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Ellis, Samuel, Franks, Daniel Wayne orcid.org/0000-0002-4832-7470 and Robinson, Elva Joan Hilda orcid.org/0000-0003-4914-9327 (2017) Ecological consequences of colony structure in dynamic ant nest networks. Ecology and Evolution. 1170–1180. ISSN 2045-7758

https://doi.org/10.1002/ece3.2749

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Eco握ogy and Evo握u試on 2017; 1�12 科|科1www.ecolevol.org

ReceivedrゲゾMayゴグゲ葦科 |科 RevisedrゲズNovemberゴグゲ葦科 |科 AcceptedrゴゲDecemberゴグゲ葦DOIrゲグsゲググゴ｠eceザsゴゼジゾ

O R I G I N A L R E S E A R C H

Ecological consequences of colony structure in dynamic ant

nest networks

Samuel Ellis1 科|科Danie旭 Ws Franks2科|科E旭va Js Hs Robinson2

Thisisanopenaccessar抄c旭eunderthetermsoftheCrea抄veCommonsAtribu抄onLicensepwhichpermitsusepdistribu抄onandreproduc抄oninanymediumpprovidedtheorigina旭workisproper旭ycitedsｻゴグゲゼTheAuthorssEco握ogy and Evo握u試onpub旭ishedbyJohnWi旭eyｹSonsLtds

1CentreforResearchinAnima旭BehaviourpUniversityofExeterpExeterpUK2YorkCentreforComp旭exSystemsAna旭ysisｹDepartmentofBio旭ogypUniversityofYorkpYorkpUK

Correspondence

Samue旭E旭旭ispCentreforResearchinAnima旭BehaviourpUniversityofExeterpExeterpUKsEmai旭rsse旭旭isｺexetersacsuk

AbstractAccesstoresourcesdependsonanindividua旭vsposi抄onwithintheenvironmentsThisispar抄cu旭ar旭y important toanima旭s that investheavi旭y innestconstruc抄onpsuchassocia旭insectssManyantspecieshaveapo旭ydomousnes抄ngstrategyrasing旭eco旭onyinhabits severa旭 spa抄a旭旭y separatednestsp otenexchanging resourcesbetween thenestssDiferentnestsinapo旭ydomousco旭onypoten抄a旭旭yhavediferen抄a旭accesstoresourcespbuttheeco旭ogica旭consequencesofthisareunc旭earsInthisstudypweinves-抄gatehownestsurviva旭andbuddinginpo旭ydomouswoodantｪFormica lugubris) colo-

nies are afected by being part of amu旭抄､nest systemsUsing ie旭d data and nove旭ana旭y抄ca旭approachescombiningsurviva旭mode旭swithdynamicnetworkana旭ysispweshowthatthesurviva旭andbuddingofnestswithinapo旭ydomousco旭onyareafectedbytheirposi抄oninthenestnetworkstructuresSpeciica旭旭ypweindthatthelowofresourcesthroughanestpwhichisbasedonitsposi抄onwithinthewidernestnetworkpdetermines a nestvs 旭ike旭ihood of surviving and of founding newnestssOur resu旭tshigh旭ighthowapparent旭ydisparateen抄抄es inabio旭ogica旭systemcanbe integratedintoafunc抄ona旭eco旭ogica旭unitsWea旭sodemonstratehowposi抄onwithinadynamicnetworkstructurecanhaveimportanteco旭ogica旭consequencess

K E Y W O R D S

dynamicnetworksppo旭ydomypredwoodantspsocia旭grouppsurviva旭ana旭ysis

ゲ科 |科INTRODUCTION

Anindividua旭vsaccesstoresourcesisstrong旭yinluencedbyitsposi抄onintheenvironmentre旭a抄vetothatresourcesThiscanhaveimportantbehaviora旭 consequencesq for examp旭ep op抄ma旭 foraging strategieshaveevo旭vedtomakebestadvantageofavai旭ab旭eresourcespgivenanindividua旭vsposi抄onintheenvironmentｪYdenbergpゴググゼｫsThisispar-抄cu旭ar旭ytrueinspeciessuchassocia旭insectspwhichformneststhatarespa抄a旭旭yixedｪat旭eastintheshorttermｫsTheposi抄onofanestintheenvironmentis旭ike旭ytoafectaccesstoresourcesandu旭抄mate旭ytheitnessoftheindividua旭swithinthenestｪMcG旭ynnpゴグゲゴｫs

Manyantspeciesinhabitmu旭抄p旭espa抄a旭旭yseparatedpbutsocia旭旭yconnectednestspastrategyca旭旭edpo旭ydomyｪDeboutpSchatzpE旭iaspｹMckeyp ゴググゼqRobinsonp ゴグゲジｫsNestswithin a po旭ydomous sys-temotenexchangeresourcesｪesgspBuczkowskipゴグゲゴqE旭旭ispProcterpBuckham､BonnetpｹRobinsonp inpressqHofmannpゴグゲジｫsAnestvsaccess toresourceswi旭旭dependnoton旭yon its 旭oca抄onwithin theforagingenvironmentbuta旭soonitsposi抄onre旭a抄vetoothernestssForexamp旭epinpo旭ydomouswoodantｪFormica lugubrisｫco旭oniesfoodandotherresourcesaretransportedthroughtheco旭onybyworkerstrave旭inga旭ongtrai旭sbetweennestsｪE旭旭ispFrankspｹRobinsonpゴグゲジqE旭旭isｹRobinsonp ゴグゲズp ゴグゲ葦ｫsThe combined nests and trai旭s of a

ゴ科 |科 科架 ELLIS ET AL.

po旭ydomouswoodantco旭onythereforeactasaresourceredistribu-

抄onnetworkrfoodresourcesaretransferreda旭ongthetrai旭sbetweenpairs of nestsp resu旭抄ng in co旭ony､旭eve旭 redistribu抄on of resourcesorganizedata旭oca旭 旭eve旭ｪE旭旭isｹRobinsonpゴグゲ葦qE旭旭iseta旭spゴグゲジｫsWoodantsvmajorsourceoffoodishoneydewpaspa抄a旭旭yandtem-

pora旭旭y stab旭e resource ｪDomischp Rischp ｹ Robinsonp ゴグゲ葦ｫs For aworkerpthereforepaccesstofoodwi旭旭dependnoton旭yontheirnestsv旭oca抄onwithin the stab旭e foraging environment but a旭so on theirnestsvposi抄oninthenestnetworkstructuresWorkersfromthesameco旭onypbutinhabi抄ngdiferentnestspthereforehavediferentaccesstoresourcessHoweverptheeco旭ogica旭consequencesofthisdiferen-

抄a旭accesstoresourcesofnestswithinthenetworkpandtheefectthat thisdiferen抄a旭accesshason thestructureof theco旭onypareunc旭ears

Inapo旭ydomousco旭onyptherearesevera旭possib旭eeco旭ogica旭con-

sequencesofanestvsaccesstoresourcessForexamp旭epanestvssur-viva旭pisespitscon抄nuedinhabita抄onpis旭ike旭ytodependonitsabi旭itytoaccessenoughresourcestosustaintheantswithinthenestsSimi旭ar旭ypworkerswithinanestmaybeinluencedbyaccesstoresourceswhenfoundingnewnestssInpo旭ydomouswoodantco旭oniespnewnestsareoten estab旭ished by buddingrDuring buddingpworkers and queens旭eave a nest on foot to found a newnest ｪBourkeｹ Franksp ゲゾゾズqE旭旭isｹRobinsonpゴグゲズｫsItwou旭dbeexpectedthatthedecisionofantswithinanesttobudanewnestisinluencedpposi抄ve旭yornega抄ve旭ypbytheirnestｪthefoundernestvsｫaccesstoresourcesｪHo旭wayｹCasepゴグググq LananpDornhauspｹBronsteinp ゴグゲゲq SorvariｹHakkarainenpゴググズｫs It is important tonote thatbothof these traitsr surviva旭andbuddingpare inherent旭y抄medependentandneedtobestudiedinadynamicframeworks

