating model 5erformance should bias our results&he out5ut of our s5ecies distribution models is a continuous 5robabilit9- %ince we were interested in the size of a s5eciesP 5otential rangeA it was necessar9 to define a threshold to convert the continuous out5ut into a binar9 classification of 6suitable7 versus 6unsuitable7 habitat- &o assess the sensitivit9 of our results to var9ing this thresholdA we calculated our anal9ses for three different threshold rules identified as best 5ractice b9 a com5rehensive com5arative stud9A $iu et al. *'((@,- We 5resent results for setting the threshold so that s5ecificit9 eLuals sensitivit9 in the main teNt and 5rovide results for two additional threshold rules *threshold U mean occurrence 5robabilit9H threshold U observed 5revalence, in a55endiN ' to demonstrate that our results were robust to the choice of threshold rule-

B(

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds "odelling was carried out in > *version '-@-)A > Develo5ment 4ore &eam '()), using 5ublished code and libraries *?arrellA '(()H Elith et al.A '((:H Freeman K "oisenA '((:,- For each Sylvia s5eciesA we calculated range filling as the area of the range from the range ma5 *realised range, divided b9 the area 5redicted as 5resence b9 the s5ecies distribution model *5otential range,-

$."." Botential determinants o% range %illing ,ompetition


We measured com5etition in two wa9s. firstA the mean number of congeneric s5ecies in grid cells in the unoccu5ied 5arts of the 5otential range and secondA the mean number of congeneric s5ecies 5er grid cell in subsets of the 5otential range *i-e- both occu5ied and unoccu5ied, based on habitat suitabilit9- &he first a55roach measures the 5otential role of com5etition in 5reventing the Sylvia warblers from eNtending their ranges into the unoccu5ied 5arts of the 5otential range- We eN5ect lower range filling for s5ecies with a high mean number of congenerics in those 5arts of their 5otential range- &he second a55roach measures how the 5otential im5act of local com5etition on large-scale distributions is affected b9 habitat suitabilit9- We eN5ect range filling to be more strongl9 im5acted b9 com5etition in areas of lower habitat suitabilit9- &o eN5lore this issueA we re5eated our anal9ses using the mean number of congenerics in all of the 5otential rangeA the least suitable @([ of the 5otential range and the least suitable '@[ of the 5otential range- ?abitat suitabilit9 was Luantified as the continuous out5ut of the boosted regression trees and ridge regressions used to identif9 the s5eciesP 5otential rangesBoth our a55roaches to measure com5etition assume that the 5resence of several Sylvia s5ecies in a grid cell signifies an increased chance that those s5ecies will interact locall9&here are several 5otential issues with this assum5tion- FirstA the number of congenerics in a cell could sim5l9 re5resents habitat heterogeneit9H in more heterogeneous cells Sylvia warbler s5ecies could occur as s5atiall9 segregated 5o5ulations in different habitats within the cell andA thereforeA not com5ete- &his is unlikel9 because habitat 5references of Sylvia warblers are similar *%hirihai et al.A '((), and local co-occurrence and use of the same habitat is well documented in Sylvia warblersA even in small areas *e-g- ElleA '((+. ' s5eciesA )-': km' H /ons et al.A '((:. + s5eciesA (-)E km'H 4od9 K WalterA )8<E. B s5eciesA (-(+ km' ,- %econdA using the mean number of congenerics in a given area as a measure of local com5etitive interactions B)

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds involves the sim5lif9ing assum5tion that all co-occurring congenerics are eLuall9 im5ortant com5etitors of a given focal s5ecies- While the im5ortance of com5etitive interactions among Sylvia warblers ma9 var9 *e-g- %chaefer K BarkowA '((B,A com5etition has been documented for numerous different s5ecies 5airs- FurtherA co-occurrences of large numbers of congenerics mainl9 reflect combinations of tem5erate and "editerranean s5ecies *Fig- B-)a, for which evidence for intrageneric com5etition is strongest *e-g- 4od9 K WalterA )8<EH 4od9A )8<:H GarciaA )8:+H "artin K &hibaultA )88EH ElleA '((+H /ons et al.A '((:,- ?enceA the mean number of congenerics likel9 5rovides a useful measure of variation in com5etitive 5ressure even though it does not eN5licitl9 incor5orate differences in interactions strength within the genus- Finall9A Sylvia warblers ma9 com5ete with birds outside the genus- While information on com5etitive interactions between Sylvia warblers and other bird genera is far from com5leteA 5articularl9 for the tro5icsA we are not 5resentl9 aware of an im5ortant eNtrageneric com5etitor- Given that we have evidence for local-scale intrageneric com5etition and that we can assume that com5etition should be most severe within the genus where foraging behaviour and mor5holog9 are most similar *Gotelli et al., '()(,A we focus on intrageneric com5etition among the Sylvia warblers-

&ispersal ability
&o Luantif9 the dis5ersal abilit9 of each s5eciesA we took an ecomor5hological a55roach- We use mor5hological traits that are related to natal dis5ersal distance in 5asserines and have been 5reviousl9 identified as the most useful surrogate measure for dis5ersal abilit9 in this grou5 *Dawideit et al.A '((8,. 1i55Ps distance *distance between ti5 of the ^rst secondar9 and ti5 of the longest 5rimar9Twing ti5 with the wing folded, divided b9 bill de5th- Birds with high 1i55Ps distance have more 5ointed wings which makes forward flight faster and more efficient *>a9nerA )8::H NorbergA )8:8H $eisler K WinklerA '((+,- %hallow bill de5th is indicative of insectivorous migrator9 birds which tend to have larger dis5ersal distances */each et al.A '(()H Dawideit et al.A '((8,-

#a)on age
0s an estimate of the age of each Sylvia s5eciesA we used data from a time-calibrated 5h9logen9 *B2hning-Gaese et al.A '((E,- %ince we had no genetic data for S. desertiA we omitted this s5ecies from our anal9ses- &aNon age in m9a was log-transformed to satisf9 distributional assum5tions of regression anal9sis- We acknowledge that taNon age as defined B'

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds from nodes in a 5h9logenetic tree ma9 underestimate the true age of a s5eciesA e-g- where it 5ersists after giving rise to daughter s5ecies *Webb K GastonA '(((,-

'abitat shift since -.(


#n order to assess how much the geogra5hic 5osition of a s5eciesP 5referred habitat has shifted since the late /leistoceneA we fitted s5ecies distribution models to current data and 5roIected them back to $G"- %5ecifications are the same as for s5ecies distribution models used to Luantif9 range fillingA but with two eNce5tions. FirstA to 5roIect models back in time we had to restrict the anal9ses to variables that were available for both time 5eriods- ?enceA for these models we used onl9 tem5erature and 5reci5itation for the 5resent *)8E(;)88(, and for the last glacial maNimum *')-((( B4, and no data on NDD# or vegetation cover- %ince estimations of 5ast climate var9 with the general circulation model *G4", usedA we conducted the anal9ses with data from two different G4"sA the communit9 climate s9stem model *44%", and the model for interdisci5linar9 research on climate *"#>=4,- $a9ers from the /aleoclimate "odelling #ntercom5arison /roIect /hase ## */"#/'H htt5.TT5mi5'-lsce-i5sl-frT, *Braconnot et al.A '((<, were downscaled using the 5roIected change in tem5erature or 5reci5itation derived from the difference between G4" out5ut for the 5ast and 5resent as a55lied to World4lim current climate *see htt5.TTwww-worldclim-orgTdownscaling,- %econdA as breeding 5henolog9 in the 5ast is unknownA we used 9earl9 means of the climate variables for both the 5resent and the $G" instead of averaging over the three 5eak breeding months- We acknowledge that 5roIecting s5ecies distribution models back in time involves a number of sim5lif9ing assum5tions?oweverA understanding the im5act of habitat shift on range filling reLuires eN5loring variation in s5atial shifts in tem5erature and 5reci5itation regimes at the continental scale- =ur conclusions are therefore robust to uncertaint9 in the s5atial delineation of 5ast s5ecies ranges at higher resolutionsWe Luantified habitat shift since $G" as the area of geogra5hic non-overla5 between the 5ast 5otential range and the 5resent 5otential range as 5redicted from the above s5ecies distribution models divided b9 the combined area of both 5otential ranges-

B+

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds

Statistical %nalysis
We used multi5le regression to assess the influence of com5etitionA dis5ersal abilit9A taNon age and habitat shift since $G" on range filling- We started with a model containing the four 5redictors as linear terms and checked for non-linearit9 in the relationshi5s b9 eNamining smoothed scatter5lot matrices and ceres 5lots *FoNA )88<,- We tested in the models successivel9 all two-wa9 interaction terms in addition to the linear terms and ke5t those in the model that were significant *4rawle9A '((<,- #n additionA we eNamined the results of a model selection 5rocedure based on 0#4cA eN5loring all models that com5lied with the 5rinci5le of marginalit9 *FoNA )88<,- &his anal9sis demonstrated that the 0#4c of our final regression models *&able B-), deviated b9 less than two from the model with the lowest 0#4c and thus belong in the grou5 of eLuall9 well su55orted best models *Burnham K 0ndersonA '(('H see 055endiN +,- /-values for individual t-tests in multi5le regressions were adIusted for multi5le inference to control for inflation of t95e # error *?othorn et al.A '((:,- We tested for 5h9logenetic signal in the residuals of the final model using "oranPs # and the 5h9logenetic distance matriN 0 */avoine et al.A '((:,- 0ll anal9ses were conducted in > *version '-)'-'A > Develo5ment 4ore &eam '()), using 5ublished code and libraries *FoNA '((+H Dra9 K DufourA '((<H ?othorn et al.A '((:H /avoine et al.A '((:,-

$.$ *esults
%5ecies distribution models for the Sylvia warblers were well validated *mean _ standard deviation over all modelsH boosted regression treesA [ deviance eN5lained U @<-'< _ ):-B(A 0 4 U )st Luartile. (-8<8A median. (-8:BA +rd Luartile. (-8:<H ridge regressionsA > 'Nagelkerke U (-BB _ (-)'A 0 4 U )st Luartile. (-8+8A median. (-8@@A +rd Luartile. (-8EB,Estimates of range filling for Sylvia s5ecies varied from (-'' to (-8) *see 055endiN B,$ong-distance migrants breeding in North-Western Eurasia like S. currucaA S. borin and S. communis filled a large 5ro5ortion of their 5otential rangeA while the lowest range filling values were shown b9 the North 0frican short-distance migrant S. deserticola and the 0frican residents S. lugens and S. boehmi-

BB

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds >ange filling was strongl9 influenced b9 the interaction between dis5ersal abilit9 and com5etition *Fig- B-'A &able B-),- Dis5ersal abilit9 had a 5ositive effect on range filling onl9 for those Sylvia s5ecies that had few com5etitors in their unoccu5ied 5otential range *FigB-',- &aNon age had no consistent effect on range filling *&able B-),- =lder Sylvia s5ecies showed higher range filling than 9ounger s5ecies in onl9 '@[ of model realizations- *FigB-+cA &able B-),- ?abitat shift since $G" had no effect on range filling *Fig- B-+dA &able B-),&hese results were robust to modelling decisionsA such as threshold used to convert range ma5s to 5resences and absencesA model algorithm and $G" general circulation model *see methodsH &able B-),- None of the final regression models showed 5h9logenetic signal in the residuals- &he effect of the interaction of com5etition and dis5ersal on range filling de5ended on the habitat suitabilit9 of the area considered for the assessment of com5etition *&able B-'A 055endiN +,- >ange filling was most strongl9 related to com5etition in areas of low habitat suitabilit9 *&able B-'A 055endiN +,-

5igure $.2+ #nteractive effect of dis5ersal abilit9 and com5etition on range fillingA calculated with )([ range ma5 conversion thresholdA boosted regression treesA and 44%" climate model *&able B-)A line ),- %hown are regression lines illustrating the effect of dis5ersal abilit9 on range filling for different levels of com5etition *FoNA '((+,- >egression line slo5es for com5etition levels '-+8 and +-@+ are not significantl9 different from zero- c U com5etition-

B@

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds =able $.1+ Effects of com5etition in the unoccu5ied 5arts of the 5otential rangeA dis5ersal abilit9A taNon ageA habitat shift since $G" and the interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,A significances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ )([ )([ )([ @([ @([ @([ @([ model algorithm 5ast climate model 44%" "#>=4 44%" "#>=4 44%" "#>=4 44%" "#>=4 com5etition dis5ersal abilit9 log *taNon age, (-B8 V *(-)E, (-B8 V *(-)@, (-++ *(-)', (-+) *(-)+, (-B) *(-):, (-B' *(-)<, (-+B *(-)', (-+( *(-)+, habitat shift since $G" (-)B *(-)E, (-'B *(-)@, -(-+' *(-)', -(-'E *(-)), (-(8 *(-):, (-'' *(-)E, -(-+B *(-)B, -(-'' *(-)+, dis5ersal abilit9 S com5etition -(-<' VV *(-'(, -(-<) VV *(-):, -(-EE VVV *(-)+, -(-EB VV *(-)+, -(-E' V *(-'), -(-E' V *(-', -(-<( VVV *(-)B, -(-E: VV *(-)B, model >'

B>& B>& >idge >idge B>& B>& >idge >idge

-(-B+ *(-'(, -(-B< *(-)8, -(-++ *(-)+, -(-B+ V *(-)+, -(-BE *(-'', -(-@' *(-'), -(-++ *(-)@, -(-B@ V *(-)B,

(-B' *(-):, (-B@ *(-)<, (-B: VV *(-)), (-B8 VV *(-)), (-B8 *(-'(, (-@' *(-)8, (-B8 VV *(-)', (-B8 VV *(-)',

(-E: VV (-<) VVV (-:@ VVV (-:B VVV (-E) VV (-EB VV (-:+ VVV (-:) VVV

V ! W (-(@H VV ! W (-()H VVV ! W (-(()-

BE

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds

5igure $."+ Effect of *a, com5etitionA *b, dis5ersal abilit9A *c, taNon age and *d, habitat shift since $G" on range filling for the multi5le regression model *n U '+, based on )([ range ma5 conversion thresholdA boosted regression trees and 44%" climate model and *&able B-)A line ),- $everage 5lots after %all *)88(,-

B<

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds =able $.2+ Effect of habitat suitabilit9 on the relationshi5 between com5etition and range filling- ?abitat suitabilit9 U subset of the 5otential range used to estimate com5etitionA "ulti5le regressions based on boosted regression treesA )([ range ma5 conversion threshold and 44%" 5ast climate model- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,A significances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+habitat suitabilit9 com5etition dis5ersal abilit9 log *taNon age, habitat shift since $G" (-)< *(-'', (-'( *(-'), (-'( *(-):, dis5ersal abilit9 S com5etition -(-<+ *(-+), -(-<< R *(-+(, -(-:@ V *(-'@, (-E< VV (-E) VV model >'

all habitat

-(-B( *(-++,

(-B< *(-'+, (-B< *(-'', (-B: R *(-)8,

(-E( V *(-)8, (-E+ V *(-):, (-@8 V *(-)<,

(-@: VV

least suitable @([

-(-BB *(-+),

least suitable '@[

-(-@+ *(-'@,

R ! W (-)H V ! W (-(@H VV ! W (-()-

$.' 3iscussion
>ange filling in Sylvia warblers was strongl9 determined b9 an interaction between dis5ersal abilit9 and com5etition- Sylvia s5ecies with higher natal dis5ersal distances filled a larger 5ro5ortion of their 5otential rangeA but onl9 when dis5ersal was not constrained b9 intrageneric com5etition- For eNam5leA S. curruca and S. rueppelli are both good dis5ersersA but onl9 S. curruca has been able to colonise most of its 5otential habitat in Northern Euro5e and 4entral 0sia where few other Sylvia warblers occurH S. rueppelli failed to invade large 5ortions of its 5otential habitat in the "editerranean where s5ecies richness of Sylvia warblers is highest *Fig- B-)b,- 4onversel9A the rather 5oor dis5ersers S. boehmi and S. undata show no marked difference in their range filling although Sylvia richness in the 5otential habitat of S. undata in 4entral Euro5e is much higher than for S. boehmi in East 0frica *Fig- B-)b,/revious studies on range filling have not addressed com5etition directl9 even though it has long been considered an im5ortant determinant for sha5ing s5ecies ranges *Dobzhansk9A )8@(H "ac0rthurA )8<',- Distribution 5atterns that concur with eN5ected effects of

