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Evolutionary Applications 201912175ndash186 emsp|emsp175wileyonlinelibrarycomjournaleva
Received1January2018emsp |emsp Revised19July2018emsp |emsp Accepted20July2018DOI 101111eva12690
O R I G I N A L A R T I C L E
Color polarization vision mediates the strength of an evolutionary trap
Bruce A Robertson1 emsp|emspGaacutebor Horvaacuteth2
ThisisanopenaccessarticleunderthetermsoftheCreativeCommonsAttributionLicensewhichpermitsusedistributionandreproductioninanymediumprovidedtheoriginalworkisproperlycitedcopy2018TheAuthorsEvolutionary ApplicationspublishedbyJohnWileyampSonsLtd
1DivisionofScienceMathematicsandComputingBardCollegeAnnandale-on-HudsonNewYork2EnvironmentalOpticsLaboratoryDepartmentofBiologicalPhysicsPhysicalInstituteELTEEoumltvoumlsLoraacutendUniversityBudapestHungary
CorrespondenceBruceARobertsonDivisionofScienceMathematicsandComputingBardCollegeAnnandale-on-HudsonNYEmailbrobertsbardedu
Funding information FundingforthisstudywasgenerouslyprovidedbyBardCollege
AbstractEvolutionarytrapsarescenariosinwhichanimalsarefooledbyrapidlychangingcon-ditionsintopreferringpoor-qualityresourcesoverthosethatbetterimprovesurvivalandreproductivesuccessThemaladaptiveattractionofaquaticinsectstoartificialsourcesofhorizontallypolarizedlight(egglassbuildingsasphaltroads)hasbecomeafirstmodelsystembywhichscientistscaninvestigatethebehavioralmechanismsthatcausetrapstooccurWeemploythisfield-basedsystemtoexperimentallyinves-tigate(a)inwhichportion(s)ofthespectrumarepolarizationallywater-imitatingre-flectorsattractivetonocturnalterrestrialandaquaticsinsectsand(b)whichmodernlamptypesresultingreaterattractioninthistypicalkindofnocturnalpolarizedlightpollutionWefound thatmostaquatic taxaexhibitedpreferences for lampsbasedupontheircolorspectramosthavinglowestpreferenceforlampsemittingblueandredlightYetdespitepreviouslyestablishedpreferenceforhigherdegreesofpolari-zationofreflectedlightmostaquaticinsectfamilieswereattractedtotrapsbasedupontheirunpolarizedspectrumChironomidmidgesaloneshowedapreferenceforthecoloroflamplightinboththehorizontallypolarizedandunpolarizedspectraindi-catingonlythisfamilyhasevolvedtouselightinthiscolorrangeasasourceofinfor-mationtoguideitsnocturnalhabitatselectionTheseresultsdemonstratethatthecolorofartificial lightingcanexacerbateorreduce itsattractivenesstoaquatic in-sectsbutthatthestrengthofattractivenessofnocturnalevolutionarytrapsandsotheirdemographicconsequences isprimarilydrivenbyunpolarized lightpollutionThisfocusesmanagementattentiononlimitingbroad-spectrumlightpollutionaswellasitsintentionaldeploymenttoattractinsectsbacktonaturalhabitats
K E Y W O R D S
aquaticinsectbehaviorcolorpreferencecolorsensitivityevolutionarytraplightpollutionmaladaptationpolarizedlightpollution
1emsp |emspINTRODUC TION
Animalsuselightasasourceofinformationtoguidebehaviorssuchasorientation(LorneampSalmon2007)reproduction(RandBridarolliDries amp Ryan 1997) and foraging (Dwyer Bearhop Campbell amp
Bryant2013)Someofthislightispolarizedmeaningthatthedirec-tionoftheelectricfieldvectoroflightisorientednonrandomlysuchthat itvibratespredominantly insomedirection (eghorizontally)Skypolarizationpatternsmaybeusedbyanimalsasacuefororien-tation (DackeNilsson Scholtz ByrneampWarrant 2003Muheim
176emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
Sjoumlberg amp Pinzon-Rodriguez 2016) but the predominant sourceof polarized light on the earthrsquos surface iswater bodies (HorvaacutethKriskaMalikampRobertson2009)Whensun-ormoonlightreflectsfromwatersurfacesittypicallybecomeshorizontallypolarizedwithdegreesofpolarization(d)dependingonthedarknessbrightnessofthewaterbodyandontheangleofreflection(Horvaacutethetal2009)Historicallywater-seekingaquaticinsectshavereliablyusedthehor-izontalpolarizationof lightreflectedwithahighenoughdegreeofpolarizationdtolocatewatersourcestheyrequiretofeedmateandlayeggs(reviewedinHorvaacutethampCsabai2014)Todayitiswell-knownthattheabundanceofsmoothdark-coloredman-madeobjects(egasphalt glass buildings black cars solar panels dasymp80ndash100 atand near the Brewsterrsquos angle θBrewster=arctan n measured fromthenormalvectorofthereflectingsurfacewithrefractiveindexn KriskaHorvaacutethampAndrikovics1998KriskaCsabaiBodaMalikampHorvaacuteth2006KriskaBernaacutethFarkasampHorvaacuteth2009Horvaacutethetal20092010)canpolarizelightmorestrongly(iewithhigherd)thananynaturalwaterbody(dasymp30ndash80FlamariqueampBrowman2001 Horvaacuteth 2014) andwater-seeking aquatic insects are pref-erentially attracted to oviposit upon these supernormally intensesources of horizontal polarization where their eggs fail to hatch(Horvaacutethetal2010Kriskaetal199820062009)Adults thendietypicallywithnosecondopportunitytoreproduceSuchcasesofanthropogenicchangewhereevolvedcue-resourcecorrelationsareuncoupledandanimalscometoprefer inferiorhabitatsareknownasecologicaltraps(DwernychukampBoag1972)whichareonetypeofthebroadercategoryofevolutionarytrapsinwhichanytypeofbehaviorissimilarlyaffected(SchlaepferRungeampSherman2002)Thesesevereformsofbehavioralmaladaptationarebeingdetectedat an increasing frequency (RobertsonampHutto 2006 RobertsonRehageampSih2013)areknowntoleadtosevereandrapidpopu-lationdeclines(DelibesFerrerasampGaona2001FletcherOrrockampRobertson2012KokkoampSutherland2001)andaffectabroadrange of insect taxa (Robertson etal 2013 Robertson Campbelletal2017Robertsonetal2018)
Whenplacedabovewaterbodiesorartificialpolarizers lampscreate both polarized (due to reflected light) and unpolarized (bydirectlight)photopollutionthatcanindividuallyorincombinationtriggermaladaptivebehaviorinadiverserangeofinsecttaxa(BodaHorvaacutethKriskaBlahoacuteampCsabai2014RobertsonCampbelletal2017Szaacutezetal2015)Yetspeciesdifferwidelyintheirsusceptibil-itytothesedifferentformsoflightpollution(RobertsonCampbelletal2017)andthespecificmechanismsbywhichtheseman-madelightsourcesinterfacewithevolvedbehavioralrulestotriggermal-adaptationremainpoorlyunderstood(Houmllkeretal2010Horvaacutethetal2009LongcoreampRich2004PerkinHoumllkerampTockner2014Robertson etal 2013) Artificial light sources (eg light-emittingdiodesmdashLEDshigh-pressure sodium [HPS] lamps)differ fromsun-lightandmoonlight in thatdifferent lamptypesemphasizeoren-tirelyomitparticularwavelengthsoflight(GastonVisserampHoumllker2015)Whenitreflectsfromaman-madeobjectlamplightwillnotbe polarized evenly across the visible spectrumWavelengths re-flectedwith highest intensity are polarized the least that iswith
lowestdegreeofpolarization (HegeduumlsampHorvaacuteth2004) Incon-trastapartfromthecasewhenthesunandmoonareclosetothehorizonsun-andmoonlightarerelativelyuniforminintensityacrossvisiblewavelengths(Gastonetal2015)andsopolarizationofnat-urallightviareflectionfromcolorless(ieblackgreyorwhite)waterbodieswillresultinasimilarlyalmostuniformdegreeofpolarizationacrossthevisiblespectrumCollectivelythismeansthatbyempha-sizingparticularwavelengthsoflightartificiallightpollutionhastheevolutionarilynoveleffectofdecouplingthehistoricallycorrelatedrelationship between the intensity of light and the degree of po-larizationofwater-reflectedlight Italsomeansthat ifnight-activewater-seekingflying insectsdisproportionatelyrelyupondifferenthorizontally polarized portions of the visible spectrum to locatewaterbodiesartificiallightinglocatednearartificialpolarizers(egasphaltglassbuildings)couldmakethoseobjectsmoreattractivetonocturnalovipositinginsectsthanactualwaterbodies
In this way lamps that emit different spectra of unpolarizedlighthave thecapabilityofexacerbating theseverityofecologicaltrapscausedbynocturnalpolarizedlightpollutionortoreducetheirdemographic impacts because stronger habitat preferences trig-ger ecological trapswithmore severedemographic consequences(Fletcheretal2012HaleampSwearer2016Robertsonetal2013)Global-scale shifts toward broader spectrum street lighting (egLEDs) for theirenergyefficiencyandsafetybenefitswill result inshiftsintheavailablewavelengthsofnocturnalpolarizedlightacrosslargelandscapeswithpotentialtoexacerbateormitigatemaladap-tivebehavioralresponsesinwater-seekinginsects
We designed a field experiment to ask (a) whether nocturnalwater-seekingaquatic insects relyupondifferent spectraofpolar-izedlighttolocatewaterbodiesand(b)whichlamptypes(andpor-tionsoftheirvisiblespectra) insectsarepreferentiallyattractedtoand sowhich sensory cuesmitigate or exacerbate the severity ofevolutionary traps triggered by nocturnal polarized light pollutionOur approach examines variation in riverside captures of sevenaquatic and six terrestrial night-active insect families in simulatedwater bodies (white and black oil-filled trays) illuminated by LEDmetal-halide(MH)orHPSlampsWeinterpretdifferentiallyhighercaptures among lamp types illuminating oil-filled horizontal whitetrays (full-spectrumreflectorsofunpolarized light)asanattractiontoaparticularspectrumofunpolarizedlightproducedbythatlampandreflectedbythetrayDifferentialattractiontoaparticularlamptreatment illuminatingblack trays (full-spectrumreflectorsofhori-zontallypolarizedlight)isinterpretedasevidenceofpreferentialat-tractiontoparticularspectraofhorizontalpolarizationInteractionsbetweentrayandlamptypessuggestpreferentialattractiontohori-zontallypolarizedlightofaspectrumthatdiffersfromthepreferredspectrumofunpolarizedlightandvice versaindicatingindependenceof detectionof thewavelength andpolarization of reflected lightTerrestrialtaxaarecommonlyattractedtobrightunpolarizedlights(Nowinszky2003NowinszkyKissSzentkiraacutelyiPuskaacutesampLadaacutenyi2012) andempirical studies suggest their ventral eye regiondoesnotperceivethepolarizationof lightcomingfrombelow(reviewedinHorvaacuteth2014RobertsonCampbelletal2017)butevidenceis
emspensp emsp | emsp177ROBERTSON aNd HORVAacuteTH