Theeco旭ogica旭interdependenceofnestswi旭旭deinethenatureofthepo旭ydomoussystemsInamonodomousco旭onyｪaco旭onyinhab-

i抄ngasing旭enestｫp thesurviva旭andbuddingofanestareafectedon旭ybyproper抄esinherenttothatnestpsuchasitssizeand旭oca抄onin the environmentsNestswithin a po旭ydomous systemmay simi-旭ar旭ysurviveandbudbasedon旭yontheirinherentproper抄espwithnoeco旭ogica旭consequencesofthenestnetworkstructuresSurviva旭andbuddingbasedon旭yoninherentproper抄esofanestwou旭dsuggestthatthereisa旭ow旭eve旭ofintegra抄onbetweennestsinthesystemandthatapo旭ydomousco旭ony issimp旭yac旭usterofmutua旭旭ynon-

aggressivenestsandnotpartofasing旭ecoopera抄veandfunc抄ona旭units In contrastp if the nests of a po旭ydomous system are part ofthesamefunc抄ona旭unitpthesurviva旭andbuddingofeachnestwi旭旭beafectednoton旭ybyinherentnestproper抄espbuta旭sobyeitheritsposi抄onintheco旭onynestnetworkormoregenera旭co旭ony､旭eve旭efectss

Inthisstudypweinves抄gatehowthesurviva旭andbuddingofnestsinpo旭ydomousco旭oniesareafectedbythree 旭eve旭soforganiza抄onrｪiｫatributesoftheindividua旭nestp ｪiiｫposi抄onoftheindividua旭nestwithinthenetworkpandｪiiiｫproper抄escommontothewho旭enetworksTheeco旭ogica旭consequencesofdiferen抄a旭accesstoresourceswithinapo旭ydomousco旭onywi旭旭giveimportantinsightsintohowpo旭ydomousco旭oniesarestructuredandpmoregenera旭旭ypthepoten抄a旭importanceofanindividua旭vsposi抄onwithinadynamicnetworks

ゴ科 |科MATERIALS AND METHODS

ゴsゲ科|科Study species and study site

Weinves抄gatedthedynamicsofthenestnetworksofthepo旭ydomousredwoodantFormica lugubrisp amemberof theeco旭ogica旭旭y impor-tantF. rufaspeciesgroupｪStockanｹRobinsonpゴグゲ葦qStockaneta旭sp2016).Wood ants are the dominant invertebrate predator in theirenvironmentq they hunt and scavenge for a variety of invertebratepreypinc旭udingotherantspeciesｪDomischeta旭spゴグゲ葦qMabe旭ispゲゾ芦ジqSavo旭ainenｹVeps士旭士inenpゲゾ芦芦ｫsHoweverpthemajorityoffoodforredwoodantco旭oniesisprovidedbyforagingforhoneydewco旭旭ectedfrom sap､feeding hemipterans in the canopy ｪDomisch eta旭sp ゴグゲ葦qRosengrenｹSundstr塾mpゲゾゾゲｫsThesehemipteranco旭oniesprovideaspa抄a旭旭yandtempora旭旭ystab旭efoodsourcefortheantssAna旭ysisofthestructureofwoodantnestnetworkshashigh旭ightedtheimpor-tanceofhoneydewtransportinstructuringtheco旭onytrai旭structureｪE旭旭iseta旭spゴグゲジｫsDetai旭edobserva抄onofthetrai旭sbetweenwoodantnestshasa旭sosuggestedthathoneydewisthemainresourcebeingtransporteda旭ongtheinternesttrai旭sｪE旭旭isｹRobinsonpゴグゲズpゴグゲ葦ｫsThisstudywasconductedattheLongshawEstateincentra旭Eng旭andsFormica lugubrisistheon旭yF. rufagroupｪtheredwoodantsｫspeciesatthissitesTheF. rufagrouparethedominantantsinEuropeanwood-

旭ands ｪJohansson ｹ Gibbp ゴグゲ葦q Savo旭ainen ｹ Veps士旭士inenp ゲゾ芦芦ｫsTheiron旭ysigniicantcompe抄torsarethereforeotherwoodantspe-

ciesｪJohanssonｹGibbpゴグゲ葦ｫsTheabsenceofotherwoodantspeciesatthissitemeanstheF. lugubrishavenosigniicantinterspeciiccom-

pe抄torssThesiteisamixtureofwood旭andpastureandhistoricp旭anta-抄onssWoodantsconstructnestmoundsfrompineneed旭esandothermateria旭 avai旭ab旭e in the 旭eaf 旭iters In Eng旭andp Formica lugubris are

po旭ygynousandeachnestoftheco旭onyis 旭ike旭ytocontainmu旭抄p旭equeensｪE旭旭isｹRobinsonpゴグゲジqProctereta旭spゴグゲ葦ｫsScotspineｪPinus

sylvestrisｫpoakｪQuercus sp.ｫp旭archｪLarix sp.ｫsycamoreｪAcer pseudopla-

tanusｫpandsi旭verbirchｪBetula pendulaｫarethemostcommontreespe-

ciesatthesitesWoodantco旭oniesatthesiteshownopreferenceforpar抄cu旭artreespeciespneitherdotheyshowanytempora旭varia抄onintreespeciespreferenceｪSamue旭E旭旭isunpublishedｫsThe旭ackofprefer-enceforpar抄cu旭artreespeciessuggeststhattheresourcesavai旭ab旭efrom thediferent tree species at the site are approximate旭y equa旭sTheir旭argesizepstab旭efoodsourcespand旭ackofsigniicantpredatorsandcompe抄torsmeanthatwoodantnestsareotenpresent inthesame旭oca抄onfora旭ongperiodof抄meｪRischpE旭旭ispｹWiswe旭旭pゴグゲ葦qRobinsonｹRobinsonpゴググ芦ｫs

ゴsゴ科|科Network mapping

Werepresentedthepo旭ydomousco旭oniesasnetworkspwiththenestsandtreesasnodesandtheinternestandforagingtrai旭sasedgesｪesgspLatyeta旭spゴグゲゲqCookpFrankspｹRobinsonpゴグゲジqE旭旭iseta旭spゴグゲジqFigureゲｫsWoodants formc旭earabove､ground trai旭sbetweennestsｪinternest trai旭sｫ and between nests and trees ｪforaging trai旭sｫs Thetrai旭sconsistofworkerstrave旭inga旭ongixedpathsotentranspor抄ngresourcesp predominant旭y honeydewp invertebrate preyp and brood

科架 科 | 科ザELLIS ET AL.

F IGURE  ゲ科Timeseriesofco旭onyIIbusedinthisstudytoi旭旭ustratesomeoftheeco旭ogica旭consequencesofnestnetworkpositioninpo旭ydomousFormica lugubris

co旭oniessAsma旭旭networkwaschosenforthepurposeofsimp旭icitysｪaｫCo旭onyIIbatthefivetimepointsusedinthisstudysNestsarerepresentascirc旭esandcategorizedashavinga旭owresourcef旭owｪnorma旭izedbetweenessof旭essthanグsゴズrpa旭eorangeｫamedium旭eve旭ofresourcef旭owｪnorma旭izedbetweenessofmorethanグsゴズand旭essthanグsゼズrorangeｫpandahigh旭eve旭ofresourcef旭owｪnorma旭izedbetweenessofgreaterthanグsゼズrdeeporange｠brownｫsNestswithab旭ackcrossarethosewhichwi旭旭notsurviveunti旭thenexttimepointｪnodataforaftersummerゴグゲジpsononestsaremarkedasbeingabandonedｫsGreentriang旭esrepresenttreessThe旭inesbetweenpointsrepresentforagingｪgreenｫandinternesttrai旭sｪgrayｫsNestswitha旭owormediumf旭owwereabandonedmoreoftenthanthosewithahighf旭owsｪbｫCo旭onyIIbatジtimepointsinthestudysAsabovepcirc旭esrepresentnestssCirc旭eco旭orrepresentthechangeinf旭owthroughthenestsincethe旭asttimepointqb旭ueindicatesadecreaseornochangeinf旭owofresourcessincethe旭asttimepointsPinkshowsnestswhichhaveincreasedinresourcef旭owsincethe旭asttimepointsB旭ackcirc旭esarethenew旭yfoundednestssB旭ackout旭inesrepresentnestsfromwhichanewnestｪsｫhasbeenfoundedｪfoundersｫqgrayout旭inesrepresentpossib旭efounderssNestswithanincreasedresourcef旭owweremore旭ike旭ytofoundnewneststhanthosewithastaticordecreasedresourcef旭ow

ジ科 |科 科架 ELLIS ET AL.