B:

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds com5etition at the regional scale have been found for mammals *0nderson et al.A '(('H %anchez-4ordero et al.A '((:, and birds *Gross K /riceA '(((,- ?oweverA Luantitative evidence of the effect of com5etitive interactions at larger s5atial scales remains ver9 limited *but see ?eikkinen et al.A '((<H Gotelli et al.A '()(,- >ecentl9A intrageneric com5etition in songbirds has been found to be im5ortant for setting elevational range limits in tro5ical bird communities *Jankowski et al.A '()(,- Where Sylvia s5ecies occur s9nto5icall9A com5etition freLuentl9 manifests itself as segregation at the microhabitat and diet level *"artin K &hibaultA )88EH /ons et al.A '((:, and sometimes as inters5ecific territorialit9 *4od9 K WalterA )8<EH 4od9A )8<:H GarciaA )8:+,- #t is conceivable that such local segregation of niches and s5ace might allow coeNistence at larger s5atial scales *WiensA )8:8H $ovette K ?ochachkaA '((E,- &his seems to be the case in the core of the s5eciesP rangesA where habitat suitabilit9 is o5timal for Sylvia warblers- 4onseLuentl9A 5atterns of congeneric s5ecies richness across the whole 5otential range are not related to range filling- NeverthelessA our results indicate that the effects of intrageneric com5etition observed at the local scale can influence the size of the realised ranges of the s5ecies at the geogra5hic scale when habitat suitabilit9 is low- 0ccordingl9A congeneric s5ecies richness was increasingl9 related to range filling when Luantified onl9 for less suitable habitat- >ange margins often have lower habitat suitabilit9 than the range core *Brown et al.A )88EH %agarin K GainesA '((',- &he unoccu5ied 5arts of the 5otential range of Sylvia warblers are mainl9 areas with low habitat suitabilit9 located at the edge of the 5otential range- ?enceA range filling was even more strongl9 related to the number of congenerics in the unoccu5ied 5otential range than to the number of congenerics in the least suitable '@[ of the 5otential range- &hese results corres5ond well to recent eN5erimental trans5lant studies showing that the harsher abiotic conditions at range edges lead to an increased im5act of intrageneric com5etition on salamanders *4unningham et al.A '((8, and of com5etition b9 neighbouring annuals on an annual legume *"ooreA '((8,&husA while Sylvia warblers engage in local-scale com5etitive interactions throughout their ranges *4od9 K WalterA )8<EH 4od9A )8<:H GarciaA )8:+H "artin K &hibaultA )88EH ElleA '((+H /ons et al.A '((:,A this translates into conseLuences for large-scale distributions onl9 in less suitable habitat#n Sylvia warblersA dis5ersal abilit9 had a clear effect on range filling but there was no evidence for dis5ersal limitation caused b9 historic habitat shift- 0 similar 5attern was observed in the 5lant famil9 /roteaceaeA where dis5ersal abilit9 influenced range filling and

B8

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds historic climate change or geogra5hic barriers had little influence *%churrA '((<,- 4onversel9A for tree and mammal taNaA where range filling is 5rimaril9 a result of historic constraints related to geogra5h9 and climateA no effect of dis5ersal abilit9 could be detected *%venning K %kovA '((BH "unguia et al.A '((:,- &hese results suggest that the effect of dis5ersal abilit9 on range filling is most 5ronounced when dis5ersal has not been constrained b9 current or historic s5atial configuration of habitats and climateWe found no consistent tendenc9 for older Sylvia warblers to fill more of their 5otential range than 9ounger s5ecies- 0n effect of taNon age on range filling has been suggested to result mainl9 from the limited time for dis5ersal available to 9ounger s5ecies */aul et al.A '((8,- 0nother 5ossible eN5lanation for a correlation between high range filling and taNon age might be that older s5ecies 5ossess a suite of traitsA such as broad environmental or habitat nichesA disturbance toleranceA or high 5o5ulation growth rates that 5romotes ra5id range filling and simultaneousl9 enables the long-term survival of a taNon *Webb K GastonA '(((,We might eN5ect older s5ecies to be less affected b9 com5etition due to their 5otentiall9 greater divergence in ecological reLuirements- &hereforeA a third eN5lanation for higher range filling in older s5ecies might be that those s5ecies are less affected b9 the 5resence of congeners than 9ounger s5ecies- ?oweverA we did not find su55ort for an interaction between taNon age and com5etition- =ther studies relating taNon age to range filling have focused on 5lantsA where no clear relationshi5 has been found *%churrA '((<H /aul et al.A '((8,- &he oldest Sylvia s5ecies with the highest range filling *e-g- S. currucaA range filling U (-8)A range size U 'A8<@A((( km', must have occu5ied their large ranges ra5idl9 after the last glacial maNimum towards the end of the /leistocene *%hirihai et al.A '((),- ?enceA the Sylvia warblers a55ear not to be dis5ersal limited- 0lsoA older s5ecies did not necessaril9 have more time for dis5ersal- &he ra5id eN5ansion of s5ecies into Northern Eurasia ma9 also eN5lain wh9 we did not find an effect of habitat shift since $G" on range filling for the Sylvia warblersA even though Luaternar9 climate change clearl9 had an effect on the current distribution of man9 taNa *e-g- %venning K %kovA '((B,We observed an increasing dis5arit9 in range filling values based on boosted regression trees and ridge regressions for s5ecies with lower 5revalence *055endiN B,- /revalence ma9 affect the accurac9 of distribution models *e-g- "armion et al.A '((8,- CetA although models for s5ecies with lower 5revalence showed increasing variation in range filling between model algorithmsA identification of range filling determinants was robust to modelling method- We

@(

+ ,ompetition and dispersal ability interact to determine geographic ranges of birds also note that the validit9 of the hindcasting a55roach used to Luantif9 habitat shift since $G" rests on the assum5tion of niche conservatism for Sylvia warblers in the last ')A((( 9ears *B2hning-Gaese et al.A '((+,- FurtherA our 5otential range estimates ma9 underestimate the true eNtent of the geogra5hic area suitable for the s5ecies since the9 are derived from realised occurrences */ulliamA '(((,- Due to being limited to bird s5ecies with strong evidence for local-scale com5etitionA our sam5le size is at the lower end of what is desirable for our anal9sis- ?enceA high model >' should not be taken as an indication that range filling is not determined b9 factors not included in our anal9sis- We also note that a measure attem5ting to ca5ture the signal of local com5etition in Sylvia warblers at larger scales could be enhanced b9 detailed com5arative studies on interaction strength within and be9ond the genus as well as abundance information across the whole intercontinental eNtent of Sylvia distributions- While the Sylvia warblers are 5robabl9 among the best-researched bird generaA such com5rehensive information is not available and we 5refer to use a measure that reflects the eNtent of current knowledge- #f this a55roach results in an oversim5lified measureA we would eN5ect to find no consistent relationshi5 with range filling- Finall9A we acknowledge this is a correlational stud9- #t would be eNciting to com5are our findings to results from largescale eN5erimentation addressing establishment success and intrageneric com5etitive 5ressure across the entire 5otential geogra5hic range of several Sylvia s5ecies#n sumA our results indicate that com5etition and dis5ersal abilit9 are among the forces that determine the distribution of s5ecies at the continental scale- ?oweverA the effect of com5etition on range-filling is contingent on s5ecies traits and habitat suitabilit9- We therefore strongl9 recommend the integration of multi5le factorsA including biological interactionsA and the eNamination of 5otential interactions among those factors in anal9ses of range characteristics- We would also caution against the assum5tion that biotic interactions are generall9 not im5ortant for large-scale range d9namics- #f com5etition generall9 5la9s a dominant role and even interacts with other factors in determining if a s5ecies is able to colonise and occu59 suitable habitat at large scalesA then it willA indeedA be a challenge to 5redict the abilit9 of s5ecies to 5ersist in the face of habitat shifts caused b9 climate change-

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Sylvia borin

Will be submitted to an international Iournal of ecolog9 as. $aubeA #-A GrahamA 4- ?- K B2hning-GaeseA 1- Niche availabilit9 in s5ace and time. migration in Sylvia warblers-

@+

1 2iche availability in space and time3 migration in Sylvia warblers

'.1 Abstract
#n the conteNt of recent advances in ecological niche modellingA both the environment and the ecological niche of a s5ecies have often been treated and Luantified as static entities- #n realit9A the environment and s5ecies? niche reLuirements are d9namic on a variet9 of scalesWe 5ro5ose a conce5tual framework of how s5ecies ? realised niches and geogra5hic ranges are sha5ed b9 the decou5led s5atio-tem5oral availabilit9 of different environmental conditions and b9 changes in niche reLuirements throughout an organism ?s lifetime- &esting 5redictions derived from the framework using migration of Sylvia warblers 9ielded new insights. 4limate niche tracking was unlikel9 to be the main driver of migration in the genus and 5otentiall9 conflicted with land-cover niche tracking- Sylvia niches were smaller during the breeding seasonA demonstrating that niche reLuirements can be d9namic in time- We suggest that taking d9namic environments and niche reLuirements into account enhances our understanding of the drivers behind s5atial movements of organisms and the d9namics in their niches and geogra5hic ranges-

'.2 0ntroduction
>ecentl9A there is a renewed conce5tual focus in ecolog9 on the ecological nicheA 5artl9 s5arked b9 develo5ments in s5ecies distribution modelling *e-g- %oberonA '((<H /earman et al., '((:H 4olwell K >angelA '((8H Wiens et al., '()(,- #n this conteNtA both the environment and the ecological niche of a s5ecies have often been treated and Luantified as static entities *Fisher et al., '()(H FranklinA '()(,- For eNam5leA it is common in s5ecies distribution modelling to use occurrence and environmental data averaged over long time 5eriodsA e-g9earsA to estimate the realised niche of a s5ecies *e-g- Dormann et al., '()(,- ?oweverA in realit9A the environmental conditions available to organisms are highl9 d9namic in s5ace and time on a variet9 of scales *e-g- da9sA seasonsA decades,- &hese d9namics result in diverse 5henomena such as the dail9 movements of zoo-5lankton in the water column *Williamson et al., '()),A annual migrations of birdsA mammalsA fish and insects *"ilner-Gulland et al., '()),A 5eriods of dormanc9 in crustaceansA fungi and 5lants *$ubzens et al., '()(,A and range shifts of organisms as a conseLuence of climate change *?untle9 et al., '((EH Barbet-"assin et al., '((8H Doswald et al., '((8,- #n additionA the niche reLuirements of organisms also var9 on a variet9 of tem5oral scales *e-g- da9sA seasonsA ontogenetic develo5ment,- ?ow d9namic @B

1 2iche availability in space and time3 migration in Sylvia warblers environments and niche reLuirements affect realised niches and geogra5hic ranges of s5ecies is a central Luestion in ecolog9 and evolution and is critical for managing s5ecies given ongoing environmental change *GastonA '((+H /earman et al., '((:,- ?ere we develo5 a framework that can enhance our understanding of niche and range d9namicsA and use Sylvia warblers to test 5redictions derived from the framework to assess its utilit9%easonal changes in the environment have long been linked to the regular movement of organisms across geogra5hic regions *$ackA )8@BH "ilner-Gulland et al., '()),- %uch movements have sometimes been associated with a niche tracking strateg9 *e-g- "artinez"e9er et al., '((BH Nakazawa et al., '((BH Batalden et al., '((<H "arini et al., '()(,- %tudies using s5ecies distribution modelling to eNamine niche tracking have described s5atiotem5oral 5atterns in the realised niches of s5eciesA but the9 have not evaluated the costs and trade-offs associated with 5articular strategies nor have the9 eN5licitl9 related the observed 5atterns to the d9namic availabilit9 of environmental niche conditions in s5ace and time/revious studies have focused on migrating birds and butterflies and the eNtent of overla5 in environmental niche dimensions between their breeding and non-breeding niches *"artinez"e9er et al., '((BH Nakazawa et al., '((BH Batalden et al., '((<H "arini et al., '()(, and on annual tem5erature tracking *Jose5h K %tockwellA '(((,- %5ecies often either show high overla5 between breeding and non-breeding niche *6niche trackers7, or little overla5 *6niche switchers7,/revious studies on niche tracking have eNamined environmental niche dimensions *most often climate, that have similar s5atio-tem5oral d9namics *"artinez-"e9er et al., '((BH Nakazawa et al., '((BH Batalden et al., '((<H "arini et al., '()(,- ?oweverA the availabilit9 of different niche dimensions is not necessaril9 s9nchronised in time and s5ace *JimenezDalverde et al., '((8,- For eNam5leA a seasonal change in local climate is not necessaril9 associated with corres5onding changes in local land cover- 0cross different seasonsA the same climatic conditions ma9 be available in locations with ver9 different land coverNiche reLuirements of organisms are usuall9 assumed to be constant across various tem5oral scales- ?oweverA the subset of niche s5ace an organism uses ma9 var9 dail9A seasonall9 and throughout an organism?s lifetime *Batalden et al., '((<H %uarez-%eoane et al., '((:,- For eNam5leA the niche for energeticall9 more demanding activitiesA such as re5roductionA ma9 be a subset of the general survival niche *GrubbA )8<<H 0lerstam K ?2gstedtA )8:'H &iteuN et al., '((<,A or s5ecies ma9 eNhibit ontogenetic niche shifts as the9

@@

1 2iche availability in space and time3 migration in Sylvia warblers mature *&akimotoA '((+H Coung et al., '((@,- 0s a resultA both niche s5ace availabilit9 and a s5ecies? niche reLuirements ma9 concurrentl9 var9 in time-