limitedWeexaminethebehavioralresponsesofterrestrialtaxatopolarizedlighttreatmentstoconfirmthesenegativeresponsesatourstudysitesandsocontextualizetheimpactsofpolarizedandunpo-larizedlighttreatmentsonaquaticinsects
2emsp |emspMATERIAL S AND METHODS
21emsp|emspStudy sites and experimental design
Weselected study siteson fivedifferent tributariesof theHudsonRiverinsouthernNewYorkStateUSA(Figure1a)Studysiteswereidentifiedinsparselypopulatedareasalongheavilyforestedrivercor-ridorsWechoseresidentialpropertiesthatmaintainedmownlawnsextendingfromthehighwaterlineinlandatleast60mtoensuresuffi-cientareaforourexperimentandsothatvegetationwouldnotimpede
insectsrsquolinesofsightormovementtowardexperimentaltestsurfacesIn2013wetrappedflyingemergentaquaticinsectstwotimesateachsiteVisit1)June17ndashJuly6andVisit2)July10ndash24Eachtrappingses-sionlastedfor120minbeginningexactly30minaftersunset
We used black and white trays (60times40cm) filled with trans-parent common salad oil to capture insects and assess their rela-tivepreferencefortestsurfacesvaryingintheira)illuminationtypeandb)degreeofpolarizationofreflectedlight(dFigures1band2)Weplaced threeofeachof the two traycolors (blackwhite) inarowparallel totheriverbankatadistanceof5metersspaced05mapartAboveeachofthetrayswesuspendeda180-wattarrayof LED a150-wattHPSbulbor a150-wattMHbulb (Figure1b)WeusedanOceanOpticsUSB2000+spectrometertomeasuretheabsoluteirradiance(μWcm2nm)ofeachlightsourceasafunctionofwavelengthWeselectedlampsofwattagesthatproducedsim-ilaroverall luminosity (11000ndash12000 lumens) among lamp typesthough at differentwavelengths (Figure3)We chose a relativelynarrowband(yellowgreen550ndash630nm)HPSsourceaLEDsourcethatpeaksattwocomplementaryrangesofwavelengths(blue420ndash480nmred620ndash680nm)withredwavelengthsbeingmorecom-monatdawnandduskandaMHlampthatproduceslightatnearlyallvisiblewavelengths(Figure3)Eachlightsourcewasplacedinsideadownwardfacing35times54cmaluminumreflectorpaintedblackontop tominimize itsvisual signatureandelevated30cmabove thetest surface In thisway therewere twotreatmentsofeach lamptypeilluminatingbothwhiteandblacktraysLampswerehunglowabove test surfaces (25cm) and shaded to illuminate the test sur-faces such that insect responseswould be based upon their per-ceptionofonlytray-reflected(notdirect)lightexceptatveryshortrange(afterRobertsonCampbelletal2017)
We exposed these six combinations of lamp spectra and traycolortreatmentstoaquaticinsectsduringeachofthetwovisitstoeachofthefivestudysitesAdjusted(seeStatisticalAnalysisbelow)capturesfromeachofthetwosamplingsessionsatastudysitewerecombinedpriortoanalysis leadingtoatotalsamplesizeofn=30forthestudyWerandomizedthelocationoftraysandlamptypesinourstudytoavoidbias incapturesassociatedwiththephysicalpositionoftreatmentsintheexperimentrelativetothesurroundingenvironmentandtheothertreatmentsThespatialarrangementoftreatmentcombinationswasrandomizedateachsitevisitwiththeexceptionthatlightsourcesofthesametypewereneverplacedim-mediatelynexttoeachotherThepositionsofthewhiteandblacktrays within a lighting treatment were exchanged every 20minAftereachsamplingsessionwepouredtraycontentsthroughfinecheeseclothtoseparateinsectswhichwerestoredin80ethanolforlateridentificationtothefamilylevel
22emsp|emspFocal taxa and experimental predictions
Wedefine aquatic insect taxa as thosewith a larval or adult life-historyphasedependentontheavailabilityofafreshwatersource(JohnsonampTriplehorn2004)Wefocusedouranalysisonaquaticinsect taxa known to exhibit stronger attraction to sources of
F IGURE 1emspa)LocationsoffieldexperimentsintheHudsonValleyregionofsouthernNewYorkStateStarsindicatelocationsofexperimentalsitesonfivetributariesoftheHudsonRiver(inwhite)locatedintwoofthesixcountiesoutlinedinblackb)Thespatialarrangementofoil-filledblackandwhitetraysilluminatedby1)high-pressuresodium(HPS)lamp2)light-emittingdiodes(LED)and3)metal-halide(MH)lampateachstudysiteOil-filledtraysoftwocolorsreflectlamplightandpolarizeittoagreater(black)orlesser(white)degreeandareplaced5metersfromtheedgeofstudystreams
178emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
horizontally polarized light with greater d-values Ephemeroptera(Horvaacuteth etal 2010 Kriska etal 1998 2006 Szaacutez etal 2015)Trichoptera(Horvaacutethetal2010KriskaMalikSzivaacutekampHorvaacuteth2008) Empididae (Robertson etal 2018) Simuliidae (Robertsonetal2018)andChironomidae(HorvaacutethMoacuteraBernaacutethampKriska2011Lerneretal20082011Robertsonetal2018)Whenseek-ingasuitableovipositionsitethesetaxatouchdownonthesurfaceofwaterbodies(Horvaacuteth2014Kriskaetal2006)butwillbecap-turedintheoiltrapsbecauseoilwetschitinpreventingtakeoff
23emsp|emspPolarizing properties of traps
Wemeasuredthereflectionndashpolarizationcharacteristicsofoil-filledtraysusingimagingpolarimetry(HorvaacutethampVarjuacute2004)inthered(650plusmn40nm=wavelength of maximal sensitivityplusmnhalf band-width of the polarimeter detector) green (550plusmn40nm) and blue(450plusmn40nm)portionsof thespectrumWeperformedpolarimet-ricmeasurementsinadarkroomwiththeopticalaxisaimeddown-wardwiththereflector(oil-filledtrays)aimedattheBrewsterrsquosangleθBrewster=arctan(n =15)=563degfromtheverticalcalculatedfortherefractiveindexn=15ofvegetableoilExposureswerestandardizedfor130thofasecondat400ISOAttheBrewsterrsquosanglesurface-reflectedlightisperpendiculartotherefractedraypenetratingthe
oilresultinginthehighestpossibledegreeofpolarizationThreeim-agesweretakenofeachobjectwiththepolarizationfilteratdiffer-entanglesThoseimageswereprocessedviaPolarworkscopysoftwareinto images illustrating thedegree (d) andangle (α)ofpolarizationat each pixelWe estimated the maximum degree of polarizationassociatedwithtraysbyquantifyingthefractionofthed-patternscomposed of black pixels using ImageJcopy softwareWe calculatedawindowsizerepresenting110ofthetraylengthandheightandestimatedmaximumdasthewindowgivingthemaximumvalueoutof20nonoverlappingsamplelocationswithintheimageofeachtraynonrandomlyfocusedonportionsofthetraywiththehighestvisualestimatesofdontheblack-to-whitescale
24emsp|emspStatistical analyses
Weexaminedtheeffectof1)lamptypeand2)thed(degreeofpolar-izationofreflectedlamplight)onthenumberofcapturedindividualswithineachinsectfamilyFamily-specificanalysesservetoillustratethe behavioral preferences of each taxon For these analyseswecreatedastandardizedabundancemetricthatwouldbereflectiveoftherelativefractionofindividualsattractedtoeachtreatmentandthatwasinsensitivetobiasassociatedwithdisproportionatelyhighor low captures at one study site relative to othersWe adjusted
F IGURE 2emspReflectionndashpolarizationcharacteristics(angleαmeasuredclockwisefromtheverticalanddegreedoflinearpolarizationofreflectedlight)ofsalad-oil-filledblack(left)andwhite(right)traysilluminatedbyhigh-pressuresodium(HPS)(top)metal-halide(center)andlight-emittingdiode(bottom)lampusedinthechoiceexperimentsmeasuredwithimagingpolarimetryinthered(650nm)green(550nm)andblue(450nm)portionsofthespectrumWhitetraysreflectedlamplightwithmuchlowerdegreesofpolarizationthanblacktraysThemaximumdvalueisgivenforeachtrayHPSlightwasreflectedwiththehighestdinthebluereflectedLEDlightwasmoststronglypolarizedinthegreenandMHlamplightwaspolarizedtoahighandmoresimilardegreeacrosstheredgreenandbluespectraBlacktraysconsistentlyreflectedhorizontallypolarizedlightWhitetraysreflectedprimarilyverticallypolarizedlightwiththeexceptionofhorizontalpolarizationofthemirrorimageofthelightsource
emspensp emsp | emsp179ROBERTSON aNd HORVAacuteTH
captures of each family associatedwith each test surface to be afractionof the total numberof captures for that visit at that sitethenmultipliedeachfractionby100roundingtothenearestwholeindividualWemodeledstandardizedcapturesusinggeneralizedlin-earmodels and fitmodels to negativebinomial or Tweedie distri-butionswitha log-link functionTweedie isa familyofprobabilitydistributionswhichhavepositivemass at zero but areotherwisecontinuousandincludethePoissondistributionwhichistypicalofcountdataNegativebinomialdistributionsarealsocommonlyusedtomodelcountdataWemodeled lampandtraytypeascategori-calvariablesinpredictingcapturesforeachfamilyandselectedtheerrordistributionmodelthatminimizedoverdispersion(ĉ)
3emsp |emspRESULTS
31emsp|emspPolarization imagery
Black trays were more efficient polarizers of reflected light thanwhite ones (Figure2) The black test surfaces reflected lightwithvery high degrees of polarization (dblack=53ndash88) including
valueshigherthanwaterisgenerallycapableof(dwater=30ndash80Horvaacuteth amp Varjuacute 2004 Horvaacuteth 2014) White trays reflectedvery low degrees of polarization (dwhite=15ndash21) Black testsurfaces reflected alwayshorizontally polarized light (because thehorizontally polarized light reflected from the airndashwater interfacewas dominating)whilewhite test surfaces reflectedmainly verti-callypolarizedlight(becausetheverticallypolarizedlightreflectedfromthetrayrsquosbottomandrefractedatthewaterndashairinterfacewasdominating)Thesmallmirrorimageoflightsourcesreflectedbythewhitetrayswashorizontallypolarized(becausethereagainthehori-zontallypolarizedlightreflectedfromthewatersurfacedominated)withmaximumd-values(Figure2)
Blacktraysreflectedlightwithlowerandhigherdinthosespec-tralrangesinwhichtheintensityofilluminatinglamplightwashigherandlowerrespectively(Figures2and3)HPSlightreflectedbytheblacktestsurfaceswasmostpolarized inthebluerange(Figure2)whereitsunpolarizedintensityisminimal(Figure3)MHlampsemit-tedmaximalintensityinthegreenspectralrange(Figure3)inwhichthereforethedof lightreflectedfromtheblacktrayswasminimal(Figure2)ContrarytothelatterlamptypetheLEDsusedwereless
F IGURE 3emspSpectraoflightreflectedfromthewhite(toprow)andblack(bottomrow)traysilluminatedbythreedifferenttypesofnocturnalilluminationhigh-pressuresodiumlamplight-emittingdiodesandmetal-halidelampWemeasuredtherelativeirradianceoflightenteringthecosine-correctingsensorofaspectrometer4cmabovethegeometriccenterofthetrayandaimingdownwardat45ointhedirectionofthemirrorimageofthelampabove