ｪE旭旭is ｹ Robinsonp ゴグゲズp ゴグゲ葦ｫsWorkers show very high ide旭ity totrai旭sprare旭yswitchingbetweentrai旭soncetheyhavebeenrecruitedｪE旭旭is ｹ Robinsonp ゴグゲ葦q Gordonp Rosengrenp ｹ Sundstr塾mp ゲゾゾゴｫsWedeineapo旭ydomousco旭onyastwoormorenestsconnectedbyinternesttrai旭sｪE旭旭iseta旭spゴグゲジｫsOurdeini抄onofaco旭onyispthere-

forep basedon func抄ona旭 resource exchangebetweennestsp ratherthanonthebasisofaggressionorre旭atednesss

Weusethesamemappingmethodemp旭oyedbyE旭旭iseta旭sｪゴグゲジｫprevious旭y at this site tomap the same co旭onies over ジ addi抄ona旭抄me points over the next ゴyearss For each co旭onyp at eachmap-

ping抄me pointpwe recorded the spa抄a旭 and topo旭ogica旭 旭ayout ofthenestsptreespandtrai旭ssForthetrai旭spwemeasuredthe旭engthofthe trai旭p compassdirec抄onof the trai旭p and the traicon the trai旭sThe traicon the trai旭wasmeasuredas the 旭engthof trai旭neededto ind ゲグworkersp which can be converted into number of antspercmoftrai旭andthennumberofantsontheen抄re旭engthofthetrai旭sTheadvantageofbasingtrai旭traicondistanceneededtoindantspratherthanarate､basedmeasurepisthatitisnotre旭iantonthespeed atwhich the ants aremovingpwhich is strong旭y afectedbytheambienttemperatureｪRosengrenpゲゾゼゼｫsAnttraicisameasureoftrai旭strengthbasedon旭yonthenumberofantspassinga旭ongthetrai旭q howeverp this is 旭ike旭y tobeafectedby thenumberofwork-ers avai旭ab旭e to trave旭 a旭ong the trai旭ssTrai旭weight is ameasureoftrai旭importanttoapar抄cu旭arnestpre旭a抄vetothepopu旭a抄onsofthenestsbeingconnectedsWeca旭cu旭atedeachtrai旭vsweightbydividingthetota旭numberofantsonthetrai旭bythemeanpopu旭a抄onofthenestsconnectedbythetrai旭ｪE旭旭iseta旭spゴグゲジｫsForforagingtrai旭sptheweightofthetrai旭sarere旭a抄vetothepopu旭a抄onoftheforagingnestｪE旭旭iseta旭spゴグゲジｫs

Foreachmappingofeachco旭onypwea旭soes抄matedthepopu-

旭a抄onsofthenestsｪChenｹRobinsonpゴグゲザｫpmeasuredthecanopy

cover over the nestsp and recorded the species of the trees usedfor foragingsTheworker popu旭a抄on in a nestwas es抄mated fromthevo旭umeofthenestmoundpca旭ibratedatthissitewithamark･re旭ease･recapture measure ｪChen ｹ Robinsonp ゴグゲザq E旭旭is eta旭spゴグゲジｫsCanopy coverwas es抄matedon thebasis of digita旭 photo-

graphstakenfromver抄ca旭旭yaboveeachnestｪE旭旭iseta旭spゴグゲジｫsEachmapwasusedtoconstructaspa抄a旭旭yembeddednetworkoftheco旭-onypwithedgesweightedbytrai旭strengthpandthenodeproper抄esofnestpopu旭a抄onｪhereaterrnestsizeｫpdistancetothenearesttreepandcanopycoversWeexaminedhowthenestnetworksofthirteenpo旭ydomouswoodantco旭onieschangedover抄mesThirteenofthe旭argestco旭oniesatthesitewerestudiedpchosenfromapre旭iminarysurveyinMayゴグゲゴｪTab旭eゲqAppendixSゲｫsTheco旭onieswereirstfu旭旭ymapped in 旭ateAugust ゴグゲゴ ｪana旭yzed as sta抄c networks inE旭旭iseta旭spゴグゲジｫsFor thenextゴyears ｪゴグゲザandゴグゲジｫp theco旭o-

nieswere fu旭旭ymapped ｪusing themethods out旭ined aboveｫ twiceperyearroncein旭atespringandagainin旭atesummersEachremap-

pingwasperformedb旭indpwithoutreferencetothemapsofprevious抄me pointssWood ants show seasona旭 ac抄vity paternsrThey arequiescent over winterp beginning foraging ac抄vity ｪand producingsexua旭ofspringｫin旭atespringpandcon抄nuingforagingthroughoutthesummerandear旭yautumnｪMaedereta旭spゴグゲ葦ｫsRemappingco旭-onies in 旭ate springandagain in 旭ate summer therefore representsthebeginning and theheight of the foraging seasonp respec抄ve旭ysThe抄mingof the 旭ate springmappingwasdependenton the抄m-

ingofspringineachyearandwasnotperformedun抄旭temperatureswerehighenoughthatbothforagingandinternesttrai旭ac抄vitywerebeing performed ｪRosengrenp ゲゾゼゼｫs Late summer mapping wasa旭waysperformed in thesecondha旭fofAugustsWoodantac抄vitycanbedependentontemperatureandweathercondi抄onssoa旭旭co旭-oniesweremappedinwarmpdrycondi抄onss

TABLE  ゲ科Detai旭softheco旭oniesusedinthisstudysNumbersrefertothenestspresentintheco旭onyatthat抄mepointsSpringrefersto旭ateMayｪthebeginningoftheforagingseasonｫandsummerin旭ateAugustｪthepeakoftheforagingseasonｫsNetchangeinnestsdescribesthediferenceinnumberｪandpercentageｫofnestsintheco旭onybetweensummerゴグゲゴandsummerゴグゲジsAveragenestpopu旭a抄onｪnestsizeinthetextｫreferstothemeannumberofantspredictedtobeinthenestsofeachco旭ony

Tota旭 number of nestsNet change in number of nests

Average nest popu旭a抄on (range)2012- Summer 2013- Spring 2013- Summer 2014- Spring 2014- Summer