'.2.1 6once!tual %rameEork %or niche d8namics in s!ace and time


We attem5t to develo5 a conce5tual framework that incor5orates the d9namic nature of nichesA geogra5hic ranges and the available environment as discussed above and which thus stimulates new Luestions about the drivers behind s5ecies ? niche and range d9namics- ?ereA we consider niches both in the Grinellian traditionA in the sense that we focus on niches as defined b9 broad environmental conditions *GrinellA )8)<H %oberonA '((<,A and in the ?utchinsonian traditionA in the sense that we regard the niche as an entit9 defined in an abstract h95ers5ace which can be limited b9 biotic interactions *?utchinsonA )8@<H 4olwell K >angelA '((8,- Building on these traditionsA we focus on realised niches Luantified from large-scale occurrence dataA which also reflect dis5ersal limitations and biotic interactions *%oberonA '((<H 4olwell K >angelA '((8,- =ur aim is to better understand how the d9namics in these realised niches are related to the d9namic availabilit9 of total niche s5aceA i-e- of the availabilit9 of niche dimensionsA such as environmental conditionsA in time and geogra5hic s5ace&o conce5tualise the strategies of organisms to co5e with the highl9 d9namic nature of available niche s5aceA we can distinguish two eNtreme scenarios of how organisms might react to a change in locall9 available environmental conditions. ), =rganisms move in geogra5hic s5ace to track their favoured environmental conditions- &husA the9 alwa9s sta9 within the s5ecific subset of niche s5ace the9 5refer *Fig- @-)A red strateg9,- &his strateg9 contributes to ecological 5henomena such as diurnal vertical 5lankton migration or the annual migrations of ungulates in %outhern 0frica *"ilner-Gulland et al., '())H Williamson et al., '()),- ', 0lternativel9A organisms sta9 where the9 are and tolerate the local change in environmental conditions- &he organismPs niche then has to encom5ass the full range of environmental conditions available locall9 through time *%oberonA '((<H /earman et al., '((:H Fig- @-)A blue strateg9,- &o endure harsh conditions in situA organisms ma9 even tem5oraril9 reduce their activit9 levelA e-g- hibernation in mammals or winter dormanc9 in trees *$ubzens et al., '()(,- #ntermediate strategies between these two scenarios of 5erfect niche tracking and no niche tracking are conceivable and have been observed *"artinez"e9er et al., '((B,- =rganisms incur an energ9 cost both for movement across regions *0lerstam et al., '((+H Wikelski et al., '((+, and for high environmental tolerance *DeWitt et @E

1 2iche availability in space and time3 migration in Sylvia warblers al., )88:H 4ale9 K "unda9A '((+H 0uld et al., '()(,- ?enceA it seems 5lausible to assume a trade-off between the abilit9 of organisms to track niches in s5ace and the abilit9 to tolerate a wide variet9 of conditions-

1 +pe(ies &2 resident 1 +pe(ies 22 spatial ni(he tra(3ing .i/e &


(ondition &
a0ailable (onditions

.i/e 2

Mini/)/ size o5 ni(he

4n0iron/ental +pa(e

(ondition &

(ondition &

(ondition 2

(ondition 2

(ondition 2

*eographi( +pa(e

y ,lat.-

x ,long.-

y ,lat.-

x ,long.-

5igure '.1+ Framework illustrating organismal strategies to co5e with d9namic niche s5ace%hown are two eNtreme scenarios of niche and range d9namics for a geogra5hic region at two 5oints in time that differ in the availabilit9 of environmental s5ace- )- >esident *s5ecies )A niches and ranges shown in blue,. %5ecies sta9ing in the same geogra5hic location have to tolerate the local change in available environmental niche s5ace- &he s5ecies ? niche then has to encom5ass the full range of environmental conditions available locall9 through timeA resulting in a broad environmental niche- '- %5atial niche tracking *s5ecies 'A niches and ranges shown in red,. %5ecies ma9 move in geogra5hic s5ace to track their favoured environmental conditions- &husA s5ecies alwa9s sta9 within the s5ecific subset of environmental niche s5ace the9 5refer and can have narrow niches&his conce5tual framework is relevant for a wide variet9 of niche dimensionsA both abiotic and biotic- #t can be a55lied to tem5orall9 highl9 d9namic *e-g- climate, and more static *e-gland cover, niche dimensions at the same time and can thus be used to eN5lore the 5otentiall9 com5leN o5timization 5roblems faced b9 s5ecies due to the decou5led availabilit9 of different @<

1 2iche availability in space and time3 migration in Sylvia warblers niche dimensions- 0lsoA b9 allowing the sha5e of the niche s5ace reLuired b9 a s5ecies *Fig@-)A dashed circles in environmental s5ace, to change in timeA the framework can incor5orate d9namic niche reLuirements across seasons and throughout s5ecies? life-c9cles-

'.2.2 A!!l8ing the %rameEork to animal migration


We derive four general 5redictions from our framework to better understand how environmental variation in s5ace and time relates to the s5atial movements and niche characteristics of organisms- We then evaluate these four 5redictions using migration behaviour in Sylvia warblers as a case stud9- &he Sylvia warblers are an eNcellent s9stem to evaluate variation in migration 5atterns as a function of s5eciesP niche characteristics and the d9namic availabilit9 of their niche in s5ace and time- &here is eNtensive information available on their rangesA ecolog9 and 5h9logen9 *e-g- %hirihai et al., '(()H B2hning-Gaese et al., '((+H Doelker K $ightA '()),- &he 'E s5ecies of Sylvia warblers evaluated here use a range of migration strategies including tem5erate and tro5ical residentsA short-A middle- and longdistance migrants *%hirihai et al., '((),- #n the followingA we state each of the four general 5rediction together with the s5ecific Luestion we evaluate for Sylvia warblers.

#rade4offs between niche breadth and migration distance3


#f both strategies in Fig- @-)A niche tracking or having a broad nicheA are costl9 for s5eciesA then we might eN5ect a trade-off between the amount of environmental variation a s5ecies can tolerate and migration distance- %5ecies that move long distances should then have more constant niches throughout the 9ear than more resident s5ecies *i-e- high niche overla5 across seasons sensu Broennimann et al., '()),A 5articularl9 for niche dimensions that are tem5orall9 highl9 d9namic such as climate- ?oweverA if niche dimensions are tem5orall9 more static *e-g- land-cover,A then s5ecies moving longer distances ma9 eN5erience large changes in these dimensions resulting in a low overla5 between different seasons- #t follows that s5ecies that move long distances should have narrower total annual niches for the niche dimensions the9 are tracking and broader total annual niches for niche dimensions that are more static in time but var9 in s5ace%5ecific Luestions. Do Sylvia warblers that migrate longer distances between breeding and non-breeding grounds show greater niche overla5 between these areas for tem5orall9 d9namic climate niche dimensions and lower overla5 in static land-cover niche dimensions` Do Sylvia warblers with longer distances between breeding and non-breeding grounds show @:

1 2iche availability in space and time3 migration in Sylvia warblers lower total annual climate niche breadth and higher total annual land-cover niche breadth` &o address these Luestions we Luantified climate and land-cover niche characteristics in both breeding and non-breading areas for each s5ecies-

2iche tracking
#f the reason for movement is niche trackingA then movement should lead to an increased niche overla5 in the seasonal niches for the tracked niche dimension com5ared to a resident strateg9%5ecific Luestion. #s the climate niche of migrant Sylvia warblers more stable as a result of migration com5ared to if these same warblers had not migrated` &o answer this LuestionA we com5ared the niche overla5 between the breeding and non-breeding conditions which the s5ecies actuall9 eN5erience to the h95othetical niche overla5 resulting from sta9ing either on the breeding or on the non-breeding grounds during the whole 9ear-

.eographic pro)imity
#f organisms that move in geogra5hic s5ace as a res5onse to d9namic niche availabilit9 minimise the cost of movementA the9 should move to the nearest available geogra5hic location with suitable conditions%5ecific Luestion. Do migrant Sylvia warblers move to the closest 5lace with suitable conditions` For this Luestion we 5roIect environmental niche conditions into geogra5hic s5ace and then locate the areas with similar climate and land-cover to the observed breeding and non-breeding grounds-

Seasonal changes in niche re5uirements


#f activities demanding high amounts of energ9A such as re5roduction or accumulation of bod9 reserves before 5u5ation or hibernationA involve more eNacting energ9 and resource reLuirementsA then the high energ9 activit9 niche should be eLual or smaller to the survival nicheA because the survival niche also incor5orates seasons of com5arativel9 lower energ9 and resource reLuirements which can be met under a broader set of environmental conditions%5ecific Luestion. #s the breeding niche of migrant Sylvia warblers eLual to or smaller than the survival niche` ?ereA we test whether niche breadth as derived from the breeding range during the breeding season is smaller or eLual than the total niche breadth derived from the combination of breeding and non-breeding environmental conditions-

@8

1 2iche availability in space and time3 migration in Sylvia warblers

'." 1ethods
'.".1 -tud8 s!ecies
&he Sylvia warblers are a genus of '< 5rimaril9 insectivorous 5asserinesA occurring in Euro5eA 0frica and western 0sia- B2hning-Gaese et al. *'((+, have classified )B of them as residentsA B as short-distance migrants and 8 as long-distance migrants- "igration distances in km for the Sylvia warblersA based on the orthodrome distance between centres of gravit9 for breeding and non-breeding ranges were also taken from B2hning-Gaese et al. *'((+,- /h9logenetic information on the relationshi5 of s5ecies within the genus was taken from Doelker K $ight *'()),-

'.".2 *anges
#nformation on breeding and non-breeding ranges of the Sylvia s5ecies was taken from %hirihai et al. *'((),A eNce5t for S. abyssinica, which was more identified as a member of the genus more recentl9 *B2hning-Gaese et al., '((+, and whose ranges were taken from Fr9 et al. *'(((,- Sylvia dohrniA an island endemic recentl9 added to the genus *Doelker et al., '((8,A was not included in the anal9ses because its eNtremel9 small range 5recluded reliable Luantification of niche characteristics- &he range ma5s in %hirihai et al. *'((), combine eN5ert knowledge with an eNtensive collation of 5oint records and are the most com5rehensive com5ilation of knowledge about Sylvia geogra5hic distributions available&he range ma5s show a high amount of s5atial detail and were thus gridded at a resolution of '@ S '@ km and converted into 5resences and absences- &o assess whether the grid resolution affects our resultsA we conducted all anal9ses for two different range gridding thresholds *i-ethe 5ercentage of minimum overla5 between the range and a grid cell for that grid cell to be classified as 5resence,. )([ and @([ 5resent- Both thresholds 9ielded ver9 similar results *see 055endiN %) in %u55orting #nformation, so we re5ort results using the )([ threshold-

'."." <nvironmental variables


We used environmental data from all biogeogra5hic realms currentl9 inhabited b9 Sylvia warblers. /alearcticA 0frotro5ic and #ndo-mala9 *=lson et al., '((),- &he data were resam5led to the same '@ km S '@ km grid as the gridded range ma5s- We chose environmental variables based on our knowledge of Sylvia ecolog9 and behaviour- We used mean tem5erature and E(

1 2iche availability in space and time3 migration in Sylvia warblers total 5reci5itation to re5resent abiotic environmental conditions *time 5eriod )8E)8(A 4> 4$ '-(H htt5.TTwww-cru-uea-ac-ukTcruTdataTtmc-htmA New et al., '(((,- &hese climatic variables ma9 affect bird distributions directl9 via 5h9siological survival limitsA through their effects on the availabilit9 and 5henolog9 of food resources or via the abundance of com5etitors and 5arasites *?untle9 et al., '((E,- We used the normalised difference vegetation indeN *NDD#H time 5eriod )8:')888H Global $and 4over Facilit9H htt5.TTglcf-umiacs-umd-eduTdataTgimmsTH &ucker et al., '((@, which correlates with green biomass and net 5rimar9 5lant 5roductivit9 *e-g- 4hong et al., )88+, and ma9 thus be linked to the availabilit9 of bird food resources such as insects *?urlbertA '((B,- We used the mean values of these variables for the three 5eak breeding and non-breeding months for each s5ecies *?eikkinen et al., '((E,- /eak breeding and non-breeding seasons for each s5ecies were defined based on information in the literature * rban et al., )88<H Fr9 et al., '(((H %hirihai et al., '(()H Bauer et al., '((@,- %ince habitat choice of Sylvia warblers is determined b9 vegetation t95e and structure *%hirihai et al., '((),A we used the "D $and 4over 4lassification data *htt5.TTglcf-umiacs-umd-eduTdataTlandcoverH )km 5iNel resolutionA ?ansen et al., '(((, to reflect vegetation cover- We distinguished between o5en shrub-landA closed shrub-landA wooded grasslandA woodland and non-needle leaf forest and calculated the 5ro5ortion of each of these classes for each grid cell- &hese eight environmental variables have been confirmed as im5ortant for sha5ing broad-scale Sylvia distributions in several studies develo5ing s5ecies distribution models for the genus *Wisz et al., '((<H Doswald et al., '((8H Barbet-"assin et al., '((8,-

'.".$ (iche characteristics


Ecologists have long debated how to best measure niche overla5 and niche breadth *e-g- ?ornA )8EEH 4olwell K Futu9maA )8<)H Warren et al., '((:H Dormann et al., '()(H >2dder K EnglerA '()),- ?ereA we follow recommendations from Broennimann et al. *'()),- For each s5eciesA we conducted a 5rinci5al com5onent anal9sis of the environmental variables described above for the whole stud9 region */alearcticA 0frotro5ic and #ndo-mala9, including the data from both the breeding and the non-breeding season- De5ending on the LuestionA we used either all environmental variables in the /40 or calculated se5arate /40s for the three climate and five land-cover variables- &he first two 5rinci5al com5onents of the /40 were used as the aNes to describe the total annual environmental s5aceA bounded b9 the minimum and maNimum environmental values found in an9 of the two seasons in the whole stud9 E)

1 2iche availability in space and time3 migration in Sylvia warblers regionFor further anal9sisA the environmental s5ace described b9 the first two /40 aNes was divided into )(( N )(( regularl9 s5aced grid cells *vi6,A with each cell thus re5resenting a uniLue set of environmental conditions- NeNtA we calculated for each s5ecies and for both breeding and non-breeding seasons the densit9 of s5ecies occurrences * oi6, and the densit9 of available environments *ei6, *i-e- the number of grid cells with these environmental conditions in the whole stud9 region during that season, in each grid cell in the environmental s5aceBoth oi6 and ei6 were calculated using a kernel smoothing function to account for im5erfect sam5ling of occurrences and to make the metrics inde5endent of the number of grid cells in environmental s5ace- Dividing oi6 b9 ei6 for each s5ecies and season then gave the occu5anc9 of the environment *zi6, in the grid cells in environmental s5ace- #f the environmental conditions corres5onding to a grid cell were unavailable in a 5articular season *i-e- e i6 U (,A $i6 was set to zero- &his 5rocedure corrects the observed occurrences for the availabilit9 of environmental conditions in each season to ensure unbiased com5arisons *Broennimann et al., '()),Niche overla5 between seasonal ranges was calculated using the D metric *%choenerA )8<(H Warren et al., '((:, on the occu5anc9 values in environmental s5ace *$i6,- &o calculate the D metricA the absolute differences in occu5anc9 values between the two ranges are summedA multi5lied b9 (-@ and then subtracted from one- &he D metric varies from ( *no niche overla5, to ) *com5lete niche overla5,&o calculate niche breadthA we converted the environmental occu5anc9 values *$i6, to 5ro5ortions and then calculated the %hannon indeN *4olwell K Futu9maA )8<),- &his measure of niche breadth thus takes into account both the number of occu5ied grid cells in environmental s5ace and the evenness in the occu5anc9 among those grid cells- For the com5arison of breeding and annual niche breadth within each s5eciesA annual niche breadth was calculated b9 first summing the occu5anc9 values * $i6, for breeding and non-breeding niches in environmental s5ace and then calculating the %hannon indeN on the summed 5ro5ortional occu5anc9 values&o determine whether a s5ecies moved during migration from its breeding range to the closest non-breeding range with suitable non-breeding environmental conditionsA we conducted the following five ste5s- FirstA we 5roIected the occu5anc9 values derived from each s5ecies? seasonal /40s including both climate and land-cover into geogra5hic s5ace to