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
176emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
Sjoumlberg amp Pinzon-Rodriguez 2016) but the predominant sourceof polarized light on the earthrsquos surface iswater bodies (HorvaacutethKriskaMalikampRobertson2009)Whensun-ormoonlightreflectsfromwatersurfacesittypicallybecomeshorizontallypolarizedwithdegreesofpolarization(d)dependingonthedarknessbrightnessofthewaterbodyandontheangleofreflection(Horvaacutethetal2009)Historicallywater-seekingaquaticinsectshavereliablyusedthehor-izontalpolarizationof lightreflectedwithahighenoughdegreeofpolarizationdtolocatewatersourcestheyrequiretofeedmateandlayeggs(reviewedinHorvaacutethampCsabai2014)Todayitiswell-knownthattheabundanceofsmoothdark-coloredman-madeobjects(egasphalt glass buildings black cars solar panels dasymp80ndash100 atand near the Brewsterrsquos angle θBrewster=arctan n measured fromthenormalvectorofthereflectingsurfacewithrefractiveindexn KriskaHorvaacutethampAndrikovics1998KriskaCsabaiBodaMalikampHorvaacuteth2006KriskaBernaacutethFarkasampHorvaacuteth2009Horvaacutethetal20092010)canpolarizelightmorestrongly(iewithhigherd)thananynaturalwaterbody(dasymp30ndash80FlamariqueampBrowman2001 Horvaacuteth 2014) andwater-seeking aquatic insects are pref-erentially attracted to oviposit upon these supernormally intensesources of horizontal polarization where their eggs fail to hatch(Horvaacutethetal2010Kriskaetal199820062009)Adults thendietypicallywithnosecondopportunitytoreproduceSuchcasesofanthropogenicchangewhereevolvedcue-resourcecorrelationsareuncoupledandanimalscometoprefer inferiorhabitatsareknownasecologicaltraps(DwernychukampBoag1972)whichareonetypeofthebroadercategoryofevolutionarytrapsinwhichanytypeofbehaviorissimilarlyaffected(SchlaepferRungeampSherman2002)Thesesevereformsofbehavioralmaladaptationarebeingdetectedat an increasing frequency (RobertsonampHutto 2006 RobertsonRehageampSih2013)areknowntoleadtosevereandrapidpopu-lationdeclines(DelibesFerrerasampGaona2001FletcherOrrockampRobertson2012KokkoampSutherland2001)andaffectabroadrange of insect taxa (Robertson etal 2013 Robertson Campbelletal2017Robertsonetal2018)
Whenplacedabovewaterbodiesorartificialpolarizers lampscreate both polarized (due to reflected light) and unpolarized (bydirectlight)photopollutionthatcanindividuallyorincombinationtriggermaladaptivebehaviorinadiverserangeofinsecttaxa(BodaHorvaacutethKriskaBlahoacuteampCsabai2014RobertsonCampbelletal2017Szaacutezetal2015)Yetspeciesdifferwidelyintheirsusceptibil-itytothesedifferentformsoflightpollution(RobertsonCampbelletal2017)andthespecificmechanismsbywhichtheseman-madelightsourcesinterfacewithevolvedbehavioralrulestotriggermal-adaptationremainpoorlyunderstood(Houmllkeretal2010Horvaacutethetal2009LongcoreampRich2004PerkinHoumllkerampTockner2014Robertson etal 2013) Artificial light sources (eg light-emittingdiodesmdashLEDshigh-pressure sodium [HPS] lamps)differ fromsun-lightandmoonlight in thatdifferent lamptypesemphasizeoren-tirelyomitparticularwavelengthsoflight(GastonVisserampHoumllker2015)Whenitreflectsfromaman-madeobjectlamplightwillnotbe polarized evenly across the visible spectrumWavelengths re-flectedwith highest intensity are polarized the least that iswith
lowestdegreeofpolarization (HegeduumlsampHorvaacuteth2004) Incon-trastapartfromthecasewhenthesunandmoonareclosetothehorizonsun-andmoonlightarerelativelyuniforminintensityacrossvisiblewavelengths(Gastonetal2015)andsopolarizationofnat-urallightviareflectionfromcolorless(ieblackgreyorwhite)waterbodieswillresultinasimilarlyalmostuniformdegreeofpolarizationacrossthevisiblespectrumCollectivelythismeansthatbyempha-sizingparticularwavelengthsoflightartificiallightpollutionhastheevolutionarilynoveleffectofdecouplingthehistoricallycorrelatedrelationship between the intensity of light and the degree of po-larizationofwater-reflectedlight Italsomeansthat ifnight-activewater-seekingflying insectsdisproportionatelyrelyupondifferenthorizontally polarized portions of the visible spectrum to locatewaterbodiesartificiallightinglocatednearartificialpolarizers(egasphaltglassbuildings)couldmakethoseobjectsmoreattractivetonocturnalovipositinginsectsthanactualwaterbodies
In this way lamps that emit different spectra of unpolarizedlighthave thecapabilityofexacerbating theseverityofecologicaltrapscausedbynocturnalpolarizedlightpollutionortoreducetheirdemographic impacts because stronger habitat preferences trig-ger ecological trapswithmore severedemographic consequences(Fletcheretal2012HaleampSwearer2016Robertsonetal2013)Global-scale shifts toward broader spectrum street lighting (egLEDs) for theirenergyefficiencyandsafetybenefitswill result inshiftsintheavailablewavelengthsofnocturnalpolarizedlightacrosslargelandscapeswithpotentialtoexacerbateormitigatemaladap-tivebehavioralresponsesinwater-seekinginsects
We designed a field experiment to ask (a) whether nocturnalwater-seekingaquatic insects relyupondifferent spectraofpolar-izedlighttolocatewaterbodiesand(b)whichlamptypes(andpor-tionsoftheirvisiblespectra) insectsarepreferentiallyattractedtoand sowhich sensory cuesmitigate or exacerbate the severity ofevolutionary traps triggered by nocturnal polarized light pollutionOur approach examines variation in riverside captures of sevenaquatic and six terrestrial night-active insect families in simulatedwater bodies (white and black oil-filled trays) illuminated by LEDmetal-halide(MH)orHPSlampsWeinterpretdifferentiallyhighercaptures among lamp types illuminating oil-filled horizontal whitetrays (full-spectrumreflectorsofunpolarized light)asanattractiontoaparticularspectrumofunpolarizedlightproducedbythatlampandreflectedbythetrayDifferentialattractiontoaparticularlamptreatment illuminatingblack trays (full-spectrumreflectorsofhori-zontallypolarizedlight)isinterpretedasevidenceofpreferentialat-tractiontoparticularspectraofhorizontalpolarizationInteractionsbetweentrayandlamptypessuggestpreferentialattractiontohori-zontallypolarizedlightofaspectrumthatdiffersfromthepreferredspectrumofunpolarizedlightandvice versaindicatingindependenceof detectionof thewavelength andpolarization of reflected lightTerrestrialtaxaarecommonlyattractedtobrightunpolarizedlights(Nowinszky2003NowinszkyKissSzentkiraacutelyiPuskaacutesampLadaacutenyi2012) andempirical studies suggest their ventral eye regiondoesnotperceivethepolarizationof lightcomingfrombelow(reviewedinHorvaacuteth2014RobertsonCampbelletal2017)butevidenceis
emspensp emsp | emsp177ROBERTSON aNd HORVAacuteTH
limitedWeexaminethebehavioralresponsesofterrestrialtaxatopolarizedlighttreatmentstoconfirmthesenegativeresponsesatourstudysitesandsocontextualizetheimpactsofpolarizedandunpo-larizedlighttreatmentsonaquaticinsects
2emsp |emspMATERIAL S AND METHODS
21emsp|emspStudy sites and experimental design
Weselected study siteson fivedifferent tributariesof theHudsonRiverinsouthernNewYorkStateUSA(Figure1a)Studysiteswereidentifiedinsparselypopulatedareasalongheavilyforestedrivercor-ridorsWechoseresidentialpropertiesthatmaintainedmownlawnsextendingfromthehighwaterlineinlandatleast60mtoensuresuffi-cientareaforourexperimentandsothatvegetationwouldnotimpede
insectsrsquolinesofsightormovementtowardexperimentaltestsurfacesIn2013wetrappedflyingemergentaquaticinsectstwotimesateachsiteVisit1)June17ndashJuly6andVisit2)July10ndash24Eachtrappingses-sionlastedfor120minbeginningexactly30minaftersunset
We used black and white trays (60times40cm) filled with trans-parent common salad oil to capture insects and assess their rela-tivepreferencefortestsurfacesvaryingintheira)illuminationtypeandb)degreeofpolarizationofreflectedlight(dFigures1band2)Weplaced threeofeachof the two traycolors (blackwhite) inarowparallel totheriverbankatadistanceof5metersspaced05mapartAboveeachofthetrayswesuspendeda180-wattarrayof LED a150-wattHPSbulbor a150-wattMHbulb (Figure1b)WeusedanOceanOpticsUSB2000+spectrometertomeasuretheabsoluteirradiance(μWcm2nm)ofeachlightsourceasafunctionofwavelengthWeselectedlampsofwattagesthatproducedsim-ilaroverall luminosity (11000ndash12000 lumens) among lamp typesthough at differentwavelengths (Figure3)We chose a relativelynarrowband(yellowgreen550ndash630nm)HPSsourceaLEDsourcethatpeaksattwocomplementaryrangesofwavelengths(blue420ndash480nmred620ndash680nm)withredwavelengthsbeingmorecom-monatdawnandduskandaMHlampthatproduceslightatnearlyallvisiblewavelengths(Figure3)Eachlightsourcewasplacedinsideadownwardfacing35times54cmaluminumreflectorpaintedblackontop tominimize itsvisual signatureandelevated30cmabove thetest surface In thisway therewere twotreatmentsofeach lamptypeilluminatingbothwhiteandblacktraysLampswerehunglowabove test surfaces (25cm) and shaded to illuminate the test sur-faces such that insect responseswould be based upon their per-ceptionofonlytray-reflected(notdirect)lightexceptatveryshortrange(afterRobertsonCampbelletal2017)
We exposed these six combinations of lamp spectra and traycolortreatmentstoaquaticinsectsduringeachofthetwovisitstoeachofthefivestudysitesAdjusted(seeStatisticalAnalysisbelow)capturesfromeachofthetwosamplingsessionsatastudysitewerecombinedpriortoanalysis leadingtoatotalsamplesizeofn=30forthestudyWerandomizedthelocationoftraysandlamptypesinourstudytoavoidbias incapturesassociatedwiththephysicalpositionoftreatmentsintheexperimentrelativetothesurroundingenvironmentandtheothertreatmentsThespatialarrangementoftreatmentcombinationswasrandomizedateachsitevisitwiththeexceptionthatlightsourcesofthesametypewereneverplacedim-mediatelynexttoeachotherThepositionsofthewhiteandblacktrays within a lighting treatment were exchanged every 20minAftereachsamplingsessionwepouredtraycontentsthroughfinecheeseclothtoseparateinsectswhichwerestoredin80ethanolforlateridentificationtothefamilylevel