I 21 16 ゲズ 11 ゲジ ┋ゼｪ┋ザザsザ鯵ｫ ゼ芦pゼ芦グｪ葦ゴズ･ゲpゼゾゲp葦ゲゼｫ

IIa ジ ザ ジ ジ ジ グｪグ鯵ｫ ゴゴpゾジゲｪゴゾグ･ゾザp芦芦ザｫ

IIb 6 6 6 9 9 ┊ザｪ┊ズグ鯵ｫ ゲゼp葦グゲｪゲズ葦グ･葦ゴp葦ジゲｫ

III 12 12 芦 9 16 ┊ジｪ┊ザザsザ鯵ｫ ザズジザズｪゲズ葦･ゲ葦葦p芦ゲズｫ

IV 12 9 6 7 7 ┋ズｪ┋ジゲs葦鯵ｫ ザ葦pズ芦芦ｪ芦ズゲ･ゲ芦ズpズググｫ

V ゲジ 11 10 芦 2 ┋ゲゴｪ┋芦芦鯵ｫ ゴゾpゴゼゴｪズゴゴ･ゴ葦ズpググズｫ

VI ゲジ 12 12 ゲザ 11 ┋ザｪ┋ゴゲ鯵ｫ ゲ芦p芦ゴゼｪゲゲゾ･ゲゲグpグザゾｫ

VII 7 7 ジ ズ 芦 ┊ゲｪ┊ゲジ鯵ｫ ゴゲpゲグゾｪゴ芦ズ･ゲゲジpゴザ芦ｫ

VIII 6 ザ ジ 6 ジ ┋ゴｪ┋ザザsザ鯵ｫ ズジpゴズズｪゴゲグ･ザ芦ジpゲ葦葦ｫ

IX 9 11 17 11 ゲズ ┊葦ｪ┊葦葦s葦鯵ｫ ザゴp葦芦ゲｪ葦芦･ゴ芦芦pザ芦グｫ

X ゲザ 芦 10 9 芦 ┋ズｪ┋ザ芦鯵ｫ ゲ芦pジゴザｪゲ葦グ･ゾグpグ葦ジｫ

XI 20 ゲズ 10 10 17 ┋ザｪ┋ゲズ鯵ｫ ゲグpゲゼズｪ芦ゾ･ゲザグp芦葦グｫ

XII 6 6 ザ 芦 ザ ┋ザｪ┋ズグ鯵ｫ ゼジpズゴ芦ｪゲ芦グズ･ゴジグpザゴゾｫ

科架 科 | 科ズELLIS ET AL.

ゴsザ科|科Ana旭ysis

Weare interested inhowdynamicproper抄esofnestswithinpo旭y-domous co旭onies are inluenced by their inherent atributes ｪnestatributesｫp their posi抄on within the nest network ｪnetwork posi-抄onｫpandatributessharedwiththewho旭eco旭onyｪco旭onyatributesqsummarized inTab旭eゴｫsA旭旭networkana旭ysiswasperformed inR ｪRDeve旭opmentCoreTeamゴグゲゲｫusingthewigraphxpackageｪCsardiｹNepuszpゴググ葦ｫs

Nestatributesarethosebaseddirect旭yoninherentatributesofthenestsThepopu旭a抄onofanestｪhereaternestsizeｫpcanopycoveroverthenestpandthedistancefromthenesttothenearesttreea旭旭havebeenshown tobeeco旭ogica旭旭y important forwoodants ｪChenｹRobinsonpゴグゲジｫandthereforehavethepoten抄a旭toinluencethesurviva旭andreproduc抄onofnestss

Network posi抄on proper抄es depend on a nestvs 旭oca抄on in theco旭ony nest networks Resource exchange between the nests of apo旭ydomouswoodantco旭onyisbasedonworkersfromagivennesttrave旭inga旭onginternesttrai旭stoneighboringnestspco旭旭ec抄nghoney-dewpandthenreturningtotheirorigina旭phomepnestｪE旭旭isｹRobinsonpゴグゲ葦ｫsThismechanismisbasedon旭oca旭resourceexchangepbetweenneighboringnestspwithoutreferencetotheeiciencyofco旭ony､旭eve旭resourceredistribu抄onｪE旭旭iseta旭spゴグゲジｫsAresourceexchangemecha-nismbasedonworkersfromagivennesttrea抄ngothernestsasfoodsourceshasthepoten抄a旭toresu旭tinresourceexchangethroughtheen抄reco旭ony ｪCookpFrankspｹRobinsonpゴグゲザqSchmo旭kepゴググゾｫs Inasystembasedon旭oca旭resourceexchangepthequan抄tyofresourceavai旭ab旭etoagivennestcanberepresentedasthelowthroughthatnestr Inwoodantco旭oniesp resourceslowfromthe treespand theninsomecasesthroughtheinternesttrai旭networkptothenestssF旭owthroughanodeinanetworkcanbemeasuredasbetweenesscentra旭-itysBetweeness is ameasureof the tota旭 numberof shortestpathsbetweenpairsofnodesinthenetworkwhichpassthroughapar抄cu旭arnodeｪesgspCrotpJamespｹKrausepゴググ芦qWhiteheadpゴググ芦ｫsWecor-rectforigraphvsreversetreatmentofweightedva旭uesbyinver抄ngthestrengthoftrai旭sforcentra旭ityana旭ysissOurmappednetworksinc旭udeboththenestsoftheco旭onyandthetreesonwhichtheyareforagingpisesp theana旭yzednetworkscontainboth theco旭onyand its foragingenvironmentsInourpo旭ydomousnetworkspa旭旭treesareattheendofanetworkontheirownbranchandthereforehaveabetweenessofzerosIncontrastpthebetweenessofanestisbasedonthenumberof

shortestpathspassingthroughitpinc旭udingthosefromtreestoothernestsinthenetworksBetweenesscanthereforeactasameasureofｪpoten抄a旭ｫresourcelowthroughapar抄cu旭arnestpdependentonthatnestvspaternoftrai旭sconnectedtoothernestsandtreesinthenet-worksWeusedaweightedmeasureofbetweenesstoaccountforthenumberofantsonatrai旭pgiventhesizeoftheconnectednestsｪtrai旭weightｫsToa旭旭owcomparisonbetweennetworksforeachco旭onypthebetweenesswasnorma旭izedtothe旭argestva旭uewithineachnetworkｪesgspLusseauｹNewmanpゴググジｫs

Co旭ony atributes are those which are shared by a旭旭 the nestswithinanestnetworksAttheco旭ony旭eve旭pweareinterestedinhowtheamountofresourcesco旭旭ectedbytheen抄reco旭onyinluencesthesurviva旭andbuddingofthenestssWeusethenumberofantsonfor-agingtrai旭sasameasureofaco旭onyvsforagingefortsThenumberofantson foraging trai旭scanbeca旭cu旭atedbymu旭抄p旭ying theantspercmforeveryforagingtrai旭bythe旭engthofthatforagingtrai旭andthensummingtheseva旭uesforthewho旭eco旭onysThisforagingmetricisameasureoftheresourceacquisi抄onefortoftheen抄reco旭onypnotacountofthenumberofforagersintheco旭onysThera抄oofthetota旭popu旭a抄onoftheco旭onyｪsummedsizeofa旭旭thenestsintheco旭onyｫto the foragingefortof theco旭onyphereaterworkerrforaging ra抄opgivesanes抄mateof foragingefortperworker in theco旭onysA 旭owworkerrforaging ra抄o suggests a high foraging efort per workerpwhereasahighworkerrforaging ra抄o indicatesa 旭ow foragingefortperworkersWeusetheworkerrforagingra抄oasameasureofco旭ony､旭eve旭 resourceacquisi抄onsWe inves抄gatehowworkerrforagingra抄opredictsvarioussurviva旭ppopu旭a抄onchangepandbuddingofnestsｪseebe旭owｫtoseehowthisco旭ony､旭eve旭measureofresourceacquisi抄oncompares to the network posi抄on､based resource acquisi抄onmea-surernorma旭izedbetweenesss

Internesttrai旭scana旭sohaveinherentpwithin､networkpandco旭onyatributessAn important inherenttraitofan internesttrai旭 istheanttraiconthattrai旭sAnttraica旭ongatrai旭doesnottakeintoaccountthesizeof thenestsconnectedby the trai旭ssTrai旭weight takes intoaccountthesizeofthenestsbeingconnectedbythetrai旭ssTheper-sistence of an internest trai旭 is 旭ike旭y to be afected by the proper-抄esandnetworkposi抄onofthenestswhichtheyjoinsSimi旭ar旭ypthenumberofinternesttrai旭sandforagingtrai旭sassociatedwiththetwoneststhataninternesttrai旭connectscana旭sobeconsideredawithin､networkatributesThebetweenessofatrai旭pun旭ikethebetweenessofanestp isaco旭ony､旭eve旭efectsTrai旭betweenessisaco旭onyatribute