E'

1 2iche availability in space and time3 migration in Sylvia warblers identif9 suitabilit9 values for each grid cell in geogra5hic s5ace- %econdA to distinguish suitable from unsuitable areas we thresholded the suitabilit9 ma5s using the sensitivit9 eLuals s5ecificit9 ruleA as recommended b9 $iu et al. *'((@, in a com5arative stud9 of thresholding rules- &hirdA we identified the closest suitable non-breeding area b9 calculating the nearest neighbour distances from all suitable non-breeding grid cells to all cells in the breeding rangeForthA we selected the suitable non-breeding grid cells that had the shortest nearest neighbour distances to the known breeding range- We selected the same number of grid cells as the number of grid cells in the known non-breeding range- FifthA we calculated the average of the 5airwise distances between all grid cells in the closest suitable non-breeding area and all grid cells in the known breeding range to obtain the minimum migration distance0nalogousl9A we calculated whether a s5ecies moved from its non-breeding range to the closest breeding range with suitable breeding environmental conditions- &he closest suitable breeding area was identified b9 calculating the nearest neighbour distances of suitable breeding grid cells to the known non-breeding range and then selecting the same number of cells as in the known breeding range with the shortest nearest neighbour distance to the known non-breeding range- &o com5are the minimum migration distances to the actual migration distances for each s5eciesA we did not use the migration distances obtained from B2hning-Gaese et al. *'((+, that were used in the other anal9ses- #nsteadA we calculated actual migration distance as the average of the 5airwise distances between all breeding range grid cells and all non-breeding range grid cells- &his ensured maNimum consistenc9 between the Luantifications of minimum and actual migration distances for this Luestion0ll anal9ses were conducted in > '-)+-) *> Develo5ment 4ore &eam '()), using 5ublished code and libraries *Broennimann et al., '())H Baddele9 K &urnerA '((@,-

'.".' -tatistical Anal8sis #rade4offs between niche breadth and migration distance
We calculated linear regressions between niche overla5 and movement distances and between total annual niche breadth and movement distances se5aratel9 for climate and land-cover niches for all Sylvia warblers- 4losel9 related s5ecies ma9 tend to have more similar traits than distantl9 related s5ecies and thus s5ecies do not necessaril9 re5resent inde5endent data 5oints *?arve9 K /agelA )88),- &o take the 5h9logenetic relationshi5 between Sylvia warblers into accountA we checked the residuals from the linear regressions for 5h9logenetic E+

1 2iche availability in space and time3 migration in Sylvia warblers autocorrelation- We tested for 5h9logenetic signal in the residuals using the 0bouheif test *0bouheifA )888, with 888 randomisations as im5lemented in the > 5ackage ade5h9lo *Jombart et al., '()(, and b9 calculating /agelPs YA a maNimum-likelihood based measure of 5h9logenetic signal */agelA )88<,A and testing for a significant difference to a lambda of zero *no 5h9logenetic structure,A as im5lemented in the > 5ackage 40#4> *FreckletonA '((8,-

2iche tracking
&o determine if the climate niche of migrant Sylvia warblers is more stable between seasons as a result of migrationA we calculated the climatic niche overla5 for the breeding and nonbreeding grounds assuming a given warbler had not migrated and com5ared this overla5 to the climate niche overla5 the migrants actuall9 eN5erience- We calculated 5aired t-tests to com5are the climate niche overla5 values for these h95othetical resident strategies to the climate niche overla5 the migrants actuall9 eN5erience between breeding and non-breeding grounds- 0dditionall9A to eNamine to what eNtent conditions in the non-breeding range differ from the conditions available on the breeding range during the non-breeding seasonA we calculated the climate niche overla5 between non-breeding conditions and the climate available on the breeding range during the non-breeding season-

.eographic pro)imity
&o evaluate if migrant Sylvia warblers move to the closest 5lace with suitable conditions based on niche Luantifications incor5orating both climate and land-coverA we used 5aired ttests to com5are the known migration distance with the distance from the breeding range to the closest suitable non-breeding area and the known migration distance with the distance from the non-breeding range to the closest suitable breeding area for each Sylvia s5ecies- We also divided the differences between known migration distance and the migration distances to the closest suitable area *i-e- distance between known breeding range and the closest nonbreeding area and the known non-breeding range and the closest breeding area, b9 the known migration distance to obtain s5ecies-s5ecific estimates of 5otential reductions in distance from ado5ting the shortest 5ossible migration route-

EB

1 2iche availability in space and time3 migration in Sylvia warblers

Seasonal changes in niche re5uirements


&o determine if the breeding niche of migrant Sylvia warblers is eLual to or smaller than the survival nicheA we calculated a 5aired WilcoNon signed rank test com5aring breeding niche breadth vs- total annual niche breadth for each Sylvia s5ecies- Niches here reflect both climate and land-cover simultaneousl9-

'.$ *esults
#rade4offs between niche breadth and migration distance
4ontrar9 to our 5redictionA movement distance between breeding and non-breeding grounds had no effect on the overla5 of breeding and non-breeding climate niches *7 U (-(((()A t U ])-(+A ! U (-+)A 82 U (-(BA Fig- @-'a, and on total annual climate niche breadth * 7 U ](-(((((('A t U -(-(BA ! U (-8<A 82 W (-()A Fig- @-'c, in Sylvia warblers- $and-cover niches showed the eN5ected relationshi5 to migration distance. Sylvia warblers with longer distances between breeding and non-breeding grounds did eNhibit a significantl9 lower overla5 in their land-cover niche *7 U -(-((((:A t U ]B-@@A ! U (-((()A 82 U (-BEA Fig- @-'b, and had significantl9 broader total annual land-cover niches *7 U (-((()8A t U '-<<A ! U (-()A 82 U (-'BA Fig- @-'d,- 0 few s5ecies migrating short distances deviated slightl9 from this 5attern due to the size difference between their ver9 small breeding ranges and larger non-breeding ranges *Fig- @-'bA scatter below regression line,- &here was no significant 5h9logenetic signal in an9 of the regression residuals *0bouheif tests. ! \ (-(@'H $ikelihood ratio tests for lambdaU(. ! \ (-@@,A indicating that our results are not affected b9 the 5h9logenetic relatedness of the s5ecies-

E@

1 2iche availability in space and time3 migration in Sylvia warblers

5igure '.2+ >elationshi5 between migration distance and *a, climate niche overla5 between breeding and non-breeding seasonA *b, land-cover niche overla5 between breeding and nonbreeding seasonA *c, total annual climate niche breadthA *d, total annual land-cover niche breadth for 'E s5ecies of Sylvia warblers-

EE

1 2iche availability in space and time3 migration in Sylvia warblers

2iche tracking
&he climate niche overla5 between breeding and non-breeding grounds for migrant Sylvia warblers was not significantl9 higher than if the9 had sta9ed either on the breeding grounds * t U (-@8A df U )'A ! U (-@E, or on the non-breeding grounds *t U ](-(+A df U )'A ! U (-8<, 9earround *Fig-@-+,- 4limate niche overla5 between the conditions migrant Sylvia warblers eN5erience on the non-breeding grounds and conditions available on their breeding grounds during the non-breeding season was low *& U (-)@ (-'@ *mean %D,A n U )+,-

5igure '."+ 4limate niche overla5 for different 5otential migration strategies in migrant Sylvia warblers- "igration does not lead to consistentl9 higher climate niche overla5 com5ared to resident strategies- migrate. niche overla5 between breeding and non-breeding grounds actuall9 eN5erienced b9 the s5eciesA sta9 breed. h95othetical niche overla5 resulting from sta9ing on the breeding grounds all 9earA sta9 non-breed. h95othetical niche overla5 resulting from sta9ing on the non-breeding grounds all 9earA dashed gre9 line U identit9 lineA n U )+-

E<

1 2iche availability in space and time3 migration in Sylvia warblers

.eographic pro)imity
#ncor5orating both climate and land-cover in niche LuantificationsA known migration distances were significantl9 longer than distances between the known breeding ranges and the closest suitable non-breeding area *t U B-@@A df U )'A ! U W (-(()H Fig- @-B,- "igrant Sylvia warblers could migrate an average of ') '([ *mean %D, less distance b9 fl9ing to the closest suitable non-breeding area- %imilarl9A known migration distances were significantl9 longer than the distances between the known non-breeding range and the closest suitable breeding area *t U @-<'A df U )'A ! UW (-(()A Fig- @-B,- /otential reductions in distance for fl9ing to the closest suitable breeding area were on average '+ ):[ *mean %D, of the s5eciesP known migration distance-

5igure '.$+ "igration distance and distance to the closest suitable non-breeding and breeding areas in migrant Sylvia warblers- /oints below the identit9 line indicate actual migration distance is longer than the distance to the closest suitable areas- actual migration. average distance between known breeding and non-breeding rangesA closest non-breed. average distance between known breeding range and closest suitable non-breeding areaA closest breed. average distance between known non-breeding range and closest suitable breeding areaA dashed gre9 line U identit9 lineA n U )+-

E:

1 2iche availability in space and time3 migration in Sylvia warblers

Seasonal changes in niche re5uirements


&aking into account both climate and land-coverA breeding niche breadth in Sylvia warblers wasA as 5redictedA significantl9 smaller than total annual niche breadth *W U )BA n U 'EA ! W (-(()A Fig- @-@,- Note that in s5ecies whose non-breeding range is environmentall9 ver9 uniformA total annual niche breadth can actuall9 be smaller than breeding niche breadthA because including the non-breeding range decreases the evenness of occu5ied environments *scatter below regression lineA Fig- @-@,-

5igure '.'+ 4om5arison of total annual niche breadth and breeding niche breadth in Sylvia warblers- &he maIorit9 of 5oints above the identit9 line indicate breeding niche breadth is smaller than total annual niche breadth- Niche breadth incor5orates both climate and landcover- dashed gre9 line U identit9 lineA n U 'E-

E8

1 2iche availability in space and time3 migration in Sylvia warblers

'.' 3iscussion
We develo5ed a new framework to understand how organisms res5ond to s5atio-tem5orall9 d9namic niche s5ace *Fig- @-), and tested the framework on seasonal migration in Sylvia warblers- We showed that Sylvia migration does not a55ear to be driven b9 climate niche tracking and that it eN5oses s5ecies to greater variation in other niche dimensions such as land-coverA which are com5arativel9 static across seasons- Niches were narrower during the breeding seasonA demonstrating that niche reLuirements can be d9namic in time- We showed that the framework is useful for testing 5redictions about Sylvia warbler migration and for stimulating new Luestions which can 5otentiall9 be transferred to other ecological 5henomena-

&rade-offs between niche breadth and migration distance


=ur results indicate that Sylvia warblers do not com5ensate for the costs of a long migrator9 Iourne9 b9 closel9 tracking their 5referred climate- ?oweverA land-cover niche breadth increased with migration distanceA highlighting the decou5led s5atio-tem5oral availabilit9 of climate and land cover- B9 a55l9ing our framework to eNamine tracking of niche dimensions other than climate we show that it ma9 be im5ossible for Sylvia s5ecies to track both climatic and land-cover conditions at the same time- #f tracking climate involves longer movement in geogra5hic s5aceA this would likel9 result in broader land-cover niches- Future research should eN5lore how migrator9 behaviour is influenced b9 and evolves under such com5leN constraintsA and how climate and land cover translate into re5roductive out5ut- &his also raises im5ortant Luestions for the transferabilit9 of correlative s5ecies distribution models in s5ace and timeA which assume a constant correlation structure between environmental variables *"orin K $echowitzA '((:H Jimenez-Dalverde et al., '((8,We also note thatA es5eciall9 for s5ecies with large rangesA Luantif9ing the niche at the s5ecies level ignores that sub5o5ulations and individuals ma9 select s5ecific environments from the total available niche s5aceA e-g- short-distance vs- trans-%aharan migrant sub5o5ulations in S. atricapilla *%hirihai et al., '(()H Doswald et al., '((8,A and hence might eN5erience ver9 different niche overla5-

<(

1 2iche availability in space and time3 migration in Sylvia warblers

2iche tracking
"igration in Sylvia warblers is not driven b9 climate niche tracking because migrating Sylvia warblers eN5erienced Iust as much climatic variation between breeding and non-breeding grounds as the9 would through annual climatic variation if the9 were resident on either the breeding or non-breeding grounds- &he low niche overla5 between winter climate on the breeding and on the non-breeding grounds suggests that overwintering on the breeding grounds would reLuire tolerating deviation from the breeding conditions in a different direction of niche s5ace com5ared to migration- 0 resident strateg9 ma9 thus eN5ose migrator9 Sylvia warblers to coldA un5roductive winters which the s5ecies ma9 be less able to tolerate- ?enceA the advantage of migration might rather be higher re5roductive success and lower nest 5redation at higher latitudes and lower winter mortalit9 at tro5ical latitudes *B2hning-Gaese et al., '(((H 0lerstam et al., '((+H $emoine K B2hning-GaeseA '((+H Griebeler et al., '()(H "c1innon et al., '()(,Evidence for climate niche tracking between breeding and non-breeding ranges in birds has been ambiguous so farA with high overla5 between breeding and non-breeding climate in some s5ecies *Jose5h K %tockwellA '(((H "artinez-"e9er et al., '((BH Nakazawa et al., '((BH "arini et al., '()(, but not in others *"artinez-"e9er et al., '((BH Nakazawa et al., '((B,- &he framework allows us to go be9ond classif9ing bird s5ecies as 6niche trackers7 or 6niche switchers7 based solel9 on climate niche overla5 between their breeding and nonbreeding grounds- #t is crucial to eNamine which 5art of the total environmental niche s5ace is available when and where to understand whether s5ecies ? movements are indeed driven b9 environmental d9namics in 5articular niche dimensions *>eside et al., '()(,- B9 taking the s5atio-tem5oral availabilit9 of environments into accountA we can com5are the niche overla5 resulting from alternative h95othetical migration strategies and evaluate if an observed migration 5attern minimises niche overla5-

.eographic pro)imity
Both during the s5ring and autumn migrationsA Sylvia warblers fl9A on averageA '([ further than to the nearest area with suitable climate and land-cover- ?enceA migration distance is likel9 driven b9 additional factors- 0lthough we Luantified niche characteristics from occurrence records and the9 thus reflect the realised nicheA we do not eN5licitl9 address the influence of biotic interactions on occu5ied niche s5ace and geogra5hic distributionsA which <)

1 2iche availability in space and time3 migration in Sylvia warblers has been demonstrated for bird migration regarding diffuse com5etition with residentsA 5arasitism and nest 5redation *0lerstam et al., '((+H $emoine K B2hning-GaeseA '((+H "c1innon et al., '()(,- 0n im5erfect o5timisation of migration distance could also be the result of genetic constraints on the migration routes of bird 5o5ulations or of the influence of geogra5hic barriers and the availabilit9 of sto5-over sites on the cost of migrating along s5ecific routes *0lerstam et al., '((+H Doswald et al., '((8,-