22emsp|emspFocal taxa and experimental predictions
Wedefine aquatic insect taxa as thosewith a larval or adult life-historyphasedependentontheavailabilityofafreshwatersource(JohnsonampTriplehorn2004)Wefocusedouranalysisonaquaticinsect taxa known to exhibit stronger attraction to sources of
F IGURE 1emspa)LocationsoffieldexperimentsintheHudsonValleyregionofsouthernNewYorkStateStarsindicatelocationsofexperimentalsitesonfivetributariesoftheHudsonRiver(inwhite)locatedintwoofthesixcountiesoutlinedinblackb)Thespatialarrangementofoil-filledblackandwhitetraysilluminatedby1)high-pressuresodium(HPS)lamp2)light-emittingdiodes(LED)and3)metal-halide(MH)lampateachstudysiteOil-filledtraysoftwocolorsreflectlamplightandpolarizeittoagreater(black)orlesser(white)degreeandareplaced5metersfromtheedgeofstudystreams
178emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
horizontally polarized light with greater d-values Ephemeroptera(Horvaacuteth etal 2010 Kriska etal 1998 2006 Szaacutez etal 2015)Trichoptera(Horvaacutethetal2010KriskaMalikSzivaacutekampHorvaacuteth2008) Empididae (Robertson etal 2018) Simuliidae (Robertsonetal2018)andChironomidae(HorvaacutethMoacuteraBernaacutethampKriska2011Lerneretal20082011Robertsonetal2018)Whenseek-ingasuitableovipositionsitethesetaxatouchdownonthesurfaceofwaterbodies(Horvaacuteth2014Kriskaetal2006)butwillbecap-turedintheoiltrapsbecauseoilwetschitinpreventingtakeoff
23emsp|emspPolarizing properties of traps
Wemeasuredthereflectionndashpolarizationcharacteristicsofoil-filledtraysusingimagingpolarimetry(HorvaacutethampVarjuacute2004)inthered(650plusmn40nm=wavelength of maximal sensitivityplusmnhalf band-width of the polarimeter detector) green (550plusmn40nm) and blue(450plusmn40nm)portionsof thespectrumWeperformedpolarimet-ricmeasurementsinadarkroomwiththeopticalaxisaimeddown-wardwiththereflector(oil-filledtrays)aimedattheBrewsterrsquosangleθBrewster=arctan(n =15)=563degfromtheverticalcalculatedfortherefractiveindexn=15ofvegetableoilExposureswerestandardizedfor130thofasecondat400ISOAttheBrewsterrsquosanglesurface-reflectedlightisperpendiculartotherefractedraypenetratingthe
oilresultinginthehighestpossibledegreeofpolarizationThreeim-agesweretakenofeachobjectwiththepolarizationfilteratdiffer-entanglesThoseimageswereprocessedviaPolarworkscopysoftwareinto images illustrating thedegree (d) andangle (α)ofpolarizationat each pixelWe estimated the maximum degree of polarizationassociatedwithtraysbyquantifyingthefractionofthed-patternscomposed of black pixels using ImageJcopy softwareWe calculatedawindowsizerepresenting110ofthetraylengthandheightandestimatedmaximumdasthewindowgivingthemaximumvalueoutof20nonoverlappingsamplelocationswithintheimageofeachtraynonrandomlyfocusedonportionsofthetraywiththehighestvisualestimatesofdontheblack-to-whitescale
24emsp|emspStatistical analyses
Weexaminedtheeffectof1)lamptypeand2)thed(degreeofpolar-izationofreflectedlamplight)onthenumberofcapturedindividualswithineachinsectfamilyFamily-specificanalysesservetoillustratethe behavioral preferences of each taxon For these analyseswecreatedastandardizedabundancemetricthatwouldbereflectiveoftherelativefractionofindividualsattractedtoeachtreatmentandthatwasinsensitivetobiasassociatedwithdisproportionatelyhighor low captures at one study site relative to othersWe adjusted
F IGURE 2emspReflectionndashpolarizationcharacteristics(angleαmeasuredclockwisefromtheverticalanddegreedoflinearpolarizationofreflectedlight)ofsalad-oil-filledblack(left)andwhite(right)traysilluminatedbyhigh-pressuresodium(HPS)(top)metal-halide(center)andlight-emittingdiode(bottom)lampusedinthechoiceexperimentsmeasuredwithimagingpolarimetryinthered(650nm)green(550nm)andblue(450nm)portionsofthespectrumWhitetraysreflectedlamplightwithmuchlowerdegreesofpolarizationthanblacktraysThemaximumdvalueisgivenforeachtrayHPSlightwasreflectedwiththehighestdinthebluereflectedLEDlightwasmoststronglypolarizedinthegreenandMHlamplightwaspolarizedtoahighandmoresimilardegreeacrosstheredgreenandbluespectraBlacktraysconsistentlyreflectedhorizontallypolarizedlightWhitetraysreflectedprimarilyverticallypolarizedlightwiththeexceptionofhorizontalpolarizationofthemirrorimageofthelightsource
emspensp emsp | emsp179ROBERTSON aNd HORVAacuteTH
captures of each family associatedwith each test surface to be afractionof the total numberof captures for that visit at that sitethenmultipliedeachfractionby100roundingtothenearestwholeindividualWemodeledstandardizedcapturesusinggeneralizedlin-earmodels and fitmodels to negativebinomial or Tweedie distri-butionswitha log-link functionTweedie isa familyofprobabilitydistributionswhichhavepositivemass at zero but areotherwisecontinuousandincludethePoissondistributionwhichistypicalofcountdataNegativebinomialdistributionsarealsocommonlyusedtomodelcountdataWemodeled lampandtraytypeascategori-calvariablesinpredictingcapturesforeachfamilyandselectedtheerrordistributionmodelthatminimizedoverdispersion(ĉ)
3emsp |emspRESULTS
31emsp|emspPolarization imagery
Black trays were more efficient polarizers of reflected light thanwhite ones (Figure2) The black test surfaces reflected lightwithvery high degrees of polarization (dblack=53ndash88) including
valueshigherthanwaterisgenerallycapableof(dwater=30ndash80Horvaacuteth amp Varjuacute 2004 Horvaacuteth 2014) White trays reflectedvery low degrees of polarization (dwhite=15ndash21) Black testsurfaces reflected alwayshorizontally polarized light (because thehorizontally polarized light reflected from the airndashwater interfacewas dominating)whilewhite test surfaces reflectedmainly verti-callypolarizedlight(becausetheverticallypolarizedlightreflectedfromthetrayrsquosbottomandrefractedatthewaterndashairinterfacewasdominating)Thesmallmirrorimageoflightsourcesreflectedbythewhitetrayswashorizontallypolarized(becausethereagainthehori-zontallypolarizedlightreflectedfromthewatersurfacedominated)withmaximumd-values(Figure2)
Blacktraysreflectedlightwithlowerandhigherdinthosespec-tralrangesinwhichtheintensityofilluminatinglamplightwashigherandlowerrespectively(Figures2and3)HPSlightreflectedbytheblacktestsurfaceswasmostpolarized inthebluerange(Figure2)whereitsunpolarizedintensityisminimal(Figure3)MHlampsemit-tedmaximalintensityinthegreenspectralrange(Figure3)inwhichthereforethedof lightreflectedfromtheblacktrayswasminimal(Figure2)ContrarytothelatterlamptypetheLEDsusedwereless
F IGURE 3emspSpectraoflightreflectedfromthewhite(toprow)andblack(bottomrow)traysilluminatedbythreedifferenttypesofnocturnalilluminationhigh-pressuresodiumlamplight-emittingdiodesandmetal-halidelampWemeasuredtherelativeirradianceoflightenteringthecosine-correctingsensorofaspectrometer4cmabovethegeometriccenterofthetrayandaimingdownwardat45ointhedirectionofthemirrorimageofthelampabove
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
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Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
emspensp emsp | emsp177ROBERTSON aNd HORVAacuteTH
limitedWeexaminethebehavioralresponsesofterrestrialtaxatopolarizedlighttreatmentstoconfirmthesenegativeresponsesatourstudysitesandsocontextualizetheimpactsofpolarizedandunpo-larizedlighttreatmentsonaquaticinsects
2emsp |emspMATERIAL S AND METHODS
21emsp|emspStudy sites and experimental design
Weselected study siteson fivedifferent tributariesof theHudsonRiverinsouthernNewYorkStateUSA(Figure1a)Studysiteswereidentifiedinsparselypopulatedareasalongheavilyforestedrivercor-ridorsWechoseresidentialpropertiesthatmaintainedmownlawnsextendingfromthehighwaterlineinlandatleast60mtoensuresuffi-cientareaforourexperimentandsothatvegetationwouldnotimpede
insectsrsquolinesofsightormovementtowardexperimentaltestsurfacesIn2013wetrappedflyingemergentaquaticinsectstwotimesateachsiteVisit1)June17ndashJuly6andVisit2)July10ndash24Eachtrappingses-sionlastedfor120minbeginningexactly30minaftersunset
We used black and white trays (60times40cm) filled with trans-parent common salad oil to capture insects and assess their rela-tivepreferencefortestsurfacesvaryingintheira)illuminationtypeandb)degreeofpolarizationofreflectedlight(dFigures1band2)Weplaced threeofeachof the two traycolors (blackwhite) inarowparallel totheriverbankatadistanceof5metersspaced05mapartAboveeachofthetrayswesuspendeda180-wattarrayof LED a150-wattHPSbulbor a150-wattMHbulb (Figure1b)WeusedanOceanOpticsUSB2000+spectrometertomeasuretheabsoluteirradiance(μWcm2nm)ofeachlightsourceasafunctionofwavelengthWeselectedlampsofwattagesthatproducedsim-ilaroverall luminosity (11000ndash12000 lumens) among lamp typesthough at differentwavelengths (Figure3)We chose a relativelynarrowband(yellowgreen550ndash630nm)HPSsourceaLEDsourcethatpeaksattwocomplementaryrangesofwavelengths(blue420ndash480nmred620ndash680nm)withredwavelengthsbeingmorecom-monatdawnandduskandaMHlampthatproduceslightatnearlyallvisiblewavelengths(Figure3)Eachlightsourcewasplacedinsideadownwardfacing35times54cmaluminumreflectorpaintedblackontop tominimize itsvisual signatureandelevated30cmabove thetest surface In thisway therewere twotreatmentsofeach lamptypeilluminatingbothwhiteandblacktraysLampswerehunglowabove test surfaces (25cm) and shaded to illuminate the test sur-faces such that insect responseswould be based upon their per-ceptionofonlytray-reflected(notdirect)lightexceptatveryshortrange(afterRobertsonCampbelletal2017)