TABLE  ゴ科Thenestatributepnetworkposi抄onpandco旭onyatributevariab旭esusedtoinves抄gatetheeco旭ogica旭consequencesofanestposi抄oninapo旭ydomousco旭ony

Nest atributes Network posi抄on Co旭ony atributes

The popu握a試on of the nestsCa旭cu旭atedbasedonnestvo旭umes

Nest betweenness.Ca旭cu旭atedfromtheweightednetworkmapspinc旭udingbothtreesandnestss

Worker器forager ra試osThetota旭popu旭a抄onoftheco旭onyｪthesumofa旭旭nestpopu旭a抄onsｫdividedbytheinstantaneousnumberofworkersonforagingtrai旭sｪfromthetrai旭strengthsｫs

The canopy coverabovethenestssCo旭旭ectedusingdigita旭photographsofthecanopyabovethenests

The distance to the nearest treesCa旭cu旭atedasthe旭ineardistanceｪisespnota旭ongforagingtrai旭sｫfromanesttothenearesttrees

葦科 |科 科架 ELLIS ET AL.

becauseitrepresentstheimportanceofatrai旭toco旭ony､旭eve旭resourcelowpratherthantheamountofresourcespassingthroughapar抄cu旭arnests

Anunder旭yingassump抄onofourmeasuresofbothnetworkposi-抄onproper抄esandco旭ony､旭eve旭proper抄esisthatthe旭eve旭oftraicon a foraging or internest trai旭 is representa抄ve of the quan抄ty ofresources ｪspeciica旭旭y honeydewｫ being transported a旭ong this trai旭sThisassump抄onisbasedondatashowingthatｪiｫthestrengthofaninternesttrai旭 isposi抄ve旭yre旭atedtothe旭eve旭offoragingbeingper-formedateachendofthetrai旭ｪE旭旭iseta旭spゴグゲジｫpｪiiｫゼグ鯵ofinternestjourneysinvo旭vetransportofhoneydewｪE旭旭isｹRobinsonpゴグゲ葦ｫpandｪiiiｫ the presence of internest trai旭s predicts transport of resourcesbetweennestsｪProctereta旭spゴグゲ葦ｫs

ゴsジ科|科Nest surviva旭

Weareinterestedinwhichfactorsｪnestatributespnetworkposi抄onporco旭onyatributesｫinluencethesurviva旭ofanestinapo旭ydomouswoodantco旭onysWeusesurviva旭ana旭ysispadaptedforusewithnet-workdataptoinves抄gatethefactorsinluencingnestsurviva旭sSurviva旭ana旭ysis is used to describe the抄meun抄旭 an event occursq for ourpurposesptheeventinques抄onisthatanestisabandonedｪK旭einbaumｹK旭einpゴグゲゴｫsNestabandonmentcanbe inferred fromtheco旭onymapsby theabsenceof aprevious旭ypresentnest at thenext抄mepointsTheadvantageofusingsurviva旭ana旭ysisratherthanmorecon-

ven抄ona旭sta抄s抄ca旭techniquesisthatcensoreddatacanbeinc旭udedsCensoreddataoccurwhensomeinforma抄onisknownaboutanindi-vidua旭pforexamp旭ewhenanestisfoundedpbutnototherinforma抄onpforexamp旭ewhenitisabandonedsThisisusefu旭forourdataasmanynestssurvive旭ongerthanourstudyperiodsSurviva旭ana旭ysisa旭旭owsustoinves抄gatehowthesurviva旭ofanestchangeswith抄meinre旭a抄ontonetworkdynamicss

WeusedanextendedCoxpropor抄ona旭hazardｪCoxPHｫmode旭toinves抄gatetheefectofexp旭anatoryvariab旭esｪXnｫonthehazardpoten抄a旭ph(t) ｪEqua抄onゲforthebasicCoxPHmode旭ｫsThehazardpoten抄a旭 is the instantaneous poten抄a旭 per unit 抄me that a nestｪor trai旭ｫ is abandonedp given that the nest ｪor trai旭ｫ has survivedupto抄met ｪK旭einbaumｹK旭einpゴグゲゴｫsThesurviva旭 func抄onpS(t)pdescribes the probabi旭ity that a nest survives 旭onger than a given抄metsTheextendedCoxPHmode旭a旭旭ows抄me､dependentexp旭an-

atory variab旭es to be inc旭uded in the ana旭ysis ｪK旭einbaumｹK旭einp2012).

IntheextendedCoxPHmode旭sreportedinthisstudyptheexp旭an-

atoryvariab旭esｪXpｫwerethenestatributepnetworkposi抄onporco旭-onyatributevariab旭eｪsｫbeing inves抄gatedsCo旭onywasa旭sousedasanexp旭anatoryvariab旭e ina旭旭mode旭ssWhen themode旭wasused todescribethesurviva旭ofatrai旭pthesurviva旭ofthenestsassociatedwiththetrai旭wasusedasanaddi抄ona旭exp旭anatoryvariab旭esThepresence

ofthenestswhichboundthetrai旭ispc旭ear旭ypvita旭tothesurviva旭pres-enceofthetrai旭itse旭fandwasa旭wayshigh旭ysigniicants

Surviva旭mode旭sassumeindependenceofdatapanassump抄onthatisvio旭atedbynetworkdatasThereforepweconstructedanu旭旭mode旭basedonthequadra抄cassignmentprocedureusingゲグpグググnodeatri-butepermuta抄onsｪCrotpMaddenpFrankspｹJamespゴグゲゲｫsWethenmeasuredtheexperimenta旭teststa抄s抄cagainstthisnu旭旭distribu抄ontoderivesta抄s抄ca旭signiicancesPermuta抄onswereconstrainedwithineachmapｪisespwithineachco旭onymapfromapar抄cu旭ar抄mepointｫsA旭旭reportedsta抄s抄csassociatedwithsurviva旭werebasedonthequa-dra抄cassignmentproceduresForsomeana旭ysespthesma旭旭estco旭oniesｪIIaandVIIIｫwerenotinc旭udedbecausethe旭ackofvaria抄onpreventedthemode旭 deining the conidence interva旭sq this is indicated in thetextby旭oweredva旭uesofnsSurviva旭ana旭ysiswasperformedinRusingthewsurviva旭xpackageｪTherneaupゴグゲゴｫs

ゴsズ科|科Nest budding

Newnestswereotenfoundedwithinthepo旭ydomousco旭oniesusedinthisstudysAnestthatwasnotpresentattheprevious抄mepointwas considered to be new旭y foundedsWe used our co旭ony 旭ayoutmaps ｪdescribedaboveｫ to inferwhichnestactedasthefounderofthenew旭yfoundednestsWerefertoanestfromwhichanewnestisbuddedasitsnata旭nestsToinferwhichnestsarethenata旭nestspweassumedthatｪiｫnew旭ybuddednestsstayconnectedtotheirnata旭nestbyatrai旭andｪiiｫthenata旭nestisthenearestnesttowhichthenew旭ybuddednestisatachedsWeusetheseassump抄onstocategorizea旭旭thenestswithinaco旭onyaseitherrnew旭yfoundedpfounders ｪthosefromwhichanewnesthasbeenbuddedｫpornonfoundersｪthosefromwhichanewnesthasnotbeenbuddedｫsInsomecasespthenearestnest toanew旭y foundednestwasanothernew旭y foundednestsAstheorderof founda抄oncannotbe inferredp thenearestestab旭ishedｪisespnotnew旭yfoundedｫnestwascharacterizedasapossib旭efoundersNew旭yfoundednestscanbeeitherforagingornonforagingandarefounded both on and of exis抄ng trai旭s ｪE旭旭isｹRobinsonp ゴグゲズｫs Inaddi抄onpthereisnoseasona旭efectofnestfounda抄onwithinthe抄meperiodsofourstudyrNestsareequa旭旭y旭ike旭ytobefoundedinspringandsummerｪE旭旭isｹRobinsonpゴグゲズｫs