Seasonal changes in niche re5uirements


=ur findings suggest that Sylvia warblers onl9 breed in a subset of their total annual niche s5aceA which is congruent with the idea that more eNacting reLuirementsA in terms of energ9A nutrients and 5rotection from 5redation during re5roductionA ma9 lead to smaller niches *GrubbA )8<<H 0lerstam K ?2gstedtA )8:'H &iteuN et al., '((<,- #n order to not obscure such seasonal niche shiftsA s5ecies distribution models should be fitted with seasonal subsets of occurrence and environmental data *?eikkinen et al., '((E,- =ur findings also indicate that niches should be regarded as d9namic entities over the life c9cle of s5ecies and that closer eNamination of how niches var9 during s5ecific 5hases of the life c9cle of organisms ma9 be beneficial for future studies aiming to model niches and distribution *Jackson et al., '((8,-

mplications
We have shown that our conce5tual framework *Fig- @-), is a useful starting 5oint to understand how the d9namics of environmental conditions in s5ace and time and d9namic niche reLuirements affect the niches and distributions of organisms- &he framework goes be9ond 5revious studies of niche tracking b9 taking the d9namic availabilit9 of niches in s5ace and time into account- #t is thus a useful a55roach to identif9 the niche dimensions that are crucial in sha5ing organisms? movements and it describes fundamental 5rocesses and constraints which are a55licable to a broad range of ecological 5henomena and taNa- For eNam5leA the framework could be used to test how diel vertical migration in zoo-5lankton is linked to the dail9 variation of tem5erature and sunlight in the water column or to eN5lore to what eNtent 5lant dormanc9A which could be seen as an eNtreme case of d9namism in s5ecies ? niche reLuirementsA is a res5onse to changes in the availabilit9 of 5articular niche dimensions&he framework is relevant for 5redicting how s5ecies ranges will res5ond to long-term tem5oral changes in conditions caused b9 climate change- &racking climate change might increasingl9 des9nchronise s5ecies with other niche dimensions such as land-cover- =ur <'

1 2iche availability in space and time3 migration in Sylvia warblers results for the Sylvia warblers suggest the9 might not track the 5redicted shifts in their 5referred climatic conditions *Barbet-"assin et al., '((8H Doswald et al., '((8,- 055l9ing the framework to animal migration highlights that niche tracking behaviour is com5leN and that assuming sim5le climate tracking when 5redicting future range shifts ma9 be too sim5listic=ur framework is 5art of an emerging trend to im5rove the mechanistic understanding of macroecological 5rocesses through anal9ses of tem5oral d9namics *Fisher et al., '()(,- We suggest that future studies aiming to model niches and distribution ma9 benefit from making use of high tem5oral resolution of occurrence and environmental data instead of using tem5oral averages as in5ut *?eikkinen et al., '((EH Jackson et al., '((8,- While the s5ecies distribution modelling 5aradigm allows the eNamination of niches and ranges at several individuall9 modelled 5oints in timeA it would be interesting to eN5lore the 5otential conseLuences of changes in niche availabilit9 and niche reLuirements in a d9namic model */agel K %churrA '()),- 055l9ing our framework to bird migration shows how we can significantl9 enhance our understanding of the drivers behind s5atial movements of organisms and the d9namics in their realised niches b9 taking changing niche reLuirements throughout organism?s life-c9cles and the s5atio-tem5oral availabilit9 of environments into account-

<+

9 ,onclusions

. 6>(6C;-0>(#n this thesisA # have attem5ted to advance our current understanding of avian range d9namics b9 modelling 5utative mechanistic linksA integrating multi5le drivers in the same anal9sisA and b9 develo5ing new conce5tual ideas- &o this endA # have combined datasets from ecolog9A evolution and the Earth sciences and used tools from information science such as geogra5hic information s9stemsA statistical 5rogramming environmentsA 5ath anal9sisA boosted regression treesA ridge regressionA bootstra55ing techniLuesA kernel estimators and niche metrics4orres5onding to the 5revious three cha5tersA the following maIor findings emerge. *i, %5ecies? traits do not account for the maIorit9 of the variation in range sizeA but the9 do 5la9 an im5ortant role- %everal traits influence range size simultaneousl9 in com5leN wa9sA both directl9 and indirectl9 through other traits- ?igh annual fecundit9A high dis5ersal abilit9A broad habitat nichesA low tro5hic levelA large bod9 size and being migrator9 emerge as the most im5ortant traits leading to large global ranges in Euro5ean 5asserines- *ii, Whether s5ecies are able to colonise 5otentiall9 suitable areas at large s5atial and tem5oral scales de5ends on multi5le drivers that interact with each other- ?igh dis5ersal abilit9 onl9 enables Sylvia warblers to fill a high 5ro5ortion of their 5otential range if s5ecies richness of congenerics in areas of low habitat suitabilit9 within their 5otential range is low- 4ontrar9 to 5revious ideas on the scale-de5endence of drivers of range d9namics */earson K DawsonA '((+H Guisan K >ahbekA '()),A this suggests that biotic interactions ma9 be im5ortant in sha5ing ranges at the continental scale and that their effect is likel9 contingent on habitat suitabilit9- *iii, %5ecies? niches and distributions are d9namic entities that de5end on the s5atio-tem5oral availabilit9 of environmental conditions- &he availabilit9 of niche dimensions ma9 be des9nchronisedA challenging s5ecies with com5leN o5timisation 5roblems when tr9ing to track them in geogra5hic s5ace- Niche reLuirements of s5ecies ma9 var9 throughout their life-c9cle"igration in Sylvia warblers is not driven b9 climate niche tracking and results in broader land-cover niches- Sylvia warblers had narrower niches during the breeding season- #n summar9A the earl9 idea that ranges are set b9 ver9 few factors *e-g- &wome9A )8+E, can be reIected- &he 5rocesses that determine the sizeA geogra5hic location and the d9namic of s5ecies distributions are highl9 com5leN and involve multi5le interacting drivers- We are Iust beginning to form a coherentA com5rehensive view of range d9namics-

<B

9 ,onclusions

..1 5uture research !ers!ectives


#t is im5ortant to kee5 in mind thatA of necessit9A the results in this thesis were obtained using different 5asserine grou5s as model s9stems- 0s a conseLuenceA the validit9 of those results remains restricted to these model grou5s until confirmed or reIected for other taNaNeverthelessA this thesis 5rovides anal9sis a55roachesA 5redictions and conce5tual ideas that can be a55lied to other grou5s of birds andA indeedA other taNa from all kingdoms of life%tudies integrating multi5le 5otential drivers of range d9namics are still rare *GastonA '((+,#t is too earl9 to comment on the likel9 differences in range d9namics across a broad taNonomic s5ectrumA but given the results from this thesisA one might formulate the cautious eN5ectation that range d9namics in other taNa will be eLuall9 com5leNA but that the relative im5ortance of different drivers is likel9 taNon-s5ecific- For eNam5leA it has been noted thatA on averageA 5lants and insects have smaller geogra5hic ranges than vertebrates *GastonA )88B, and birds generall9 have larger geogra5hic ranges than mammals *0ndersonA )8:B,?95otheses involving dis5ersal abilit9A bod9 size and habitat use have been 5ro5osed as eN5lanations *0ndersonA )8:B,A but this 5attern has so far not been demonstrabl9 linked to characteristics of these animal grou5s- %tud9ing the Luestions raised in this thesis for other taNa seems a worthwhile endeavour>ange d9namics are com5leN because a s5ecies? geogra5hic distribution is an emergent 5ro5ert9 of a number of highl9 elaborate subs9stems such as 5o5ulation geneticsA meta5o5ulation d9namics and communit9 ecolog9H all influenced b9 short-term environmental d9namicsA long-term environmental change and historic effects *GastonA '((+H NewtonA '((+H B2hning-Gaese et al.A '((EH /rice K 1irk5atrickA '((8,- Few of the dataA 5atterns and 5otential relationshi5s are directl9 observable b9 an9 individual researcher and man9 cannot be studied b9 eN5erimentation *BrownA )88@,- #t a55ears that there might be a trade-off between breadth and de5th- %tudies at large s5atio-tem5oral scales that include man9 s5ecies sometimes ma9 become divorced from the local ecolog9 and histor9 of the individual taNa *Wiens K DonoghueA '((B,- For eNam5leA for a com5arative anal9sis of the global ranges of all Euro5ean 5asserines as in cha5ter threeA one has to resort to relativel9 broad classifications to characterise the s5ecies? habitat and diet 5references- =n the other handA detailed studies of occu5anc9A 5o5ulation d9namics and biotic interactions can be so restricted in taNonomic and geogra5hic focus that it is hard to know how to generalise be9ond regarding <@

9 ,onclusions them as a collection of single case studies *BrownA )88@H $adle K WhittakerA '()),H e-gstudies investigating intrageneric biotic interactions in Sylvia warblers t95icall9 cover s5atial eNtents below '-@ km' *e-g- 4od9 K WalterA )8<EH 4od9A )8<:H GarciaA )8:+H "artin K &hibaultA )88EH ElleA '((+H /ons et al.A '((:,- =ne fundamental challenge facing research on s5ecies distributions is to s9nthesise these two 5ers5ectives */earson K DawsonA '((+H Wiens K DonoghueA '((BH Guisan K >ahbekA '()),#n working towards such a s9nthesis of breadth and de5th in range d9namicsA there are two current trends. one is to combine data from ecolog9A evolution and geoscience and the other is to integrate an increasing number of factors and more com5leN relationshi5s into the same Luantitative frameworkA which then allows us to look at the emergent 5ro5erties of s9stems that are too com5leN to be antici5ated b9 the human mind- #n this thesisA # have 5ursued both a55roachesH the first a55roach 5articularl9 in cha5ters B and @A and the second a55roach 5articularl9 in cha5ter +- &wo 5otential difficulties in further develo5ing this line of research have become a55arent. *i, Even when working with eNce5tionall9 well-studied taNaA data availabilit9 Luickl9 becomes an issue- &he $innean and Wallacean shortfallsA the incom5leteness of our knowledge of s5ecies and their distributions *$omolino et al., '()(,A are acutel9 felt- For the Sylvia warblersA advances in our taNonomic knowledge have resulted in s5ecies being attributed to the genus as recentl9 as '((8 *Doelker et al., '((8, and there is a geogra5hic bias in the Lualit9 of the information on s5ecies? rangesA with data for North 0frica and the "iddle East being notabl9 5oorer than for Euro5e *%hirihai et al., '((),- 0lsoA to full9 understand the d9namic nature of avian geogra5hic ranges we will need to build sufficientl9 standardised and integrated datasets about tem5oral changes in the abundance structure within the area we 5resentl9 consider a s5ecies? geogra5hic range and about migrator9 movements *Wikelski K 1a9sA '()),- >egarding 5otential drivers of range d9namicsA com5utationall9 accessible Luantitative information on man9 s5ecies? traitsA such as dis5ersal abilit9 or brain sizeA is still limited to 5articular bird grou5sA and some driversA such as biotic interactionsA are eNceedingl9 difficult to Luantif9- *ii, Develo5ing models that retain breadth *i-e- a55l9 to man9 s5ecies over large scales, without loosing de5th *i-e- take into account the local ecolog9 of individual s5ecies, im5lies higher model com5leNit9- ?ence the models need large amounts of data when we want to test h95otheses about the factors driving range d9namics- Even if we have com5arativel9 good data for a grou5A we ma9 be faced with the conundrum that the grou5 ma9 sim5l9 not have enough members to result in a

<E

9 ,onclusions sam5le size with enough degrees of freedom to test more com5leN h95otheses- /urel9 5ractical constraints that are still of relevance when conducting studies on range d9namics are that 5re5aration and integration of data from different sources into a common geogra5hical and statistical environment can be ver9 time-consuming and increasingl9 com5leN models ma9 sim5l9 take a ver9 long time to runFinall9A # want to 5oint out several areas where the incor5oration of additional com5leNit9 has the 5otential to advance our understanding of s5ecies? distributions and which should be considered when tr9ing to 5redict range shifts under global change. *i, &here is still am5le sco5e to integrate more of the drivers of range d9namics into one anal9sis *Botkin et al.A '((<,- For eNam5leA it might be interesting to incor5orate all of the s5ecies traits identified as im5ortant determinants of range size in cha5ter + into anal9ses similar to those 5resented in cha5ter BA which attem5t to assess how im5ortant s5ecies? traits are relative to other 5otential drivers of range d9namicsA such biotic interaction and historic constraints- Ba9esian methods ma9 be a 5romising route for the simultaneous anal9sis of multi5le drivers as the9 allow for great model com5leNit9 andA b9 using 5riorsA can take information into account that is difficult to integrate into other methods *EllisonA '((BH 4ho9 et al., '((8,- *ii, 4ha5ter B highlights that the mechanisms which facilitate or 5revent colonisation of suitable habitat are contingent on the environmental conditions of the area under consideration- &his is consistent with studies of range edges which demonstrate that range limits for the same s5ecies can be set b9 different mechanisms in different 5laces *e-g- BarnesA )8@<H Gross K /riceA '(((,- #t ma9 be fruitful to eN5lore techniLues such as geogra5hicall9 weighted regression and make more use of the ca5abilities of data-driven s5ecies distribution modelling methods such as boosted regression trees to visualise com5leN interactions *0ustinA '((<H Elith et al., '((:,- *iii, We should be cautious about assuming that biotic interactions can be safel9 ignored for 5redicting range shifts at large scales *Brooker et al., '((<,- &here is an increasing trend to take biotic interactions into account in s5ecies distribution modelling b9 using the occurrence of other s5ecies as 5redictors *0raOIo K $uotoA '((<H ?eikkinen et al., '((<H /reston et al.A '((:,- #f biotic interactions are im5ortant at large scalesA s5ecies cannot be modelled in isolation when tr9ing to 5redict range shiftsA and methods which iterativel9 consider shifts in several s5ecies have to be develo5ed *1eith et al., '((:H Baselga K 0raOIoA '((8,- *iv, 4ha5ter @ 5rovides an eNam5le of how the in-de5th consideration of the ecolog9 of a 5articular s5ecies grou5 can stimulate the develo5ment of conce5tual frameworks and ideas that are relevant for man9 taNa

<<

9 ,onclusions and ecological 5henomena- &he d9namic nature of niche reLuirements demonstrated in cha5ter @ indicates that ranges for different life-c9cle stages ma9 have to be modelled se5aratel9 to 5ro5erl9 account for the associated change in s5ecies? niches *Doswald et al., '((8H Jackson et al., '((8,- 0lsoA understanding how s5ecies deal with s5atio-tem5oral des9nchronisation in different niche dimensions ma9 hel5 us understand how the9 will res5ond to the 5redicted future 5revalence of novel climates *Williams et al., '((<,- While we discuss the framework mainl9 in the conteNt of short-term tem5oral d9namicsA it can in 5rinci5le be eLuall9 a55lied to how ranges and niches change in the course of evolution and to eNamine 5henomena such as niche conservatism *Wiens K GrahamA '((@H 4ris5 et al., '((8,- Focusing on the strategies s5ecies can develo5 in res5onse to the d9namic availabilit9 of environmental conditions can 5otentiall9 9ield new insights into how organisms ma9 deal with environments that are d9namic both on ecological and evolutionar9 time-scales *Fisher et al., '()(H /earman et al., '((:,-