We exposed these six combinations of lamp spectra and traycolortreatmentstoaquaticinsectsduringeachofthetwovisitstoeachofthefivestudysitesAdjusted(seeStatisticalAnalysisbelow)capturesfromeachofthetwosamplingsessionsatastudysitewerecombinedpriortoanalysis leadingtoatotalsamplesizeofn=30forthestudyWerandomizedthelocationoftraysandlamptypesinourstudytoavoidbias incapturesassociatedwiththephysicalpositionoftreatmentsintheexperimentrelativetothesurroundingenvironmentandtheothertreatmentsThespatialarrangementoftreatmentcombinationswasrandomizedateachsitevisitwiththeexceptionthatlightsourcesofthesametypewereneverplacedim-mediatelynexttoeachotherThepositionsofthewhiteandblacktrays within a lighting treatment were exchanged every 20minAftereachsamplingsessionwepouredtraycontentsthroughfinecheeseclothtoseparateinsectswhichwerestoredin80ethanolforlateridentificationtothefamilylevel
22emsp|emspFocal taxa and experimental predictions
Wedefine aquatic insect taxa as thosewith a larval or adult life-historyphasedependentontheavailabilityofafreshwatersource(JohnsonampTriplehorn2004)Wefocusedouranalysisonaquaticinsect taxa known to exhibit stronger attraction to sources of
F IGURE 1emspa)LocationsoffieldexperimentsintheHudsonValleyregionofsouthernNewYorkStateStarsindicatelocationsofexperimentalsitesonfivetributariesoftheHudsonRiver(inwhite)locatedintwoofthesixcountiesoutlinedinblackb)Thespatialarrangementofoil-filledblackandwhitetraysilluminatedby1)high-pressuresodium(HPS)lamp2)light-emittingdiodes(LED)and3)metal-halide(MH)lampateachstudysiteOil-filledtraysoftwocolorsreflectlamplightandpolarizeittoagreater(black)orlesser(white)degreeandareplaced5metersfromtheedgeofstudystreams
178emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
horizontally polarized light with greater d-values Ephemeroptera(Horvaacuteth etal 2010 Kriska etal 1998 2006 Szaacutez etal 2015)Trichoptera(Horvaacutethetal2010KriskaMalikSzivaacutekampHorvaacuteth2008) Empididae (Robertson etal 2018) Simuliidae (Robertsonetal2018)andChironomidae(HorvaacutethMoacuteraBernaacutethampKriska2011Lerneretal20082011Robertsonetal2018)Whenseek-ingasuitableovipositionsitethesetaxatouchdownonthesurfaceofwaterbodies(Horvaacuteth2014Kriskaetal2006)butwillbecap-turedintheoiltrapsbecauseoilwetschitinpreventingtakeoff
23emsp|emspPolarizing properties of traps
Wemeasuredthereflectionndashpolarizationcharacteristicsofoil-filledtraysusingimagingpolarimetry(HorvaacutethampVarjuacute2004)inthered(650plusmn40nm=wavelength of maximal sensitivityplusmnhalf band-width of the polarimeter detector) green (550plusmn40nm) and blue(450plusmn40nm)portionsof thespectrumWeperformedpolarimet-ricmeasurementsinadarkroomwiththeopticalaxisaimeddown-wardwiththereflector(oil-filledtrays)aimedattheBrewsterrsquosangleθBrewster=arctan(n =15)=563degfromtheverticalcalculatedfortherefractiveindexn=15ofvegetableoilExposureswerestandardizedfor130thofasecondat400ISOAttheBrewsterrsquosanglesurface-reflectedlightisperpendiculartotherefractedraypenetratingthe
oilresultinginthehighestpossibledegreeofpolarizationThreeim-agesweretakenofeachobjectwiththepolarizationfilteratdiffer-entanglesThoseimageswereprocessedviaPolarworkscopysoftwareinto images illustrating thedegree (d) andangle (α)ofpolarizationat each pixelWe estimated the maximum degree of polarizationassociatedwithtraysbyquantifyingthefractionofthed-patternscomposed of black pixels using ImageJcopy softwareWe calculatedawindowsizerepresenting110ofthetraylengthandheightandestimatedmaximumdasthewindowgivingthemaximumvalueoutof20nonoverlappingsamplelocationswithintheimageofeachtraynonrandomlyfocusedonportionsofthetraywiththehighestvisualestimatesofdontheblack-to-whitescale
24emsp|emspStatistical analyses
Weexaminedtheeffectof1)lamptypeand2)thed(degreeofpolar-izationofreflectedlamplight)onthenumberofcapturedindividualswithineachinsectfamilyFamily-specificanalysesservetoillustratethe behavioral preferences of each taxon For these analyseswecreatedastandardizedabundancemetricthatwouldbereflectiveoftherelativefractionofindividualsattractedtoeachtreatmentandthatwasinsensitivetobiasassociatedwithdisproportionatelyhighor low captures at one study site relative to othersWe adjusted
F IGURE 2emspReflectionndashpolarizationcharacteristics(angleαmeasuredclockwisefromtheverticalanddegreedoflinearpolarizationofreflectedlight)ofsalad-oil-filledblack(left)andwhite(right)traysilluminatedbyhigh-pressuresodium(HPS)(top)metal-halide(center)andlight-emittingdiode(bottom)lampusedinthechoiceexperimentsmeasuredwithimagingpolarimetryinthered(650nm)green(550nm)andblue(450nm)portionsofthespectrumWhitetraysreflectedlamplightwithmuchlowerdegreesofpolarizationthanblacktraysThemaximumdvalueisgivenforeachtrayHPSlightwasreflectedwiththehighestdinthebluereflectedLEDlightwasmoststronglypolarizedinthegreenandMHlamplightwaspolarizedtoahighandmoresimilardegreeacrosstheredgreenandbluespectraBlacktraysconsistentlyreflectedhorizontallypolarizedlightWhitetraysreflectedprimarilyverticallypolarizedlightwiththeexceptionofhorizontalpolarizationofthemirrorimageofthelightsource
emspensp emsp | emsp179ROBERTSON aNd HORVAacuteTH
captures of each family associatedwith each test surface to be afractionof the total numberof captures for that visit at that sitethenmultipliedeachfractionby100roundingtothenearestwholeindividualWemodeledstandardizedcapturesusinggeneralizedlin-earmodels and fitmodels to negativebinomial or Tweedie distri-butionswitha log-link functionTweedie isa familyofprobabilitydistributionswhichhavepositivemass at zero but areotherwisecontinuousandincludethePoissondistributionwhichistypicalofcountdataNegativebinomialdistributionsarealsocommonlyusedtomodelcountdataWemodeled lampandtraytypeascategori-calvariablesinpredictingcapturesforeachfamilyandselectedtheerrordistributionmodelthatminimizedoverdispersion(ĉ)
3emsp |emspRESULTS
31emsp|emspPolarization imagery
Black trays were more efficient polarizers of reflected light thanwhite ones (Figure2) The black test surfaces reflected lightwithvery high degrees of polarization (dblack=53ndash88) including
valueshigherthanwaterisgenerallycapableof(dwater=30ndash80Horvaacuteth amp Varjuacute 2004 Horvaacuteth 2014) White trays reflectedvery low degrees of polarization (dwhite=15ndash21) Black testsurfaces reflected alwayshorizontally polarized light (because thehorizontally polarized light reflected from the airndashwater interfacewas dominating)whilewhite test surfaces reflectedmainly verti-callypolarizedlight(becausetheverticallypolarizedlightreflectedfromthetrayrsquosbottomandrefractedatthewaterndashairinterfacewasdominating)Thesmallmirrorimageoflightsourcesreflectedbythewhitetrayswashorizontallypolarized(becausethereagainthehori-zontallypolarizedlightreflectedfromthewatersurfacedominated)withmaximumd-values(Figure2)
Blacktraysreflectedlightwithlowerandhigherdinthosespec-tralrangesinwhichtheintensityofilluminatinglamplightwashigherandlowerrespectively(Figures2and3)HPSlightreflectedbytheblacktestsurfaceswasmostpolarized inthebluerange(Figure2)whereitsunpolarizedintensityisminimal(Figure3)MHlampsemit-tedmaximalintensityinthegreenspectralrange(Figure3)inwhichthereforethedof lightreflectedfromtheblacktrayswasminimal(Figure2)ContrarytothelatterlamptypetheLEDsusedwereless
F IGURE 3emspSpectraoflightreflectedfromthewhite(toprow)andblack(bottomrow)traysilluminatedbythreedifferenttypesofnocturnalilluminationhigh-pressuresodiumlamplight-emittingdiodesandmetal-halidelampWemeasuredtherelativeirradianceoflightenteringthecosine-correctingsensorofaspectrometer4cmabovethegeometriccenterofthetrayandaimingdownwardat45ointhedirectionofthemirrorimageofthelampabove
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
178emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
horizontally polarized light with greater d-values Ephemeroptera(Horvaacuteth etal 2010 Kriska etal 1998 2006 Szaacutez etal 2015)Trichoptera(Horvaacutethetal2010KriskaMalikSzivaacutekampHorvaacuteth2008) Empididae (Robertson etal 2018) Simuliidae (Robertsonetal2018)andChironomidae(HorvaacutethMoacuteraBernaacutethampKriska2011Lerneretal20082011Robertsonetal2018)Whenseek-ingasuitableovipositionsitethesetaxatouchdownonthesurfaceofwaterbodies(Horvaacuteth2014Kriskaetal2006)butwillbecap-turedintheoiltrapsbecauseoilwetschitinpreventingtakeoff
23emsp|emspPolarizing properties of traps
Wemeasuredthereflectionndashpolarizationcharacteristicsofoil-filledtraysusingimagingpolarimetry(HorvaacutethampVarjuacute2004)inthered(650plusmn40nm=wavelength of maximal sensitivityplusmnhalf band-width of the polarimeter detector) green (550plusmn40nm) and blue(450plusmn40nm)portionsof thespectrumWeperformedpolarimet-ricmeasurementsinadarkroomwiththeopticalaxisaimeddown-wardwiththereflector(oil-filledtrays)aimedattheBrewsterrsquosangleθBrewster=arctan(n =15)=563degfromtheverticalcalculatedfortherefractiveindexn=15ofvegetableoilExposureswerestandardizedfor130thofasecondat400ISOAttheBrewsterrsquosanglesurface-reflectedlightisperpendiculartotherefractedraypenetratingthe
oilresultinginthehighestpossibledegreeofpolarizationThreeim-agesweretakenofeachobjectwiththepolarizationfilteratdiffer-entanglesThoseimageswereprocessedviaPolarworkscopysoftwareinto images illustrating thedegree (d) andangle (α)ofpolarizationat each pixelWe estimated the maximum degree of polarizationassociatedwithtraysbyquantifyingthefractionofthed-patternscomposed of black pixels using ImageJcopy softwareWe calculatedawindowsizerepresenting110ofthetraylengthandheightandestimatedmaximumdasthewindowgivingthemaximumvalueoutof20nonoverlappingsamplelocationswithintheimageofeachtraynonrandomlyfocusedonportionsofthetraywiththehighestvisualestimatesofdontheblack-to-whitescale
24emsp|emspStatistical analyses
Weexaminedtheeffectof1)lamptypeand2)thed(degreeofpolar-izationofreflectedlamplight)onthenumberofcapturedindividualswithineachinsectfamilyFamily-specificanalysesservetoillustratethe behavioral preferences of each taxon For these analyseswecreatedastandardizedabundancemetricthatwouldbereflectiveoftherelativefractionofindividualsattractedtoeachtreatmentandthatwasinsensitivetobiasassociatedwithdisproportionatelyhighor low captures at one study site relative to othersWe adjusted