Weusedgenera旭旭inearmixedefectmode旭sｪGLMMsｫtoana旭yzehow budding re旭ates to various nestp nestwithin the networkp andco旭ony atributess In theGLMMsp founder status ｪisesp founderp non-

founderporpossib旭efounderｫwasusedastheresponsevariab旭ewiththevariab旭eｪsｫofinterestastheixedefectsCo旭onypnestIDpandsea-sonwereinc旭udedasrandomefectssFurtherdetai旭softhetestsaregiveninthesuppor抄nginforma抄onqthesuperscriptinthetextrefersto the rowof the tab旭e ｪAppendix SゴｫsA旭旭GLMMsused a binomia旭errorstructureanda旭ogit旭inkfunc抄onsWetestedsigniicanceusingachi､squaredana旭ysisofdevianceｪAoDｫwhichcomparesthefu旭旭mode旭to a nu旭旭 mode旭without the ixed efects If the nu旭旭 mode旭 and fu旭旭mode旭aresigniicant旭ydiferentpitindicatesthattheixedefecthasasigniicantimpactinexp旭ainingthedatasHerewereporttheresu旭tsoftheAoDsGLMMswereperformedinRusingthew旭meジxpackageｪRDeve旭opmentCoreTeamRpゴグゲゲｫs

ｪゲｫh�

t,X�

=h0(t)e

p∑

i=1

βiXi

X= (X1,X2,…Xp)

科架 科 | 科ゼELLIS ET AL.

ザ科 |科RESULTS

ザsゲ科|科Nest surviva旭

The posi抄on of a nestwithin the network is a key predictor of itssurviva旭sNestswithahighernorma旭izedbetweenessaresigniicant旭ymore 旭ike旭y to survive than nestswith a 旭ower norma旭ized betwee-

nessｪCoxPHrz┎┋ザs芦pn┎ズズ芦pp┎sグググゴqFigureゴｫsThisre旭a抄onshipbetween norma旭ized betweeness and surviva旭 is robust evenwhennestsize is introduced intothesurviva旭mode旭sLargernestsaresig-niicant旭ymore旭ike旭ytosurvivethansma旭旭ernestsｪCoxPHpz┎┋ゴsゼpn┎ズズ芦pp┎sググゴ葦ｫpbutwheninc旭udedinthesamemode旭asanaddi-抄ona旭ixedefectpnestswithahighernorma旭izedbetweenessares抄旭旭signiicant旭ymore 旭ike旭y to survive thannestswith a 旭owernorma旭-izedbetweenessｪCoxPHrnorma旭izedbetweenessrz┎┋ゴsグ葦pn┎ズズ芦pp┎sグゲジ葦qnestsizerz┎┋ゴsゴゲpn┎ズズ芦pp┎sグゲゾ芦ｫs

Thesurviva旭ofinternesttrai旭sispsimi旭ar旭yppredictedbytheposi-

抄onwithinthenestnetworkoftheneststheyconnectsInternesttrai旭s

connecttwonestsqeachnesthasava旭ueofnorma旭izedbetweenesss

The旭owerofthesetwonorma旭izedbetweenessva旭uesissigniicant旭y

re旭atedtothesurviva旭oftheinternesttrai旭ｪCoxPHrz┎┋ゲsザグpn┎ジゼ葦p

p┎sグザゼザｫsThere旭a抄onshipisnega抄verAtrai旭associatedwithanest

witha旭ownorma旭izedbetweenessis旭ess旭ike旭ytosurvivethanatrai旭

associatedwithanestwithahighnorma旭izedbetweenesssNestswith

ahighnorma旭izedbetweenessaremore 旭ike旭ytosurvivethannests

witha旭ownorma旭izedbetweenessｪaboveｫwhichmayexp旭aintheneg-

a抄vere旭a抄onshipbetweentrai旭surviva旭andnestnorma旭izedbetwee-

nesssTrai旭s associatedwithanestwith 旭ownorma旭izedbetweeness

maybe旭ess旭ike旭ytosurvivebecausethenestis旭ess旭ike旭ytosurvivep

ratherthanduetotheposi抄onofthetrai旭withinthenetworksWe found no re旭a抄onships between surviva旭 and co旭ony､旭eve旭

efectss Nest surviva旭 is not signiicant旭y re旭ated to co旭onyworkerr-foragingra抄oｪCoxPHrz┎グsゴジpn┎ズズ芦pp┎sザゼザゾｫpevenwhennestsizeisa旭soinc旭udedinthesurviva旭mode旭ｪCoxPHrz┎グsザ芦pn┎ズズ芦pp┎sザジ芦ズｫsSimi旭ar旭yptrai旭surviva旭isnotsigniicant旭yre旭atedtoeithertrai旭 betweeness ｪCox PHr z┎┋ゲs葦芦p n┎ジゼ葦p p┎sグ葦ジ葦ｫ or co旭onyworkerrforagingra抄oｪCoxPHrz┎┋ゲsザゼpn┎ジゼ葦pp┎sゲグゲ葦ｫs

Atributesofthenestsandtrai旭scana旭soinluencetheirsurviva旭sLargernestsaresigniicant旭ymore旭ike旭ytosurvivethansma旭旭ernestsｪCoxPHpz┎┋ゴsゼpn┎ズズ芦pp┎sググゴ葦ｫsHoweverptheothernestatri-buteswemeasureddonotafectsurviva旭sThesurviva旭ofagivennestis not signiicant旭y afected by either the distance from the nest tothenearesttreeｪCoxPHrz┎┋ゲsゴジpn┎ズ芦ゲpp┎sゲグゲゼｫorthecanopycoveroverthenestｪCoxPHrz┎┋グsゲゼpn┎ズ葦ザpp┎sザゼゾ芦ｫsInternesttrai旭swithahighant traicweresigniicant旭ymore 旭ike旭y tosurvivethantrai旭swith旭owanttraicｪCoxPHrz┎┋ゴsジpn┎ジゼ葦pp┎sググジゴｫsHoweverp there is no signiicant re旭a抄onship between trai旭 weightｪwhichisadjustedforthesizesoftheconnectednestsｫandtrai旭sur-viva旭ｪCoxPHrz┎┋グsズゾpn┎ジゼ葦pp┎sゴ葦ゾゾｫs

ザsゴ科|科Nest budding

Nests fromwhichnewnests havebeen founded ｪfoundersｫ have asigniicant旭y higher norma旭ized betweeness than those from whichno new nests have been founded ｪnonfoundersｫ ｪAoD1: χ

2┎ゲゴsジpdf┎ゲpp┑sググゲqFigureザｫsNestsotenchangeinnorma旭izedbetwee-

ness between two 抄me pointss Founder nests have a signiicant旭ygreaterincreaseinnorma旭izedbetweenessovertheperiodwhichthenewnestwasfoundedthannonfoundernestsｪAoD2: χ2┎ゲジsゼpdf┎ゲpp < .0001).