..2 6oncluding remarks


>ange d9namics are com5leN- 4onsidering the scale of our ignorance and the data reLuired to remed9 itA it seems doubtful whether our abilit9 to make meaningful 5redictions of the effect of global change on biodiversit9 will im5rove sufficientl9 within the time-frame of a few 9ears that is relevant for 5olitical action and management decisions *"E0A '((@bH %4BDA '()(,- &he urgenc9 of making wide-ranging societal decisions sha5ing a traIector9 of global change that cannot be controlled or reversed *?annah et al., '(('H 1ingA '((@H %ternA '((E, contrasts with the inevitabl9 smallA incremental ste5s and recursive discussions in which science can deci5her wh9 organism are distributed the wa9 the9 are *$adle K WhittakerA '()),"ore research on the d9namics that give rise to s5ecies ? geogra5hic distributions is urgentl9 needed *Davis et al.A )88:H "E0A '((@bH /armesanA '((EH %4BDA '()(,Notwithstanding 5ressure to obtain funding in a scientific s9stem whose members are increasingl9 5art of the 5recariatA ecologists should be cautious about enabling the notion that global change im5acts can be 5recisel9 5redicted and managed b9 overstating the certaint9 and 5ractical relevance of their 5redictions *%utherst et al., '((<H Willis K BhagwatA '((8H %inclair et al., '()(,- ELuall9A the9 should be war9 of 5roviding eNcuses for societal agents to 5ost5one difficult decisions b9 eNaggerating the imminence of results that would allow such <:

9 ,onclusions management *GlanzA )8::H ?ulmeA '((@H %utherst et al., '((<H %4BDA '()(,- ?ighlighting what we 5resentl9 do not know and cannot 5redict ma9 encourage societal discourse about risk acce5tance and 5lanning under uncertaint9 *Dasgu5taA '((:H 44%/A '((8H Dawson et al.A '()),-

<8

: 8eferences

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: 8eferences 4aseA &- D-A ?oltA >- D-A "c/eekA "- 0- K 1eittA &- ?- *'((@, &he communit9 conteNt of s5eciesP borders. ecological and evolutionar9 5ers5ectives- <ikosA 192A ':-BE4atch5oleA 4- 1- *)8<:, #nters5ecific territorialism and com5etition in %crocephalus warblers as revealed b9 5la9back eN5eriments in areas of s9m5atr9 and allo5atr9- %nimal *ehaviourA 2.A )(<'-)(:(44%/A "organA "- G-A DowlatabadiA ?A ?enrionA "-A 1eithA D-A $em5ertA >-A "cBrideA %-A %mallA "-A K WilbanksA &- *eds-, *'((8, *est !ractice %pproaches for ,haracteri$ing, ,ommunicating, and ncorporating Scientific Uncertainty in ,limate &ecision (aking. 0 >e5ort b9 the 4limate 4hange %cience /rogram and the %ubcommittee on Global 4hange >esearch- National =ceanic and 0tmos5heric 0dministrationA WashingtonA D44ha5inA F- %-A 3avaletaA E- %-A EvinerA D- &-A Na9lorA >- $-A DitousekA /- "-A >e9noldsA ?- $-A ?oo5erA D- -A et al. *'(((, 4onseLuences of changing biodiversit9- 2atureA $9'A '+B'B'4hongA D- $- %-A "ouginA E- K Gastellu-Etchegorr9A J- /- *)88+, >elating the global vegetation indeN to net 5rimar9 5roductivit9 and actual eva5otrans5iration over 0fricanternational =ournal of 8emote SensingA 1$A )@)<-)@BE4ho9A $-A =?$ear9A %- J-A "engersenA >- 0- K 1errieA $- *'((8, Elicitation b9 design in ecolog9. using eN5ert o5inion to inform 5riors for Ba9esian statistical models- ;cologyA /9A 'E@-'<<4larkA G- 0- *)8<8, Bod9 weights of birds - review- ,ondorA 21A )8+-'('4od9A "- $- *)8<:, ?abitat selection and inters5ecific territorialit9 among the s9lviid warblers of England and %weden- ;cological (onographsA $2A +@)-+8E4od9A "- $- K WalterA ?- *)8<E, ?abitat selection and inters5ecific interactions among "editerranean s9lviid warblers- <ikosA 24A ')(-'+:4olwellA >- 1- K Futu9maA D- J- *)8<), "easurement of niche breadth and overla5- ;cologyA '2A @E<-@<E4olwellA >- 1- K >angelA &- F- *'((8, ?utchinson?s dualit9. &he once and future niche !roceedings of the 2ational %cademy of Sciences if the United States of %merica, 19.A )8E@)-)8E@:4onnorA E- F- K BowersA "- 0- *)8:<, &he s5atial conseLuences of inters5ecific com5etition%nnales Zoologici CenniciA 2$A ')+-''E4rawle9A "- J- *'((<, #he 8 *ook. Wile9A 4hichester4ris5A "- D-A 0rro9oA "- &- 1-A 4ookA $- G-A GandolfoA "- 0-A JordanA G- J-A "cGloneA "- %-A WestonA /- ?-A et al. *'((8, /h9logenetic biome conservatism on a global scale- 2atureA $'2A <@B-<@E4rutzenA /- J- *'((', Geolog9 of mankind- 2ature, $1'A '+4unninghamA ?- >-A >isslerA $- J- K 05odacaA J- J- *'((8, 4om5etition at the range boundar9 in the slim9 salamander. using reci5rocal trans5lants for studies on the role of biotic interactions in s5atial distributions- =ournal of %nimal ;cologyA 42A @'-E':+

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: 8eferences $eislerA B- K WinklerA ?- *'((+, "or5hological conseLuences of migration in 5asserines%vian (igration *eds- /- BertholdA E- Gwinner K E- %onnenschein,A 55- )<@-):E%5ringerA Berlin$emoineA N- K B2hning-GaeseA 1- *'((+, /otential im5act of global climate change on s5ecies richness of long-distance migrants- ,onservation *iology, 14A @<<-@:E$esterA %- E-A >uttenbergA B- #-A GainesA %- D- K 1inlanA B- /- *'((<, &he relationshi5 between dis5ersal abilit9 and geogra5hic range size- ;cology -ettersA 19A <B@-<@:$iuA 4-A Berr9A /- "-A DawsonA &- /- K /earsonA >- G- *'((@, %electing thresholds of occurrence in the 5rediction of s5ecies distributions- ;cographyA 22A +:@-+8+$oboA J- "-A Jimenez-DalverdeA 0- K >ealA >- *'((:, 0 4. a misleading measure of the 5erformance of 5redictive distribution models- .lobal ;cology and *iogeographyA 14, )B@-)@)$omolinoA "-D- K >iddleA B- >-A WhittakerA >- J- K BrownA J- ?- *'()(, *iogeographyA Bth edn- %inauerA %underlandA "0$ovetteA #- J- K ?ochachka W- "- *'((E, %imultaneous effects of 5h9logenetic niche conservatism and com5etition on avian communit9 structure- ;cologyA 24A %)B-%':$ubzensA E-A 4erdaA J-A 4larkA "- *eds-, *'()(, &ormancy and 8esistance in 'arsh ;nvironmentsA %eries. &o5ics in 4urrent GeneticsA Dol- ')A %5ringerA ?eidelberg"ac0rthurA >- ?- *)8<', .eographical ;cology3 !atterns in the &istribution of Species/rinceton niversit9 /ressA /rinceton"anelA %-A WilliamsA ?- 4- K =rmerodA %- J- *'((), Evaluating 5resence-absence models in ecolog9. the need to account for 5revalence- =ournal of %pplied ;cologyA "2A 8')-8+)"ariniA "- f-A Barbet-"assinA "-A "artinezA J-A /restesA N- /- K JiguetA F- *'()(, 055l9ing ecological niche modelling to 5lan conservation actions for the >ed-s5ectacled 0mazon *%ma$ona pretrei,- *iological ,onservation, 1$"A )('-))'"armionA "-A $uotoA "-A ?eikkinenA >- 1- K &huillerA W- *'((8, &he 5erformance of stateof-the-art modelling techniLues de5ends on the geogra5hical distribution of s5ecies;cological (odellingA 229A +@)'-+@'("artinA J- $- K &hibaultA J- 4- *)88E, 4oeNistence in "editerranean warblers. Ecological differences or inters5ecific territorialit9` =ournal of *iogeographyA 2"A )E8-)<:"artinez-"e9erA E-A /etersonA 0- &- K Navarro-%iguenzaA 0- G- *'((B, Evolution of seasonal ecological niches in the /asserina buntings *0ves . 4ardinalidae,- !roceedings of the 8oyal Society * *iological Sciences, 241A ))@)-))@<"c1innonA $-A %mithA /- 0-A NolA E-A "artinA J- $-A Do9leA F- #-A 0brahamA 1- F-A GilchristA ?G-A "orrisonA >- #- G- K Bgt9A J- *'()(, $ower 5redation risk for migrator9 birds at high latitudes- Science, "24A +'E-+'<-

:8

: 8eferences "illennium Ecos9stem 0ssessment *'((@a, 4ha5ter )). Biodiversit9 >egulation of Ecos9stem %ervices- #n. ?assan >A %choles >A 0sh N *eds-,A ;cosystems and 'uman Well4being3 ,urrent State and #rends, 0olume 1, Cindings of the ,ondition and #rends Working .roup- #sland /ressA WashingtonA D-4-A 55- '8<-+'8"illennium Ecos9stem 0ssessment *'((@b,- ;cosystems and 'uman Well4being3 Synthesis. #sland /ressA WashingtonA D4"ilner-GullandA E- J-A Fr9NellA J- "- K %inclairA 0- >- E- *eds-, *'()), %nimal (igration3 % SynthesisA =Nford niversit9 /ressA =Nford"itchellA >- J- *)88', &esting evolutionar9 and ecological h95otheses using 5ath-anal9sis and structural eLuation modelling- Cunctional ;cologyA .A )'+-)'8"ooreA 1- 0- *'((8, Fluctuating 5atch boundaries in a native annual forb. the roles of niche and dis5ersal limitation- ;cologyA /9A +<:-+:<"orinA h- K $echowitzA "- J- *'((:, 4ontem5orar9 5ers5ectives on the niche that can im5rove models of s5ecies range shifts under climate change- *iology -ettersA $A @<+@<E"unguiaA "-A /etersonA 0- &- K %anchez-4orderoA D- *'((:, Dis5ersal limitation and geogra5hical distribution of mammal s5ecies- =ournal of *iogeographyA "'A ):<8-)::<NakazawaA C-A /etersonA 0- &-A "artinez-"e9erA E- K Navarro-%iguenzaA 0- G- *'((B, %easonal niches of Nearctic-Neotro5ical migrator9 birds. #m5lications for the evolution of migration- %uk, 121A E)(-E):NewA "-A ?ulmeA "- K JonesA /- *'(((, >e5resenting twentieth-centur9 s5ace-time climate variabilit9- /art ##. Develo5ment of )8()-8E monthl9 grids of terrestrial surface climate=ournal of ,limateA 1"A '')<-''+:NewtonA #- *'((+, Speciation and *iogeography of *irds. 0cademic /ressA $ondonNorbergA - "- *)8:8, 0ertebrate Clight, (echanics, !hysiology, (orphology, ;cology and ;volution- %5ringerA Berlin=lsonA D- "-A DinersteinA E-A Wikramana9akeA E- D-A BurgessA N- D-A /owellA G- D- N-A nderwoodA E- 4-A D?0micoA J- 0-A #touaA #-A %trandA ?- E-A "orrisonA J- 4-A $oucksA 4J-A 0llnuttA &- F-A >ickettsA &- ?-A 1uraA C-A $amoreuNA J- F-A WettengelA W- W-A ?edaoA /K 1assemA 1- >- *'((), &errestrial ecoregions of the world. 0 new ma5 of life on Earth- *ioscienceA '1A 8++-8+:=rmeA 4- D- $-A DaviesA >- G-A =lsonA D- 0-A &homasA G- ?-A DingA &- %-A >asmussenA /- 4-A >idgel9A >- %-A %tattersfieldA 0- J-A BennettA /- "-A =wensA #- /- F-A BlackburnA &- "- K GastonA 1- J- *'((E, Global 5atterns of geogra5hic range size in birds !-<S *iologyA $A e'(:/agelA J- K %churrA F- "- *'()), Forecasting s5ecies ranges b9 statistical estimation of ecological niches and s5atial 5o5ulation d9namics- .lobal ;cology and *iogeographyA doi. )(-))))TI-)BEE-:'+:-'())-((EE+-N /agelA "- *)88<, #nferring evolutionar9 5rocesses from 5h9logenies- Zoologica Scripta, 2.A ++)-+B:8(

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8)

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8'

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8+

: 8eferences DirkkalaA >- *)88+, >anges of northern forest 5asserines - a fractal anal9sis- <ikosA .4A '):''EDoelkerA G- K $ightA J- E- *'()), /aleoclimatic eventsA dis5ersal and migrator9 losses along the 0fro-Euro5ean aNis as drivers of biogeogra5hic distribution in Sylvia warblers*(, ;volutionary *iologyA 11A)E+DoelkerA G-A "eloA "- K BowieA >- 4- 1- *'((8, 0 Gulf of Guinea island endemic is a member of a "editerranean-centered bird genus- bisA 1'1A @:(-@:+WallaceA 0- >- *):<E, #he .eographical &istribution of %nimalsD With % Study of the 8elations of -iving and ;)tinct Caunas as ;lucidating the !ast ,hanges of the ;arths Surface- "acmillan K 4o-A $ondonWarrenA D- $-A GlorA >- E- K &urelliA "- *'((:, Environmental niche eLuivalenc9 versus conservatism. Luantitative a55roaches to niche evolution- ;volution, .2A ':E:-'::+WebbA &- J-A K GastonA 1- J- *'(((, Geogra5hic range size and evolutionar9 age in birds!roceedings of the 8oyal Society * *iological SciencesA 2.4A ):B+-):@(WhiteA $- *)8E<, &he ?istorical >oots of =ur Ecological 4risis- Science 1''A )'(+-)'(<. WiensA J- 0- *)8:8, %5atial scaling in ecolog9- Cunctional ;cologyA "A +:@-+8<WiensA J- J- K DonoghueA "- J- *'((B,- ?istorical biogeogra5h9A ecolog9 and s5ecies richness- #rends in ;cology and ;volutionA 1/A E+8-EBBWiensA J- J- K GrahamA 4- ?- *'((@, Niche conservatism. #ntegrating evolutionA ecolog9A and conservation biolog9- %nnual 8eview of ;cology, ;volution and SystematicsA ".A @)8@+8WiensA J- J-A 0ckerl9A D- D-A 0llenA 0- /-A 0nackerA B- $-A Buckle9A $- B-A 4ornellA ?- D- et al. *'()(, Niche conservatism as an emerging 5rinci5le in ecolog9 and conservation biolog9- ;cology -etters, 1"A )+)(-)+'BWikelskiA "- K 1a9sA >- *'((), "ovebank. archiveA anal9sis and sharing of animal movement data- World Wide Web electronic 5ublication- htt5.TTwww-movebank-orgWikelskiA "-A &arlowA E- "-A >aimA 0-A DiehlA >- ?-A $arkinA >- /- K DisserA G- ?- *'((+, 0vian metabolism. 4osts of migration in free-fl9ing songbirds- 2ature, $2"A <(B-<(BWilliamsA J- W-A JacksonA %- &- K 1utzbachA J- E- *'((<, /roIected distributions of novel and disa55earing climates b9 ')(( 0D- !roceedings of the 2ational %cademy of Sciences, 19$, @<+:-@<B'. WilliamsonA 4- E-A FisherA J- "-A BollensA %- "-A =verholt E- /- K Breckenridge J- 1- *'()), &oward a more com5rehensive theor9 of zoo5lankton diel vertical migration. #ntegrating ultraviolet radiation and water trans5arenc9 into the biotic 5aradigm-imnology and <ceanography, '.A )E(+-)E'+WillisA 1- J- K BhagwatA %- 0- *'((8, Biodiversit9 and 4limate 4hange- Science, 2$2A :(E:(<WinklerA ?- K $eislerA B- *)88', =n the ecomor5holog9 of migrants- bisA 1"$A ')-':8B