F IGURE 2emspReflectionndashpolarizationcharacteristics(angleαmeasuredclockwisefromtheverticalanddegreedoflinearpolarizationofreflectedlight)ofsalad-oil-filledblack(left)andwhite(right)traysilluminatedbyhigh-pressuresodium(HPS)(top)metal-halide(center)andlight-emittingdiode(bottom)lampusedinthechoiceexperimentsmeasuredwithimagingpolarimetryinthered(650nm)green(550nm)andblue(450nm)portionsofthespectrumWhitetraysreflectedlamplightwithmuchlowerdegreesofpolarizationthanblacktraysThemaximumdvalueisgivenforeachtrayHPSlightwasreflectedwiththehighestdinthebluereflectedLEDlightwasmoststronglypolarizedinthegreenandMHlamplightwaspolarizedtoahighandmoresimilardegreeacrosstheredgreenandbluespectraBlacktraysconsistentlyreflectedhorizontallypolarizedlightWhitetraysreflectedprimarilyverticallypolarizedlightwiththeexceptionofhorizontalpolarizationofthemirrorimageofthelightsource
emspensp emsp | emsp179ROBERTSON aNd HORVAacuteTH
captures of each family associatedwith each test surface to be afractionof the total numberof captures for that visit at that sitethenmultipliedeachfractionby100roundingtothenearestwholeindividualWemodeledstandardizedcapturesusinggeneralizedlin-earmodels and fitmodels to negativebinomial or Tweedie distri-butionswitha log-link functionTweedie isa familyofprobabilitydistributionswhichhavepositivemass at zero but areotherwisecontinuousandincludethePoissondistributionwhichistypicalofcountdataNegativebinomialdistributionsarealsocommonlyusedtomodelcountdataWemodeled lampandtraytypeascategori-calvariablesinpredictingcapturesforeachfamilyandselectedtheerrordistributionmodelthatminimizedoverdispersion(ĉ)
3emsp |emspRESULTS
31emsp|emspPolarization imagery
Black trays were more efficient polarizers of reflected light thanwhite ones (Figure2) The black test surfaces reflected lightwithvery high degrees of polarization (dblack=53ndash88) including
valueshigherthanwaterisgenerallycapableof(dwater=30ndash80Horvaacuteth amp Varjuacute 2004 Horvaacuteth 2014) White trays reflectedvery low degrees of polarization (dwhite=15ndash21) Black testsurfaces reflected alwayshorizontally polarized light (because thehorizontally polarized light reflected from the airndashwater interfacewas dominating)whilewhite test surfaces reflectedmainly verti-callypolarizedlight(becausetheverticallypolarizedlightreflectedfromthetrayrsquosbottomandrefractedatthewaterndashairinterfacewasdominating)Thesmallmirrorimageoflightsourcesreflectedbythewhitetrayswashorizontallypolarized(becausethereagainthehori-zontallypolarizedlightreflectedfromthewatersurfacedominated)withmaximumd-values(Figure2)
Blacktraysreflectedlightwithlowerandhigherdinthosespec-tralrangesinwhichtheintensityofilluminatinglamplightwashigherandlowerrespectively(Figures2and3)HPSlightreflectedbytheblacktestsurfaceswasmostpolarized inthebluerange(Figure2)whereitsunpolarizedintensityisminimal(Figure3)MHlampsemit-tedmaximalintensityinthegreenspectralrange(Figure3)inwhichthereforethedof lightreflectedfromtheblacktrayswasminimal(Figure2)ContrarytothelatterlamptypetheLEDsusedwereless
F IGURE 3emspSpectraoflightreflectedfromthewhite(toprow)andblack(bottomrow)traysilluminatedbythreedifferenttypesofnocturnalilluminationhigh-pressuresodiumlamplight-emittingdiodesandmetal-halidelampWemeasuredtherelativeirradianceoflightenteringthecosine-correctingsensorofaspectrometer4cmabovethegeometriccenterofthetrayandaimingdownwardat45ointhedirectionofthemirrorimageofthelampabove
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
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BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
emspensp emsp | emsp179ROBERTSON aNd HORVAacuteTH
captures of each family associatedwith each test surface to be afractionof the total numberof captures for that visit at that sitethenmultipliedeachfractionby100roundingtothenearestwholeindividualWemodeledstandardizedcapturesusinggeneralizedlin-earmodels and fitmodels to negativebinomial or Tweedie distri-butionswitha log-link functionTweedie isa familyofprobabilitydistributionswhichhavepositivemass at zero but areotherwisecontinuousandincludethePoissondistributionwhichistypicalofcountdataNegativebinomialdistributionsarealsocommonlyusedtomodelcountdataWemodeled lampandtraytypeascategori-calvariablesinpredictingcapturesforeachfamilyandselectedtheerrordistributionmodelthatminimizedoverdispersion(ĉ)
3emsp |emspRESULTS
31emsp|emspPolarization imagery
Black trays were more efficient polarizers of reflected light thanwhite ones (Figure2) The black test surfaces reflected lightwithvery high degrees of polarization (dblack=53ndash88) including
valueshigherthanwaterisgenerallycapableof(dwater=30ndash80Horvaacuteth amp Varjuacute 2004 Horvaacuteth 2014) White trays reflectedvery low degrees of polarization (dwhite=15ndash21) Black testsurfaces reflected alwayshorizontally polarized light (because thehorizontally polarized light reflected from the airndashwater interfacewas dominating)whilewhite test surfaces reflectedmainly verti-callypolarizedlight(becausetheverticallypolarizedlightreflectedfromthetrayrsquosbottomandrefractedatthewaterndashairinterfacewasdominating)Thesmallmirrorimageoflightsourcesreflectedbythewhitetrayswashorizontallypolarized(becausethereagainthehori-zontallypolarizedlightreflectedfromthewatersurfacedominated)withmaximumd-values(Figure2)
Blacktraysreflectedlightwithlowerandhigherdinthosespec-tralrangesinwhichtheintensityofilluminatinglamplightwashigherandlowerrespectively(Figures2and3)HPSlightreflectedbytheblacktestsurfaceswasmostpolarized inthebluerange(Figure2)whereitsunpolarizedintensityisminimal(Figure3)MHlampsemit-tedmaximalintensityinthegreenspectralrange(Figure3)inwhichthereforethedof lightreflectedfromtheblacktrayswasminimal(Figure2)ContrarytothelatterlamptypetheLEDsusedwereless
F IGURE 3emspSpectraoflightreflectedfromthewhite(toprow)andblack(bottomrow)traysilluminatedbythreedifferenttypesofnocturnalilluminationhigh-pressuresodiumlamplight-emittingdiodesandmetal-halidelampWemeasuredtherelativeirradianceoflightenteringthecosine-correctingsensorofaspectrometer4cmabovethegeometriccenterofthetrayandaimingdownwardat45ointhedirectionofthemirrorimageofthelampabove
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
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Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
180emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
intenseinthegreen(Figure3)thusdofLEDlightreflectedbytheblacktestsurfaceswasmaximalinthegreen(Figure2)
32emsp|emspBehavioral responses of arthropods
Wecapturedatotalof75992adultaquaticinsectsin27familiesWefocusedouranalysison13familiesforwhichwewereabletofitmod-elswithoutoverdispersion(ĉlt40alldf=30Table1)Theseincludedsevenfamiliesofaquaticinsects(CaenidaeChironomidaeEmpididaeGlossosomatidae Heptageniidae Hydropsychidae and Isonychiidae)and six families of terrestrial insects (Cecidomyiidae CercopidaeCicadellidaeScarabaeidaeTipulidaeandGeometridae)HydropsychidandisonychiddatawerefitusingmodelswithaTweediedistributionwhiletheremainingfamilieswerefitwithnegativebinomialdistribu-tionsFitofdatatomodelswasgood(ĉ=095ndash391)
Allsevenaquaticfamiliesexhibitedapreferencefor lampandortraytype(Figure4Table1)Isonychidmayflieswerecapturedinpro-gressivelygreaternumbersinLEDandMHlampsthansodiumlamps(Figure4a) while caenid mayflies exhibited a preferential attractiontoLEDsandtowhitetestsurfacesingeneral(Figure4b)Danceflies(Empididae)exhibitedpreferenceonlyforwhitetestsurfacesregard-less of lamp type (Figure4c) Flat-headed mayflies (Heptageniidae)were captured in lowest abundance in LED treatments (Figure4d)Glossomatidandhydropsychidmayflieswerealsocapturedinlowestnumbers in LED-illuminated traps but were also more attracted towhite test surfaces (Figures4ef) Nonbiting midges (Chironomidae)weredifferentially attracted toLEDsassociatedwithwhite test sur-faces while exhibiting lowest preference for themwhen they were
associatedwithblackones(Figure4g)Amongterrestrialinsectsonlyone familyexhibiteddifferential attraction toanexperimental treat-ment Geometrid moths were attracted to sodium and MH lamps relativetoLEDs(Figure4m)
4emsp |emspDISCUSSION
Thereducedintensityofphotonsavailableatnighthasbeenthoughttomaketheprocessofcolorvisiondifficultandsoselectagainstitsuse(WarrantampDacke2011)Yetthespectralcompositionofarti-ficiallightisknowntoaffectdegreeofattractionbybothterrestrialandaquaticinsects(vanGrunsvenetal2014Longcoreetal2015PawsonampBader2014)andwefoundcolorpreferencewasubiqui-tousamongthenight-activeaquaticinsectswestudiedThisstudyrepresentsthefirstexperimentalinvestigationoftheroleofspec-tralpreferenceintriggeringnocturnalevolutionarytrapsinducedbypolarized-light-pollutingartificialreflectorsWefoundthataquaticinsect families exhibited distinct andmaladaptive preferences forspecificwavelengths of both unpolarized light and polarized lightreflectedfromblackorwhitetestsurfacesthathaveimplicationsfortheirconservationandthemanagementoflightpollution
Colorvisionandpreferenceareknownamongdiurnal(HobackSvatos SpomerampHigley 1999 RadwellampCamp 2009) and cre-puscular(Schwind19911995)aquaticinsectsWefoundspectrum-dependentphototaxis innocturnalaquatic insects to fall into fourcategories(a)AttractiontoMHlamps(Isonychiidae) (b)attractionto LED lamps (Caenidae) (c) preference for MH and HPS lamps
TABLE 1emspAbundancemodeloutputinformationrelativetothesevenmostabundantfamiliesofemergentaquaticinsectsandsixfamiliesofterrestrialinsectswithnoaquaticlife-historystagecapturedinthefieldexperiments
Family Common nameNo of individuals
of captures Model
χ2 values
Lamp Tray Interaction
Aquatic
Caenidae Squaregillmayflies 889 11 NB 695 407
Chironomidae Nonbitingmidges 14465 184 NB 1078 5864 5469
Empididae Danceflies 3622 46 NB 265 935