Nestsinco旭onieswitha旭owworkerrforagingra抄oｪisespahighfor-agingefortperworkerｫarenotsigniicant旭ymore旭ike旭ytobefound-

ers than nests in co旭onieswith a highworkerrforaging ra抄o ｪAoDザ: χ

2┎グsゲズp df┎ゲp p┎sゼグｫs Simi旭ar旭yp nests in co旭onies which have a旭oweredworkerrforagingra抄o ｪisespan increase inforagingefortperworkerｫarenotsigniicant旭ymore 旭ike旭y tobefounders thannest inco旭onieswitha raisedworkerrforaging ra抄o ｪAoDジ: χ2┎グsグジpdf┎ゲpp┎s芦グｫs

F IGURE  ゴ科Nestsurviva旭dependsonthef旭owｪmeasuredasnorma旭izedbetweenessｫofresourcessｪaｫThemeanｪ┓standarderrorｫnorma旭izedbetweennessｪpotentia旭resourcef旭owｫofnestswhichsurvivetothenexttimepointpandthenestswhichareabandonedbeforethenexttimepointsｪbｫSurviva旭ofnestspredictedbytheextended､coxPHsurviva旭mode旭sCurvesrepresenthowthesurviva旭ofnestswithadefinedｪandunchangingｫresourcef旭owispredictedtochangewithtimesAhighresourcef旭ownestisanestwithanorma旭izedbetweenessofグsゾpmediumresourcef旭ownestisanestwithanorma旭izedbetweenessofグsズpanda旭owresourcef旭ownestisanestwithanorma旭izedbetweenessofグsゲsCurvesareca旭cu旭atedusingtheKap旭an･MeiermethodsTheerrorｪSEｫisthedifferencebetweensurviva旭indifferentco旭oniessEachpointrepresentsthesurviva旭ofanestatt+xtimepointsafterthenestisfoundedqfouristhemaximumtimepointsafterfoundationasourstudyon旭ycoveredfivetimepoints

(a)

(b)

芦科 |科 科架 ELLIS ET AL.

Nest atributes do not predictwhether the nest has acted as afounders Founder nests are not signiicant旭y 旭arger thannonfoundernests ｪAoDズ: χ

2┎グsゴグp df┎ゲp p┎s葦ズｫs Simi旭ar旭yp nestswhich had agreaterincreaseinsizearenotsigniicant旭ymore旭ike旭ytobefound-

ersthannestswhichhavehada旭owerincreaseoradecreaseinsizeｪAoD.6: χ

2┎グsグザp df┎ゴp p┎s芦ゼｫs Founder nests are neither signii-

cant旭y c旭oser to trees ｪAoD7: χ2┎グsゲゾp df┎ゲp p┎s葦葦ｫp nor have a

signiicant旭y旭owercanopycovernestｪAoD芦: χ2┎グsゼ芦pdf┎ゲpp┎sザ芦ｫthannonfoundernestssNestsarenotsigniicant旭ymore 旭ike旭ytoactasfoundersinspringthaninsummerｪAoD9: χ2┎グsゲ葦pdf┎ゲpp┎s葦ゾｫs

We a旭so inves抄gated how the popu旭a抄on change of nests isafectedbynestatributespnestposi抄onpandco旭onyatributessWefoundnosigniicantefectsｪresu旭tsreportedinAppendixSザｫs

ジ科 |科DISCUSSION

In this studypwe found that the posi抄on of a nestwithin the net-workofpo旭ydomousFormica lugubrisco旭onieshasimportanteco旭ogi-ca旭consequences for thatnestandthestructureand integra抄onoftheco旭onysNestswithahigherlowofresourcespevenifthiscomesindirect旭yviaothernestsphaveanincreasedchanceofsurvivingandfoundingnewneststhannestswitha 旭owerlowofresourcesqnestsizeisa旭soaccountedforanddoesnote旭iminatethisefectsDistanceto thenearest foraging treedoesnotafectnestsurviva旭sResourcelowthroughanestdependsonitsconnec抄onstotheothernestsandhowititsintothebroaderstructureofthenetworksThesurviva旭andbuddingofanestisdependentonitsre旭a抄onshipwithothernestsandthewiderpaternofinterac抄onbetweenthenestsinthepo旭ydomousco旭onysOurresu旭tsshowthatpdespitebeingspa抄a旭旭yseparatedptheinterconnected nests of a po旭ydomous co旭ony can be considered asing旭e eco旭ogica旭 unitp at 旭east in terms of resource acquisi抄onsWea旭sodemonstratethatdynamicnetworkposi抄oncanhaveimportanteco旭ogica旭consequencess

Theviewofthenestsofpo旭ydomouswoodantco旭oniesasforminga sing旭e eco旭ogica旭 unitp supported by our resu旭tsp suggests that thefactorsinluencingtheitnessofindividua旭sinagivennestare旭ike旭ytobestrong旭y 旭inkedtotheitnessof individua旭s inothernestssTheabi旭ityofanesttosurviveandbuddependspinpartponitsposi抄onintheco旭onynestnetworksThisdependenceshows that the resourcemovementthroughtheco旭onyhasanimportanteco旭ogica旭inluencesChanges in theenvironmentnearanygivennesthave thepoten抄a旭toafectthesurviva旭andbuddingofneststhroughoutthenetworksHoweverp it is important tonote that the surviva旭 and reproduc抄onofnestsaredrivenbyproximateprocesspname旭ytheaccessofnestsvia the workers within themp to resourcess This inding supportsotherworkthathasdetectednoevidencefortop､downpco旭ony､旭eve旭efectsonthestructureofpo旭ydomousco旭oniesｪE旭旭iseta旭spゴグゲジｫsTheextenttowhichnestscanbeconsideredaspartofthesameco旭onypsuper､co旭onyporsuper､organismisanimportantconsidera抄onwhenassessingp forexamp旭ep the 旭eve旭atwhichse旭ec抄onacts inaco旭onyｪHe旭anter士p StrassmannpCarri旭旭opｹQue旭旭erp ゴググゾqKennedypU旭旭erpｹHe旭anter士pゴグゲジqMofetpゴグゲゴｫs

Resourcesareotendistributedheterogeneous旭y in theenviron-

mentqpo旭ydomymaybeawaytomoreeicient旭yexp旭oit thesedis-persedresourcesｪCooketa旭spゴグゲザqHo旭wayｹCasepゴグググqLananeta旭spゴグゲゲqSchmo旭kepゴググゾｫsThenestandforagingnetworkofpo旭ydomousco旭oniescanbeviewedasatransporta抄onnetworktomoveresourcesfromfoodsourcestothenestsandthenbetweennestsｪCooketa旭spゴグゲジqLatyeta旭spゴグゲゲｫsTransporteiciencyreferstotheeasewithwhichresourcescanlowthroughanetworksInthepo旭ydomousnestsystempnestswithahighresourcelowareatpointsinthenetworkimportant for co旭ony､旭eve旭 resource redistribu抄on and thereforeco旭ony､旭eve旭transporteiciencyｪesgspCroteta旭spゴググ芦qPernaｹLatypゴグゲジｫs Ifnetworktransporteiciency isbeingretainedwithinaco旭-onypnestsandtrai旭swithahigherresourcelowpandthereforegreaterimportance for eiciencypmaybemore 旭ike旭y to survive than thosewitha 旭owerbetweenesssWefoundthatnestswithahighresourcelowaremore 旭ike旭ytosurvivethannestswitha 旭owerbetweenesssHoweverptrai旭swithahigherresourcelowarenotmore旭ike旭ytosur-vivethantrai旭switha旭owerbetweenesssEicienttransportstructuresare therefore not preferen抄a旭旭y being retained in the nest networksAddi抄ona旭旭yptheprocessofnestfounda抄onwi旭旭a旭sodegradeeicienttransportstructuressAsnewnestsarefoundedbynestswithahightransportva旭ueｪahighlowofresourcesｫpthiswi旭旭a旭terthestructureoftheco旭onyaroundthatnestsThisestab旭ishmentofnewnestsandtrai旭swi旭旭changetheprevious旭yexis抄ngpeicientp transportstructuress Inasystemwhichisunderstrongse旭ec抄vepressureforeiciencypit isexpectedthathigh旭yefec抄vetransportstructureswi旭旭beretainedsAsthisisnotthecaseintheredwoodantpo旭ydomousco旭oniespitmaybethattransporteiciencyisnotunderstrongse旭ec抄vepressures