: 8eferences WiszA "- %-AWaltherA B- 0 K >ahbek 4- *'((<, sing 5otential distributions to eN5lore determinants of Western /alaearctic migrator9 songbird s5ecies richness in sub-%aharan 0frica- =ournal of *iogeography, "$A :':-:B)CoungA &- /-A /etersenA D- 0- K 4lar9A J- J- *'((@, &he ecolog9 of restoration. historical linksA emerging issues and uneN5lored realms- ;cology -ettersA 2A EE'-E<+-

8@

E %cknowledgements

2 A6

(>@C<3,<1<(=-

=ur achievements are not our own- &he9 grow from the fabric of su55ort and o55ortunit9 created in familiesA work5lacesA communitiesA social and 5rofessional networks and the societies we live in- &he work 5resented in this thesis was made 5ossible onl9 b9 the su55ort and contribution of numerous other 5eo5le and institutions- # am most grateful to all of them/rof- Dr- 1atrin B2hning Gaese acted as a 5rinci5al su5ervisor for this thesis and su55orted it through her guidanceA ideasA comments and constructive criticism and her remarkable willingness to alwa9s make the time to discuss science or give feedback on a manuscri5t- %he 5rovided the institutional setting for this work with 5articular financial su55ort from the $=EWE eNcellence initiative b9 the ?essian "inistr9 for %cience and the 0rts/rof- Dr- 4atherine ?- Graham kindl9 agreed to act as co-su5ervisor and su55orted me through scientific ideas and discussionsA her talent for scientific networkingA her generous hos5italit9 and b9 regularl9 giving me a nudge in the right direction via %k95e and email# have learned a lot from both of them/rof- Dr- 4arsten >ahbek contributed hel5ful comments and data on global geogra5hic bird ranges- %usanne Fritz eNtracted the global range data from the original database and was alwa9s ha559 to let me 5ick her brains- # would like to thank ?eiko 1orntheuer for granting me full access to his work and for being so well-organised- "onika %chwager and %ven &rautmann contributed statistical and methodological eN5ertise and hel5ful comments on cha5ter +- $luds Brotons and %hai "eiri made hel5ful comments on manuscri5t drafts- =livier Broennimann engaged in stimulating discussions on niche metrics and shared his > codeGerald "a9r from the %enckenberg "useum Frankfurt and %9lke Frahnert and /ascal Eckhoff from Berlin Natural ?istor9 "useum 5rovided access to bird collections# would like to thank 5resent and 5ast members of the ecolog9 lab at the Johannes Gutenbergniversit9 "ainz and the Biodiversit9 and 0rea D9namics of Dertebrates 5roIect grou5 at the Biodiversit9 and 4limate >esearch 4entre *Bi1-F, j in 5articular NilsA 1athrinA "onikaA DanaA "atthiasA EviA "atzeA JohannaA "arcA &anIaA %usanneA 4hristianA %venA 4laudiaA %venIaA NinaA &onka and Birgit j for friendshi5A scientific and technical su55ortA hel5ful feedback on

8E

E %cknowledgements talks and manuscri5ts and a great working atmos5here# gratefull9 acknowledge the financial andA in 5articularA invaluable idealistic su55ort from the German National "erit Foundation which # had the 5rivilege of enIo9ing throughout m9 undergraduate and 5ostgraduate studies and during a doctoral scholarshi5- /rof- Dr- Dolker Bach has acted as m9 tutor in all this time- # am dee5l9 humbled that the German 5eo5le gave of their wages to su55ort m9 education and workDuring m9 /hDA # also received financial and idealistic su55ort from the ELual =55ortunities =ffice of the Johannes Gutenberg- niversit9 "ainzA the #nternational Biogeogra5hic %ociet9 and the Biogeogra5h9 >esearch Grou5 of the >o9al Geogra5hic %ociet905art from the work 5resented in this thesisA # had the o55ortunit9 to contribute m9 eN5ertise to a number of collaborative 5roIects ; including field work on avian seed dis5ersalH modelling the 5astA 5resent and future ranges of aLuatic invertebratesH assessing 5otential climate change im5acts on Euro5ean birds and Luantif9ing avian dis5ersal abilit9 ; which so far resulted in @ 5ublications and B manuscri5ts- # am grateful to 1athrin &heissingerA Nils BreitbachA Britta DawideitA 0llie /hillimoreA "atthias %chleuningA "onika %chwagerA %ven &rautmannA %teffen - /aulsA Julia &aubmannA Franz BadeckA "iklos Bilint and ?olger ?erl9n for 5roductive and enIo9able collaborations and for broadening m9 scientific horizon# am dee5l9 grateful to m9 5arentsA #ngeborg and >olf $aubeA for their loveA encouragement and their generousA consistent and infallible su55ort- "an9 thanks to $aube and to $enaA 1athaA Nici and %ina?uwA thank 9ouli for being the best of 5ossible brothers and to m9 uncle and aunt Gerhard and #lse ?engstA m9 grandmother $isa

8<

F %ppendices

/ ABB<(306<A!!endi7 1+ -tud8 s!ecies


=able A1.1+ $ist of )E@ Euro5ean 5asserine bird s5ecies used for anal9ses in cha5ter + Famil9
0egithalidae 0laudidae 0laudidae 0laudidae 0laudidae 0laudidae 0laudidae 0laudidae 0laudidae 0laudidae Bomb9cillidae 4erthiidae 4erthiidae 4erthiidae 4inclidae 4isticolidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae 4orvidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae

Genus
%egithalos %lauda ,alandrella ,alandrella ,hersophilus ;remophila .alerida .alerida -ullula (elanocorypha *ombycilla ,erthia ,erthia #roglodytes ,inclus ,isticola ,orvus ,orvus ,orvus ,orvus ,yanopica .arrulus 2ucifraga <riolus !erisoreus !ica !yrrhocora) !yrrhocora) ,alcarius ,arduelis ,arduelis ,arduelis ,arduelis ,arduelis ,arduelis ,arduelis ,arpodacus

%5ecies
caudatus arvensis brachydactyla rufescens duponti alpestris cristata theklae arborea calandra garrulus brachydactyla familiaris troglodytes cinclus 6uncidis cora) corone frugilegus monedula cyanus glandarius caryocatactes oriolus infaustus pica graculus pyrrhocora) lapponicus cannabina carduelis chloris flammea flavirostris hornemanni spinus erythrinus

8:

F %ppendices Famil9
Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae Fringillidae ?irundinidae ?irundinidae ?irundinidae ?irundinidae ?irundinidae $aniidae $aniidae $aniidae $aniidae $aniidae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae

Genus
,occothraustes ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a ;mberi$a Cringilla Cringilla -o)ia -o)ia -o)ia -o)ia (iliaria !inicola !lectrophena) !yrrhula Serinus Serinus &elichon 'irundo 'irundo 'irundo 8iparia -anius -anius -anius -anius -anius ,ercotrichas ;rithacus Cicedula Cicedula Cicedula -uscinia -uscinia -uscinia (onticola (onticola (uscicapa <enanthe <enanthe

%5ecies
coccothraustes aureola caesia cia cirlus citrinella hortulana melanocephala pusilla rustica schoeniclus coelebs montifringilla curvirostra leucoptera pytyopsittacus scotica calandra enucleator nivalis pyrrhula citrinella serinus urbicum daurica rupestris rustica riparia collurio e)cubitor minor nubicus senator galactotes rubecula albicollis hypoleuca parva luscinia megarhynchos svecica sa)atilis solitarius striata hispanica isabellina

88

F %ppendices Famil9
"uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae "uscica5idae /aridae /aridae /aridae /aridae /aridae /aridae /aridae /aridae /aridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae /asseridae >egulidae >egulidae %ittidae %ittidae %ittidae %ittidae %turnidae

Genus
<enanthe <enanthe <enanthe !hoenicurus !hoenicurus Sa)icola Sa)icola #arsiger #urdus #urdus #urdus #urdus #urdus #urdus ,yanistes -ophophanes !arus !eriparus !oecile !oecile !oecile !oecile 8emi$ %nthus %nthus %nthus %nthus %nthus %nthus (ontifringilla (otacilla (otacilla (otacilla !asser !asser !asser !etronia !runella !runella 8egulus 8egulus Sitta Sitta Sitta #ichodroma Sturnus

%5ecies
leucura oenanthe pleschanka ochruros phoenicurus rubetra rubicola cyanurus iliacus merula philomelos pilaris tor5uatus viscivorus caeruleus cristatus ma6or ater cinctus lugubris montanus palustris pendulinus campestris cervinus petrosus pratensis spinoletta trivialis nivalis alba cinerea flava domesticus hispaniolensis montanus petronia collaris modularis ignicapilla regulus europaea neumayer whiteheadi muraria roseus

)((

F %ppendices Famil9
%turnidae %turnidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae %9lviidae

Genus
Sturnus Sturnus %crocephalus %crocephalus %crocephalus %crocephalus %crocephalus %crocephalus %crocephalus ,ettia 'ippolais 'ippolais 'ippolais 'ippolais -ocustella -ocustella -ocustella !anurus !hylloscopus !hylloscopus !hylloscopus !hylloscopus !hylloscopus !hylloscopus Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia Sylvia

%5ecies
unicolor vulgaris arundinaceus dumetorum melanopogon paludicola palustris schoenobaenus scirpaceus cetti icterina olivetorum pallida polyglotta fluviatilis luscinioides naevia biarmicus bonelli borealis collybita sibilatri) trochiloides trochilus atricapilla borin cantillans communis conspicillata curruca hortensis melanocephala nisoria rueppelli sarda undata

)()

F %ppendices

A!!endi7 2+ -31 threshold sensitivit8 anal8sis


=able A2.1+ Effects of com5etition in the unoccu5ied 5arts of the 5otential rangeA dis5ersal abilit9A taNon ageA habitat shift since $G" and the interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- %5ecies distribution model out5ut was converted from continuous 5robabilities into binar9 out5ut using the mean !robabilit8 value as threshold- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,Asignificances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ model algorithm B>& 5ast climate model 44%" com5etition dis5ersal abilit9 (-E) VV *(-)+, (-E( VV *(-)B, (-B@ V *(-)+, (-BE V *(-)+, (-E( VV *(-)', (-@8 VV *(-)+, (-BB V *(-)+, (-BE V *(-)+, log *taNon age, (-++ *(-)', (-++ *(-)+, (-'B *(-)+, (-'+ *(-)+, (-+' *(-)', (-+' *(-)', (-'@ *(-)+, (-'B *(-)+, habitat shift since $G" -(-': *(-)+, -(-): *(-)+, -(-)E *(-)+, -(-(8 *(-)), -(-') *(-)B, -(-)) *(-)B, -(-)) *(-)+, -(-(B *(-)), dis5ersal abilit9 S com5etition -(-@' V *(-)@, -(-@< VV *(-)@, -(-E+ V *(-)E, -(-E+ VV *(-)<, -(-@: VV *(-)B, -(-E+ VV *(-)@, -(-E< VV *(-)@, -(-E< VV *(-)@, (-:( VVV (-:) VVV (-:( VVV (-:' VVV (-:) VVV (-:) VVV (-:( VVV model >'

-(-+@ *(-)E,

(-:+ VVV

)([

B>&

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-(-BB *(-)E,

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@([

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V! W (-(@HVV! W (-()H VVV! W (-(()

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F %ppendices =able A2.2+ Effects of the com5etition in unoccu5ied 5arts of the 5otential range with the least suitable habitatA dis5ersal abilit9A taNon ageA habitat shift since $G" and interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- %5ecies distribution model out5ut was converted from continuous 5robabilities into binar9 out5ut using the observed !revalence value as threshold- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,Asignificances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ )([ )([ )([ @([ @([ @([ @([ model algorithm B>& B>& >idge >idge B>& B>& >idge >idge 5ast climate model 44%" "#>=4 44%" "#>=4 44%" "#>=4 44%" "#>=4 com5etition dis5ersal abilit9 (-E) VV *(-)+, (-E( VV *(-)B, (-B@ V *(-)+, (-BE V *(-)+, (-E( VV *(-)', (-@8 VV *(-)+, (-BB V *(-)+, (-BE V *(-)+, log *taNon age, (-++ *(-)', (-++ *(-)+, (-'B *(-)+, (-'+ *(-)+, (-+' *(-)', (-+' *(-)', (-'@ *(-)+, (-'B *(-)+, habitat shift since $G" -(-': *(-)+, -(-): *(-)+, -(-)E *(-)+, -(-(8 *(-)), -(-'' *(-)B, -(-)) *(-)+, -(-)) *(-)+, -(-(B *(-)), dis5ersal abilit9 S com5etition -(-@+ VV *(-)@, -(-@< VV *(-)@, -(-E+ VV *(-)B, -(-E+ VV *(-)@, -(-@: V *(-)E, -(-E+ VV *(-)<, -(-E: VV *(-)@, -(-E: VV *(-)@, model >'

-(-+@ *(-)E, -(-BB *(-)E, -(-BB V *(-)@, -(-@) VV *(-)B, -(-BE *(-):, -(-@E V *(-):, -(-@( V *(-)@, -(-@@ VV *(-)B,

(-:+ VVV (-:( VVV (-:) VVV (-:)VVV (-:' VVV (-:( VVV (-:) VVV (-:( VVV

V! W (-(@HVV! W (-()H VVV! W (-(()

)(+

F %ppendices =able A2."+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the unoccu5ied 5arts of the 5otential range and for different range ma5 conversion thresholdsA model algorithms and 5ast climate models%5ecies distribution model out5ut was converted from continuous 5robabilities into binar9 out5ut using the mean !robabilit8 value as threshold- d U dis5eral abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold )([ model algorithm B>& 5ast climate model "#>=4 d N N )([ B>& 44%" N N N @([ @([ B>& B>& "#>=4 44%" N N N N )([ >idge "#>=4 N N N )([ >idge 44%" N N N @([ >idge "#>=4 N N @([ >idge 44%" N N N N N N N a N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N c N N N N N N N N N N N N h d.a d.c N N N d.h a.c a.h c.h 0#4c -:-+8 -E-E' -)(-@E -8-<: -:-@: -8-: -8-: -8-<' -:-<) -)<-E) -)<-+E -)<-) -)<-B) -)<-+E -)<-) -)<-BE -)<-)@ -)<-BE -)<-)@

)(B

F %ppendices =able A2.$+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the whole 5otential range and for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- %5ecies distribution model out5ut was converted from continuous 5robabilities into binar9 out5ut using the observed !revalence value as threshold - d U dis5ersal abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold )([ model algorithm B>& 5ast climate model "#>=4 d N N )([ B>& 44%" N N @([ @([ B>& B>& "#>=4 44%" N N N N )([ >idge "#>=4 N N N )([ >idge 44%" N N N @([ >idge "#>=4 N N @([ >idge 44%" N N N N N N N a N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N c N N N N N N N N N N h d.a d.c N N N d.h a.c a.h c.h 0#4c -:-B8 -E-EE -)(-E' -8-:@ -8-:< -8-:< -8-:@ -:-: -)<-E) -)<-+E -)<-) -)<-B) -)<-+E -)<-) -)<-BE -)<-)@ -)<-BE -)<-)@