Glossosomatidae Saddle-casemayflies 4955 63 NB 913 480
Heptageniidae Flat-headedmayflies 28375 362 T 722 008
Hydropsychidae Net-spinningcaddisflies 16010 204 T 2310 1033
Isonychiidae Brush-leggedmayflies 1187 15 T 854 091
Terrestrial
Cecidomyiidae Gallmidges 5279 67 NB 058 280
Cercopidae Froghoppers 652 08 NB 495 126
Cicadellidae Leafhoppers 2624 33 NB 052 281
Scarabaeidae Scarabbeetles 222 03 NB 090 261
Tipulidae Craneflies 84 01 NB 399 360
Geometridae Inchwormmoths 85 01 NB 2870 078
NotesCapturesof individuals inthesegroupsweremodeledasfunctionsof lampandtraytypeDatawerefittoeithernegativebinomial (NB)orTweedie(T)distributionsWaldchi-squaredvaluesassociatedwitheachindependentvariablearegivenalongwiththeirstatisticalsignificance(plt005plt001plt0001)
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
emspensp emsp | emsp181ROBERTSON aNd HORVAacuteTH
relativetoLEDs(HeptageniidaeGlossosomatidaeHydropsychidae)and(d)polarization-dependentpreferencerelativetoblue-redLEDs(Chironomidae)Broad-spectrumMHlampswerethemostattractive
lamptypeforfiveofthesevenfamiliesThespectraofthespecial-izedblue-redLEDsweused in this studywere foundconsistentlymostattractiveonlybyafamilyoftinymayflies(Caenidae)known
F IGURE 4emspStandardizedabundanceofinsectfamiliescapturedbythedifferentlyilluminatedoiltrapsasafunctionofthetypeofillumination(Lamp)andthecolor(blackorwhite)oftheinsecttrap(tray)Blackandwhiteoil-filledtrayswereplacedbelowdownward-facinglampsofthreetypeshigh-pressuresodiumlight-emittingdiodeandmetal-halide(MseeFigure1)BlacktraysstronglypolarizethevisiblewavelengthsaswoulddarklakesandotherdarkwaterbodiesStandardizedabundanceistherelativenumberofcapturespertreatment(plusmnstandarderror)foreachfamilyofaquatic(a-g)andentirelyterrestrial(h-m)insectsStatisticalsignificanceofindependentvariablesisshownineachfigureplt005plt001plt0001(seeTable1)Lettersabandcaboveeachtreatmentrepresentresultsofpairwisecontrastposthoctestssuchthatcolumnsthatdonotsharethesameletterrepresentmeansthataredifferentatthep=005levelImagesaretypicalmorphologicalrepresentationsofeachtaxonomicgroup
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
182emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
frompreviousstudiestofindthesetwocolorsrelativelyunattractive(RadwellampCamp2009)OthermayfliesinthisstudypreferredMHandHPS lamplight over blue-red LEDswhich is themore typicalbehaviorofdiurnalEphemeroptera(Hobacketal1999RadwellampCamp2009)FiveofthesixterrestrialinsectfamiliesweexaminedexhibitednoclearpreferencewithrespecttospectraExceptionallygeometridmothswereequallyandmoreattractedtotheHPSandMHlampsrelativetoLEDsThisgroupisknownforitsaffinityforbroad-spectrumlightingandlampsemphasizingbluelight(BarghiniampdeMedeiros2012Batesetal2013LampharampKocifaj2013vanLangeveldeEttemaDonnersWallis-DeVriesampGroenendijk2011Somers-YeatesHodgsonMcGregorSpaldingampffrench-Constant2013)butinthisstudywereleastattractedtoLEDswhichemittedthewidest diversity of bluewavelengths and at highest intensityWhether the spectral preferenceswe identifiedoriginally evolvedtoguideanimalsduringanydiurnalorcrepuscularactivityperiodsisunclearItisclearthatthesecolorpreferencesarecurrentlyguid-ingnocturnalbehaviorandcouldrepresentadaptationsdesignedtoguidenocturnalhabitatselectionnavigationormateselectionandsomaybe adaptive non-adaptive ormaladaptive indifferentbe-havioralcontext
Onlyoneofthesevenaquaticandnoneoftheterrestrialinsectfamilies in this study exhibitedpatterns of capture indicating dif-ferentialattractiontoreflectedpolarizedlightspectraIfinsectsin
thisstudywereexhibitingattractionforhighdegreesofpolarizationof reflected light independent of its color they should have beencaptured more frequently in our water-imitating traps associatedwithHPSandLEDlampswhichproducedthehighestdegreesofre-flectedpolarizationbutpatternsofcapturesuggestedthatinsectswerenotusingpolarizedlighttoguidetheirhabitatselectiondeci-sions Insteadcaptureswerecommonly (fiveof thesevenaquaticfamilies)higherintreatmentswithnonpolarizingwhitetestsurfacesindicatingthatphototaxiswasdominantoverpolarotaxisNotablytheseaquaticinsectgroupshavebeenexperimentallydemonstratedtoexhibitpreferenceforwater-reflectedlightwithhigherdegreesofpolarization(EphemeropteraKriskaetal2009trichopteraKriskaetal 2008 Lerner etal 2008 Chironomidae Robertson etal2018) yet ignore thesehabitat selectionpreferenceswhen in thepresence of unpolarized lamp light which simulates their primarynavigationalcuethemoon(Bodaetal2014RobertsonCampbelletal 2017 Szaacutez etal 2015) Robertson Campbell etal (2017)concludedthatthisresultindicatesthatabehavioralcueevolvedtoguidebehaviorinonecontextcanbeexploitedtotriggeranevolu-tionarytrapinanotherAmoreproximateexplanationmaybethatpolarizationsensitivityishousedintheommatidialcellsthatdetectbrightness in insecteyes (eg thebrightnessndashpolarizationhypoth-esisLabhart2016)muchthesamewaythatPapiliobutterflyom-matidiacontainbothpolarizationandcolorsensitivityleadingtoan
F I G U R E 4emspContinued
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
emspensp emsp | emsp183ROBERTSON aNd HORVAacuteTH
inabilitytodistinguishbetweenthesetwomodalitiesoflightunderartificial laboratoryconditions (HegeduumlsampHorvaacuteth2004Kelber1999KelberThunellampArikawa2001)
Onlychironomidmidgesexhibitedpatternsofcapturesconsis-tentwithwavelength-specificpreferenceforhorizontallypolarizedlightatnightBecauseunpolarizedwavelengthsproducedathigh-estintensitybylampsarepolarizedtheleastbyreflection(Hegeduumlsamp Horvaacuteth 2004 Figures2 and 3) inference about evidence forcolor-dependentpolarizationpreferencemustalsofollowthisrulePreviousstudiesindicatedthatday-activechironomidsaredifferen-tiallyattractedtounpolarizedgreenbluelight(Burkett1998)butexperimental studiesof colorpreference in thisgroupare rare Inour studymidges homed to the blue andor red unpolarized LEDlightbuttheirreducedattractiontothehorizontallypolarizedlighttreatmentof this lamp type is actually consistentwithpreferencefor polarized light in this same spectrum In accordance with theinverse relationship between intensity and degree of reflectionndashpolarizationredblue lightwaspolarizedthe leastandgreenlightthemostbyourLEDlamppairedwiththeblackpolarizingreflector(Figure2)Andbecausepatternsofchironomidcaptureareinconsis-tentwithpreferencefortreatmentforthehighestdegreeofpolar-izationindependentofcolorwecaninferthatchironomidsexhibitcolor-dependent preference for red or blue horizontally polarizedlightThisresultissurprisingbecauseSchwindrsquos(19911995)stud-iesshowthatall18speciesofday-activeaquaticinsectshestudiedusedspecificspectraofpolarizedlighttoguidetheirhabitatselec-tionbehavior
Whymightsuchwavelength-specificpolarizationsensitivityorpreference evolveThe first possibility is that lakes and rivers ortheirbottomsdifferintheirabsorbancespectruminwaysthataretied to thesurvivalofeggsandyoungShallowwaterbodieswithabundantgreenplantoralgalsubstrates forexample reflect lightwiththehighestdegreesofhorizontalpolarizationintheultravioletandbluerangesofthespectrum(HorvaacutethampVarjuacute2004Schwind1995) Because blue light can penetratemore deeply into naturalclearwaterthanUVandred lightpreferencescouldreflectadap-tationtothedepthatwhichaquaticlife-stageslive(Schwind1995)Alternativelypreferencesforunpolarizedlightofdifferentcolorinnocturnal insectsmaybe aby-productof the evolutionary tuningof light sensitivity to spectra associated with food (HillWells ampWells1997)refugiafrompredationorconspecifics(EllersBoggsampMallet2003)
Nocturnalpolarizedlightpollution(Horvaacutethetal2009)differsfundamentallyfromitsdiurnalcounterpartinthatincontrasttothefullspectrumofsunlightlampspectracanbeaconstrainedsubsetofwavelengthsbut thosethatareproduceddifferdramatically intheintensityatwhichtheyareproduced(Gastonetal2015)Thusthedegreeofpolarizationofareflector(naturalorartificial)canbedecoupled from thewavelength it has been associatedwith overevolutionaryhistoryandtheavailabilityofwavelengthsof lighttobepolarizedcanvarydramaticallydependingonthelampitselfThisrepresentsanovelmechanismfortheformationofanevolutionarytrapFinallybycapturinginsectsimmediatelyupontouchingtheoil
surfaceourtrappingmethodpreventedinsectsfromusingolfactorycues toavoidunsuitablehabitatswhichmaybemorecommon inlongerlivedinsects(Lerneretal2011)
Collectivelyourresultsindicatethatnocturnalaquaticinsectsare capable of color-dependent polarization sensitivity that canexacerbateormitigatetheirresponsestolightpollutiondepend-ing on the spectrum of light pollution they are exposed to andthequalitiesofthereflectingsurfaces(egbuildingsasphalt) inthe vicinity Our results confirm that of previous studies (Bodaetal 2014 Robertson Campbell etal 2017 Szaacutez etal 2015)that nocturnal polarized light pollution is less important in trig-geringevolutionarytrapsthanunpolarizedlamplightandindicateapartialtrade-offbetweenspectrumintensityininducingevolu-tionary trapsviaunpolarizedvspolarized lightBroad-spectrumlamps thatemphasizeallwavelengthsat relativelyhigh intensity(egsomecommercialLEDs)mayhaveveryweakpolarizedlightsignaturesbutattractaquatic insects to theirunpolarizedspec-traBecausereducing theattractivenessofevolutionary traps isthemostimportantmethodtoreducetheirdemographicimpacts(Fletcheretal2012HaleampSwearer2016RobertsonOstfeldampKeesing2017RobertsonCampbelletal2017)ourresultssug-gest thatselectionofmorenarrowband (especiallyblue-redem-phasizing)lampsbyindividualsandgovernmentswillhavegreaterconservationbenefitsformoretaxaofnocturnalaquatic insectsthaneffortstoreducetheuseofpolarizingbuildingandroadwaymaterialsnearlakesandrivers