Thelowofresourcesthroughapar抄cu旭arnestcanchangeover抄meduetoothernestsinthenetworkbeinggainedand旭ostsTheinte-

gratednatureofthesystemmeansthatagivennestcou旭dmaintainthesameconnec抄onstoneighboringnestsandtreesbuts抄旭旭undergoachangeintheamountofresourcesavai旭ab旭etoitｪandthereforeitschancesofsurvivingandreproducingｫpduetonestsbeingabandoned

F IGURE  ザ科Nestswithahighernorma旭izedbetweenessaremore旭ike旭ytoactasfoundersofnewneststhannestswitha旭owernorma旭izedbetweenessｪAoDrχゴ┎ゾsゼpdf┎ゴpp ┎ sググ芦ｫsForthefigurepbetweenessiscategorizedasグpゲp旭owｪ┑グsゴズｫpmediumｪグsゴズ･グsゼズｫporhighｪ┒グsゼズｫ

科架 科 | 科ゾELLIS ET AL.

orfoundede旭sewhereintheco旭onysNestsinunproitab旭eareaspandthereforewitha旭owresourcelowparemore旭ike旭ytobeabandonedthannests inproitab旭eareassThesedynamicswi旭旭 resu旭t in theco旭-onymovingtowardresourcesandawayfromunproitab旭eareassFora spa抄a旭旭y embeddednetworkp suchas apo旭ydomousnetworkp thismovementisphysica旭movementofnodessInnetworkswhicharenotspa抄a旭旭yembeddedpsuchassocia旭networkspthisprocesscou旭dresu旭tin anetworkc旭usteringaroundcertainnodesp forexamp旭e individu-

a旭swith informa抄onsThe reversecou旭da旭sooccurqanetworkcou旭dc旭uster away from speciic nodesp for examp旭e diseased individua旭sinasocia旭networksThesechangesinthenetworkstructurearese旭f､organizedpresu旭抄ngfromse旭ec抄vepressurebasedonan individua旭vsposi抄oninthenetworks

Thenestnetworksofpo旭ydomousantco旭oniesarepinsomewayspana旭ogous to the socia旭 networks of individua旭 organismss Like indi-vidua旭spantnestscansurviveandreproduceｪinthesenseoffoundingnewnestsｫsTherearep howeverp crucia旭 diferencessForexamp旭ep thedeathofanindividua旭anima旭inasocia旭networkhasdirectitnesscon-

sequencess In contrastp a旭though abandoning anestwi旭旭 resu旭t in the旭ossoftheresourcesinvestedinconstruc抄ngthenestｪwhichmaybeconsiderab旭eｫpitisun旭ike旭ytoresu旭tinthedeathoftheantsinthenestqtheywi旭旭simp旭yjoinothernestsintheco旭onysDespitetheseimportantdiferencesppo旭ydomousant co旭oniesmaybeusefu旭mode旭sof socia旭networkssSimi旭artotheantnestnetworksptheposi抄onofanindividua旭inasocia旭networkcanhaveimportantconsequencesfortheiraccesstop forexamp旭ep informa抄on ｪesgspB旭onderｹDornhauspゴグゲゲqFarinepAp旭inpShe旭donpｹHoppitpゴグゲズｫanddisease ｪesgspCrosseta旭spゴググジqOterstaterｹThomsonpゴググゼｫsHoweverp旭inkingthesenetworkposi-抄onefectstothe旭ifehistoryofindividua旭sischa旭旭engingpduetothedif-icu旭抄esinco旭旭ec抄ngsuicient旭yhigh､qua旭itytempora旭datatoa旭旭owthenetworkstobeexamineddynamica旭旭yｪCroteta旭spゴググ芦pゴグゲゲqKurverseta旭spゴグゲジqWhiteheadpゴググ芦ｫsUsingthepo旭ydomousnestnetworkspwehavedemonstrated thatnetworkposi抄oncanhavean importantinluenceonthesurviva旭ppopu旭a抄onchangepandbuddinginadynamicsystemsThisprovidesausefu旭basis forexamining the importanceofnetworkposi抄oninotherbio旭ogica旭systemssuchassocia旭systemss

The network dynamics observed in these po旭ydomous co旭oniesi旭旭ustratethepoten抄a旭feedbackbetweentheindividua旭旭eve旭andthesystem旭eve旭inbio旭ogica旭networkssTheposi抄onofanindividua旭withinabio旭ogica旭systemcanafectthatindividua旭vsexposuretopforexam-

p旭epfoodpmatespinforma抄onpanddiseaseｪesgspAp旭inpFarinepMorand､FerronpｹShe旭donpゴグゲゴqChrist旭eyeta旭spゴググズqOhｹBadyaevpゴグゲグｫsThestructureofthenetworkispinturnpafectedbythenodeswithinthenetworksForexamp旭eptheovera旭旭paternofinterac抄onsbetweenindividua旭s in a systemcanbe inluencedby avarietyof bio抄c andabio抄cfactorssuchasfoodavai旭abi旭itypsexdemographicspandseasonｪBrentpMacLarnonpP旭atpｹSemp旭epゴグゲザqDardenpJamespRamnarinepｹCrotpゴググゾqFostereta旭spゴグゲゴｫsThenestswithinpo旭ydomousco旭-onieshigh旭ighthowtheseefectscanbereciproca旭inadynamicsys-temsDiferen抄a旭surviva旭andreproduc抄onofnodes inasystemwi旭旭changethestructureofthenetworkasnewnodesappearandothersdisappears Thiswi旭旭p in turnp change an individua旭vs re旭a抄ve posi抄onwithinthenetworkpa旭teringitschancesofsurvivingandreproducings

Thenetworkpthereforepwi旭旭becon抄nua旭旭yrestructuringpresu旭抄nginadynamicsystemwhichisnotstab旭ethrough抄mesDynamicprocesseswi旭旭reactdiferent旭ytosta抄csystemswhenfacingeco旭ogica旭andenvi-ronmenta旭changesｪKurverseta旭spゴグゲジｫs

In conc旭usionpwe found that the surviva旭 and budding of nestswithinpo旭ydomousFormica lugubrisco旭oniesarere旭atedtotheirposi-抄oninthetrai旭networksTheseresu旭tshigh旭ighthowapparent旭ydispa-rateen抄抄esinabio旭ogica旭systemcanbeintegratedintoafunc抄ona旭eco旭ogica旭unitsIta旭soshowshowindirectaccesstoresourcespthroughothersinaresourceexchangesystempcanhaveimportanteco旭ogica旭consequencess

ACKNOWLEDGMENTS

We wou旭d 旭ike to thank Duncan Procterp Phi旭ip Buckham､BonnetpJonPitchfordpKanchonDasmahapatrapandLotaSundstr塾mforva旭u-

ab旭ediscussionandcommentsonthemanuscriptsWea旭sothanktheEvo旭u抄onandTheore抄ca旭Eco旭ogyresearchgroupsattheUniversityofYorkforhe旭pfu旭discussionssWewou旭daddi抄ona旭旭y旭iketothankChrisMi旭nerpCar旭Hawkepandthestafandvo旭unteersattheNa抄ona旭TrustvsLongshawestatefortheirhe旭pandsupportsSEacknow旭edgesfundingfromtheNa抄ona旭Environmenta旭ResearchCounci旭andtheNa抄ona旭TrustsEJHRacknow旭edgesfundingfromtheRoya旭Societys

CONFLICT OF INTEREST

Nonedec旭areds

DATA ACCESSIBILITY

Dataareprovidedinthesupp旭ementarymateria旭ss

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ゲグ科 |科 科架 ELLIS ET AL.

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