)(@

F %ppendices

A!!endi7 "+ Akaike model selection : %ull model regression results %or all habitat suitabilit8 levels
=able A".1+ Effects of com5etition in the !otential rangeA dis5ersal abilit9A taNon ageA habitat shift since $G" and the interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,A significances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ model algorithm 5ast climate model 44%" com5etition dis5ersal abilit9 log *taNon age, (-E( V *(-)8, (-E) V *(-)<, (-@B V *(-)@, (-B8 V *(-)<, (-@+ *(-'), (-@< V *(-'(, (-@B V *(-)E, (-B: *(-):, habitat shift since $G" (-)< *(-'', (-+) *(-)8, -(-BE V *(-)@, -(-': *(-)@, (-(@ *(-'@, (-'@ *(-'', -(-B8 *(-)<, -(-'E *(-)<, dis5ersal abilit9 S com5etition -(-<+ *(-+), -(-<< V *(-'<, -(-<@ V *(-'', -(-E+ *(-'@, -(-@E *(-+B, -(-E< *(-+), -(-E: V *(-'B, -(-@@ *(-'E, (-E+ VV (-<) VVV (-@+ V (-@( V (-E: VV (-<@ VVV (-E+ VV model >'

B>&

-(-B( *(-++,

(-B< *(-'+, (-@B *(-'', (-E: VV *(-)<, (-<( V *(-'(, (-B@ *(-'E, (-@B *(-'@, (-E@ V *(-):, (-EE V *(-'),

(-@: VV

)([

B>&

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-(-@) *(-'8,

)([

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44%"

-(-+( *(-'),

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@([

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-(-)8 *(-'+,

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F %ppendices =able A".2+ Effects of com5etition in the '9F o% the !otential range Eith the least suitable habitatA dis5ersal abilit9A taNon ageA habitat shift since $G" and the interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,A significances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ model algorithm 5ast climate model 44%" com5etition dis5ersal abilit9 log *taNon age, (-E+ V *(-):, (-EB VV *(-)<, (-B' V *(-)), (-B( V *(-)', (-@B *(-'(, (-@: V *(-)8, (-B' V *(-)', (-+: *(-)+, habitat shift since $G" (-'( *(-'), (-+B *(-):, -(-++ V *(-)), -(-'B *(-)), (-(E *(-'+, (-'E *(-'), -(-+: *(-)B, -(-') *(-)+, dis5ersal abilit9 S com5etition -(-<: *(-+(, -(-:' V *(-'E, -(-8' VVV *(-)@, -(-8) VVV *(-)E, -(-E( *(-+), -(-E8 *(-':, -(-8B VVV *(-)<, -(-8( VV *(-)8, (-:) VVV (-:@ VVV (-@E V (-@E V (-:@ VVV (-:< VVV (-EE VV model >'

B>&

-(-BB *(-+),

(-B< *(-'', (-@@ *(-'), (-@' VV *(-)(, (-@+ VV *(-)), (-B@ *(-'@, (-@B *(-'B, (-@+ VV *(-)), (-@+ VV *(-)+,

(-E) VV

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F %ppendices =able A"."+ Effects of com5etition in the 2'F o% the !otential range Eith the least suitable habitatA dis5ersal abilit9A taNon ageA habitat shift since $G" and the interaction between dis5ersal abilit9 and com5etition on range filling- "ulti5le regressions for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- B>& U boosted regression treesH ridge U ridge regression- %hown are standardised 5artial regression coefficientsA standard errors *in 5arentheses,Asignificances adIusted for simultaneous inference and whole model >' and significances- >es5onse asin*sLrt*N,, transformed- n U '+range ma5 conversion threshold )([ model algorithm B>& 5ast climate model 44%" com5etition dis5ersal abilit9 (-B: *(-)8, (-@+ V *(-):, (-B@ VV *(-)), (-BB VV *(-)), (-B@ *(-'), (-@( *(-'(, (-B@ VV *(-)(, (-BB VV *(-)), log *taNon age, (-@8 V *(-)<, (-@: VV *(-)@, (-+: V *(-)), (-+< V *(-)', (-@) *(-):, (-@' V *(-)<, (-+< V *(-)), (-+@ V *(-)', habitat shift since $G" (-'( *(-):, (-+) *(-)E, -(-'@ V *(-)), -(-'( *(-)(, (-(8 *(-'(, (-'@ *(-)8, -(-+( *(-)', -(-)8 *(-)', dis5ersal abilit9 S com5etition -(-:@ V *(-'@, -(-:E VV *(-'', -(-:B VVV *(-)B, -(-:B VVV *(-)@, -(-<' *(-'<, -(-<E V *(-'@, -(-8) VVV *(-)@, -(-8( VVV *(-)E, (-:@ VVV (-:< VVV (-E) VV (-@: VV (-:E VVV (-:< VVV (-<) VVV model >'

-(-@+ *(-'@,

(-E< VV

)([

B>&

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44%"

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@([

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@([

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@([

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V! W (-(@HVV! W (-()H VVV! W (-(()

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F %ppendices =able A".$+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the unoccu!ied !arts o% the !otential range and for different range ma5 conversion thresholdsA model algorithms and 5ast climate modelsd U dis5ersal abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold )([ model algorithm B>& 5ast climate model "#>=4 d N N N )([ B>& 44%" N N N @([ B>& "#>=4 N N N @([ B>& 44%" N N )([ >idge "#>=4 N N )([ @([ >idge >idge 44%" "#>=4 N N N N @([ >idge 44%" N N N N N a N N N N N N N N N N N N c N N N N N N N N N N N N N N N N N N N N N N N N N h d.a d.c N N N N N N N N N N N N N N N N N N N N N N N N d.h a.c a.h c.h 0#4c -))-@8 -)(-:+ -)(-'+ -))-@8 -)(-'+ -)(-)) -:-(' -<-+) -E-(< -:-(' -<-+) -):-+' -)E-8+ -'(-)E -)@-:+ -)@-E: -)@-)E -):-@:

)(8

F %ppendices =able A".'+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the Ehole !otential range and for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- d U dis5ersal abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold )([ model algorithm B>& 5ast climate model "#>=4 d a c h d.a d.c d.h a.c a.h c.h 0#4c

N N N

N N N N N N N N N N N N N N N N N N N N N N N N N N N N

-E-) -@-8B -@-<) -@-): -@-)+ -@-(: -@-(< -B-@B -B-'< -B-)E -<-@' -E-) N N N N N N N N -@-8: -@-8B -@-<) -B-:8 N N -B-E8 -B-B@ N N N N N N N N N N N N N N N N N N N N N N N N N N -B-)B -+-88 -+-+) -+-(: -<-)+ -<-:: -)(-B: -@-'< -:-<8

N N

N N N

N N )([ B>& 44%" N N N N N @([ B>& "#>=4 N

N N N N N N N N N N

N N N

N N N N

@([ )([ )([ @([ @([

B>& >idge >idge >idge >idge

44%" "#>=4 44%" "#>=4 44%"

N N N N N

N N N N N

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F %ppendices =able A"..+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the '9F o% the !otential range Eith least suitable habitat and for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- d U dis5ersal abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold
)([

model algorithm
B>&

5ast climate model


"#>=4

d
N N N N N

a
N N N N N N N N N N N N N N N N

c
N N N N

h d.a

d.c
N

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a.c
N

a.h

c.h

0#4c
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N N N N N N N

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N N N N N N N N

N N

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N N N N N N N N N N N N N N N N N N N N N N N N N

N N N N N N N N N N N N N N N N N N N

-B-)B -+-8' -:-B: -'(-)E -):-+B -'+-8 -)@-B' -)B-E8 -)B-B -)+-B: -)8-:8

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F %ppendices =able A".4+ Best models *corrected 0#4 difference of less than ' from best model, for multi5le regressions focusing on com5etition in the 2'F o% the !otential range Eith least suitable habitat and for different range ma5 conversion thresholdsA model algorithms and 5ast climate models- d U dis5ersal abilit9H a U log *taNon age,H c U com5etitionH h U habitat shift since $G"H B>& U boosted regression treesH ridge U ridge regressionH N U variable or interaction term included in modelrange ma5 conversion threshold )([ model algorithm B>& 5ast climate model "#>=4 N N N N )([ B>& 44%" N N N @([ B>& "#>=4 N N N N @([ )([ B>& >idge 44%" "#>=4 N N N )([ >idge 44%" N N @([ >idge "#>=4 N N @([ >idge 44%" N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N N -)(-)E -8-E@ -:-8' -:-)8 -8-E@ -:-8' -:-)8 -E-B: -E-(@ -B-8E -B-:8 -:-@E -')-++ -'(-8' -''-@' -'(-8' -'(-+ -)8-)B -''-E) d a c h d.a d.c d.h a.c a.h c.h 0#4c

))'

F %ppendices

A!!endi7 $+ *ange %illing estimates %or di%%erent Sylvia s!ecies and distribution modelling methods
=able A$.1+ >ange filling estimates for different Sylvia s5ecies distribution modelling methods- B>& U boosted regression treesH >idge U ridge regression)([ range ma5 conversion threshold s5ecies S. GcurrucaH curruca S. communis S. borin S. atricapilla S. GsardaH sarda S. nisoria S. subcaeruleum S. GnanaH nana S. layardi S. melanocephala S. GhortensisH hortensis S. leucomelaena S. mystacea S. undata S. cantillans S. conspicillata S. GhortensisH crassirostris S. buryi S. abyssinica S. rueppelli S. boehmi S. lugens S. deserticola B>& (-8) (-:8 (-:E (-:@ (-<: (-<B (-E8 (-E@ (-E+ (-@: (-B@ (-B@ (-B@ (-BB (-B) (-B) (-+8 (-+: (-+E (-++ (-+) (-+) (-'' >idge (-<B (-E< (-E' (-@: (-(@ (-BB (-'@ (-+@ (-(E (-)< (-)( (-)E (-)@ (-)+ (-)) (-)) (-)+ (-(< (-)( (-(B (-(E (-(E (-(@ @([ range ma5 conversion threshold B>& (-8) (-:: (-:@ (-:+ (-<< (-<' (-E8 (-EE (-E@ (-@+ (-B' (-@) (-B: (-B( (-+< (-+@ (-B( (-B) (-+) (-+: (-+) (-+' (-'( >idge (-<' (-E: (-E' (-@< (-(E (-B' (-'E (-+@ (-(E (-): (-(< (-)E (-)+ (-)+ (-)) (-)) (-)) (-(< (-(E (-(B (-(E (-(@ (-(@

))+

F %ppendices

A!!endi7 '+ ,ridding threshold sensitivit8 anal8sis


>esults of anal9ses in cha5ter @ using a @([ gridding threshold *i-e- the 5ercentage of minimum overla5 between the range 5ol9gon and a geogra5hic grid cell for that grid cell to be classified as 5resence, for the Sylvia ranges-

#rade4offs between niche breadth and migration distance


=able '.1+ >elationshi5 between migration distance and different niche characteristics- Given are 7A tA !A 82 from sim5le regression anal9ses- %ignificant relationshi5s are 5rinted in boldNo 5h9logenetic signal in an9 regression residuals *all 0bouheif tests. ! \ (-(EA all $ikelihood ratio tests for lambdaU(. ! \ (-@+,- n U 'E(iche characteristic climate niche overla5 between breeding and nonbreeding season total annual climate niche breadth landGcover niche overla! betEeen breeding and nonG breeding season total annual landGcover niche breadth (-(((() -(-((((((E G9.9999/ 9.9992 t -(-:8 (-() P (-+: (-88 R2 (-(+ W (-() 9.$$ 9.2$

G$.". H 9.991 2.4" 9.91

2iche tracking
=able '.2+ 4limate niche overla5 for different 5otential migration strategies in migrant Sylvia warblers- sta9 breed. h95othetical climate niche overla5 resulting from sta9ing on the breeding grounds all 9earA sta9 non-breed. h95othetical climate niche overla5 resulting from sta9ing on the non-breeding grounds all 9earA Given are tA dfA pA from 5aired t-tests6limate niche overla! betEeen breeding and nonGbreeding season vs. sta9 breed. sta9 non-breed. t (-<( -(-(: df )' )' P (-@( (-8B

4limate niche overla5 between the conditions migrant Sylvia warblers eN5erience on the nonbreeding grounds and conditions available on their breeding ranges during the non-breeding season was low *& U (-)B _ (-'@ *mean _ %D,A n U )+,))B

F %ppendices

.eographic pro)imity
=able '."+ "igration distance and distance to the closest suitable non-breeding and breeding areas in migrant Sylvia warblers- 0rea suitabilit9 incor5orates both climate and land-coveractual migration. average distance between known breeding and non-breeding rangesA closest non-breed. average distance between known breeding range and closest suitable non-breeding areaA closest breed. average distance between known non-breeding range and closest suitable breeding areaA Given are tA dfA !A from 5aired t-tests and 5otential savings in distance for the shortest 5ossible route as a 5ercentage of actual migration distanceActual migration distance vs. closest non-breed. closest breed. t B-+E E-() df )' )' P W (-(() W (-(() potential savings (mean SD) ') '( [ '@ ): [

Seasonal changes in niche re5uirements3


Breeding niche breadth in Sylvia warblers was significantl9 smaller than total annual niche breadth *5aired WilcoNon signed rank testA W U )@A n U 'EA ! W (-((),-

))@

,urriculum 0itae

6;**06;C;1 V0=A<
#rina $aube
Biodiversit!t und 1lima Forschungszentrum *Bi1-F, %enckenberganlage '@ E(+'@ Frankfurt am "ain German9 Email. irina-laubeksenckenberg-de /hone. ((B8 E8 <@B' ):') (8T)( ; 5resent /hD at the #nstitute for Ecolog9A Evolution and Diversit9H Goetheniversit9 FrankfurtA German9 in coo5eration with the Biodiversit9 and 4limate >esearch 4entre *Bi1-F,A FrankfurtA German9H su5ervisor. /rofDr- 1atrin B2hning-Gaese (<T(< ; (:T)( /ostgraduate studies at the De5artment of Ecolog9A Johannes Gutenbergniversit9 "ainzA German9 )'T(E ; (ET(< >esearcher at the De5artment of Ecolog9A Johannes Gutenberg- niversit9 "ainzA German9H 5roIect 6/rotected 0reas in German9 under Global 4hange ; >isks and /olic9 =5tions7 for the Federal 0genc9 of Nature 4onservation ))T(E Di5loma in Biolog9H De5artment of Ecolog9A Johannes Gutenbergniversit9A "ainzA German9H thesis. 6$andsca5e structureA avian diversit9 and seed removal of the wild cherr9 *!runus avium $-,7H su5ervisor. /rofDr- 1atrin B2hning-Gaese )(T(' ; (ET(+ E>0%" % scholarshi5 visiting studentH Facult9 of Biomedical and $ife %ciencesA niversit9 of GlasgowA 1 )(T(( ; )(T(' ndergraduate studies in Biolog9H Johannes Gutenberg- niversit9 "ainzA German9

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