Giventhestrongnegativefitnessconsequencesthatevolution-arytrapshaveforindividualsnaturalselectionagainstmaladaptivebehaviors shouldbe strong and so traps shouldbeephemeral intime (Schlaepfer Sherman Blossey amp Runge 2005 Delibes etal2001) Indeeda recentstudy illustrates theevolutionof reducedsensitivitytounpolarizedlightpollutionbyurbanmoths(AltermattampEbert2016)Yet theymaypersisteven in the faceofpopula-tioncollapseifuseofpolarizedlighttolocatewaterorunpolarizedlighttonavigateiscanalized(Fletcheretal2012)Theintentionalconstructionofevolutionary traps tocontrolpestspecieshas re-cently been highlighted as a powerful new wildlife managementtool(HorvaacutethBlahoacuteEgriampLerner2014RobertsonOstfeldetal2017)theprinciplesofwhichcanbeadaptedtomanipulatenativespeciesexposed toevolutionary traps intomakingadaptivedeci-sions (egHorvaacutethetal2010)Forexampleaquatic insectsareattractedupwardtothesurfacesofbridgesbyunpolarizedbridge-lightingwheretheythenovipositonhorizontallypolarizingasphaltsurfaces(Maacutelnaacutesetal2011)butdownstream-facingLEDsplacedat thewater surface canpreferentially attract insects toovipositonthewater instead(Egrietal2017)Becauseunpolarizedlightisprimarilyresponsiblefordrawingaquaticinsectsawayfromsuit-ablehabitatsasimilarstrategyofplacingabrighterormorebroad-spectrum lamp above water can be employed where residencesor businesses expose lakes and streams to artificial light Globaltrends toward the wholesale adoption of broad-spectrum lampsfor nocturnal illumination portend the expansion of evolutionarytrapsforbothterrestrialandaquaticinsectsacrosstheglobeata
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
184emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
timewheninsectpopulationsarealreadyundercollapse(Hallmannetal 2017) and when continent-scale evolutionary traps havebeen implicatedascausal (monarchbutterfliesFaldynHunterampElderd 2018 European honey bee Kessler etal 2015 Rundloumlfetal2015)Anestimateoftheeffectsoftheevolutionarytrapef-fectassociatedwithunpolarizedstreetlampsinGermanyindicatedthat the light couldwipe outmore than 60 billion insects over asingle summer (Eisenbeis 2006) Selectionof narrow-wavelengthlamp types where safety andor cost is allowed will lead to themostsignificantreductionsintheattractionofaquatic(thisstudyRobertsonCampbelletal2017RobertsonOstfeldetal2017)andterrestrial(PawsonandBader2014Longcoreetal2015)in-sectstononhabitatswhilemorecarefulselectionofpotentialar-tificialpolarizersnearwaterbodies(egHorvaacutethetal2010)willhaveanadditionalthoughminorbenefit
ACKNOWLEDG EMENTS
We thank the logistical support of the Estrato Research andDevelopment Ltd (Dr Andraacutes Barta and Dr Ramoacuten HegeduumlsBudapestHungary)
CONFLIC T OF INTERE S T
Wedeclarenocompetinginterests
DATA ARCHIVING S TATEMENT
Data are available from theDryadDigital Repository httpsdoiorg105061dryad7m5cv66
ORCID
Bruce A Robertson httporcidorg0000-0002-6101-1569
R E FE R E N C E S
Altermatt F amp Ebert D (2016) Reduced flight-to-light behaviour ofmothpopulationsexposedtolong-termurbanlightpollutionBiology Letters1220160111httpsdoiorg101098rsbl20160111
Barghini A amp de Medeiros B A S (2012) UV radiation as an at-tractor for insects Leukos 9 47ndash56 httpsdoiorg101582LEUKOS20120901003
BatesAJSadlerJPEverettGGrundyDLoweNDavisGhellipYoungH(2013)AssessingthevalueoftheGardenMothSchemecitizen science dataset how does light trap type affect catchEntomologia Experimentalis et Applicata 146 386ndash397 httpsdoiorg101111eea12038
BodaPHorvaacutethGKriskaGBlahoacuteMampCsabaiZ(2014)PhototaxisandpolarotaxishandinhandNightdispersalflightofaquaticinsectsdistracted synergistically by light intensity and reflectionpolariza-tion Naturwissenschaften 5 385ndash395 httpsdoiorg101007s00114-014-1166-2
Burkett D A (1998) Light Color Attraction and Dietary SugarComposition for Several Mosquito (Diptera Culicidae) Species
FoundinNorthCentralFloridaDissertationUniversityofFloridaGainesville159pp
Dacke M Nilsson D E Scholtz C H Byrne M ampWarrant E J(2003) Insect orientation to polarizedmoonlightNature424 33httpsdoiorg101038424033a
DelibesMFerrerasPampGaonaP(2001)AttractivesinksorhowindividualbehaviouraldecisionsdeterminesourcendashsinkdynamicsEcology Letters4401ndash403httpsdoiorg101046j1461-0248200100254x
DwernychukLWampBoagDA (1972)Ducksnesting inassociationwithgulls-anecological trapCanadian Journal of Zoology50559ndash563httpsdoiorg101139z72-076
Dwyer R G Bearhop S Campbell H A amp Bryant D M (2013)Shedding lighton lightBenefitsofanthropogenic illuminationtoanocturnally foraging shorebird Journal of Animal Ecology82 478ndash485httpsdoiorg1011111365-265612012
EisenbeisG (2006)Artificial night lighting and insectsAttraction ofinsectstostreetlampsinaruralsettinginGermanyInCRichampTLongcore(Eds)Ecological consequences of artificial night lighting(pp281ndash304)WashingtonDCIslandPress
EgriAacute SzaacutezD FarkasA PereszleacutenyiAacuteHorvaacutethGampKriskaG(2017)Methodtoimprovethesurvivalofnight-swarmingmayfliesnearbridgesinareasofdistractinglightpollutionRoyal Society Open Science4171166httpsdoiorg101098rsos171166
EllersJBoggsCLampMalletJ (2003)TheevolutionofwingcolorMalematechoiceopposesadaptivewingcolordivergenceincoliasbutterfliesEvolution571100ndash1106
FaldynMJHunterMDampElderdBD(2018)ClimatechangeandaninvasivetropicalmilkweedAnecologicaltrapformonarchbut-terfliesEcologyl991031ndash1038httpsdoiorg101002ecy2198
FlamariqueINampBrowmanHI(2001)Foragingandprey-searchbe-haviourofsmalljuvenilerainbowtrout(Oncorhynchus mykiss)underpolarizedlightJournal of Experimental Biology2042415ndash2422
FletcherRJOrrockJLampRobertsonBA(2012)Howthetypeofanthropogenicchangealters theconsequencesofecological trapsProceedings of the Royal Society of London B2792546ndash2552httpsdoiorg101098rspb20120139
GastonK JVisserMEampHoumllkerF (2015)Thebiological impactsof artificial light at night The research challenge Philosophical Transactions of the Royal Society of London B37020140133httpsdoiorg101098rstb20140133
vanGrunsvenRHDonnersMKoekeeKTichelaarIvanGeffenKGroenendijkDhellipVeenendaalE (2014)Spectralcompositionoflightsourcesandinsectphototaxiswithanevaluationofexistingspectral responsemodels Journal of Insect Conservation18 225ndash231httpsdoiorg101007s10841-014-9633-9
HaleRampSwearerSE(2016)EcologicaltrapsCurrentevidenceandfuturedirectionsProceedings of the Royal Society of London B28320152647httpsdoiorg101098rspb20152647
Hallmann C A Sorg M Jongejans E Siepel H Hofland NSchwan H hellip de Kroon H (2017) More than 75 percent de-cline over 27 years in total flying insect biomass in protectedareasPLoS ONE12 e0185809 httpsdoiorg101371journalpone0185809
Hegeduumls R amp Horvaacuteth G (2004) Polarizational colours could helppolarization-dependent colour vision systems to discriminate be-tween shiny and matt surfaces but cannot unambiguously codesurface orientation Vision Research 44 2337ndash2348 httpsdoiorg101016jvisres200405004
HillPSMWellsPHampWellsH (1997)Spontaneousflowercon-stancy and learning in honey bees as a function of colourAnimal Behaviour54615ndash627httpsdoiorg101006anbe19960467
HobackWWSvatosTMSpomerSMampHigleyLG(1999)Trapcolorandplacementaffectsestimatesofinsectfamily-levelabundanceanddiversityinaNebraskasaltmarshEntomologia Experimentalis et Appli cata91393ndash402httpsdoiorg101046j1570-7458199900507x
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
emspensp emsp | emsp185ROBERTSON aNd HORVAacuteTH
HoumllkerFMossTGriefahnBKloasWVoigtCCHenckelDhellipTocknerK(2010)ThedarksideoflightAtransdisciplinaryresearchagendaforlightpollutionpolicyEcology and Society1513[online]URLhttpwwwecologyandsocietyorgvol15iss4art13
HorvaacutethG (Ed) (2014)Polarized light and polarization vision in animal sciencesBerlinHeidelbergSpringer
Horvaacuteth G BlahoacuteM Egri A Kriska G Seres I amp Robertson B(2010)Reducing themaladaptive attractivenessof solarpanels topolarotactic insectsConservation Biology 24 1644ndash1653 httpsdoiorg101111j1523-1739201001518x
Horvaacuteth G Blahoacute M Egri A amp Lerner A (2014) Applying polar-ization-based traps to insect control InGHorvaacuteth (Ed)Polarized Light and Polarization Vision in Animal Sciences(pp561ndash584)BerlinHeidelbergSpringer
HorvaacutethGampCsabaiZ (2014)Polarizationvisionofaquatic insectsInGHorvaacuteth(Ed)Polarized light and polarization vision in animal sci-ences(pp113ndash145)BerlinHeidelbergSpringer
Horvaacuteth G Kriska G Malik P amp Robertson B (2009) Polarizedlight pollution A new kind of ecological photopollutionFrontiers in Ecology and the Environment 7 317ndash325 httpsdoiorg101890080129
Horvaacuteth GMoacutera A Bernaacuteth B amp Kriska G (2011) Polarotaxis innon-bitingmidgesFemalechironomidsareattractedtohorizontallypolarizedlightPhysiology and Behavior1041010ndash1015httpsdoiorg101016jphysbeh201106022
Horvaacuteth G amp Varjuacute D (2004) Polarized light in animal vision BerlinHeidelbergSpringerhttpsdoiorg101007978-3-662-09387-0
Johnson N F amp Triplehorn C A (2004) Borror and DeLongrsquos intro-duction to the study of insects 7th ed SilverwaterNSWAustraliaCengageLearning
KelberA(1999)WhylsquofalsersquocoloursareseenbybutterfliesNature402251httpsdoiorg10103846204
KelberAThunellCampArikawaK(2001)Polarisation-dependentco-lourvisioninPapiliobutterfliesJournal of Experimental Biology2042469ndash2480
Kessler S C Tiedeken E J Simcock K LDerveau SMitchell JSoftleyShellipWrightGA(2015)Beespreferfoodscontainingne-onicotinoidpesticidesNature521(7550)74
KokkoHampSutherlandWJ(2001)Ecologicaltrapsinchangingenviron-mentsEcologicalandevolutionaryconsequencesofabehaviourallymediatedAlleeeffectEvolutionary Ecology Research3537ndash551
Kriska G Bernaacuteth B Farkas R amp Horvaacuteth G (2009) Degrees ofpolarization of reflected light eliciting polarotaxis in dragonflies(Odonata)mayflies (Ephemeroptera) and tabanid flies (Tabanidae)Journal of Insect Physiology551167ndash1173httpsdoiorg101016jjinsphys200908013
KriskaGCsabaiZBodaPMalikPampHorvaacutethG(2006)Whydored and dark-coloured cars lure aquatic insects The attraction ofwater insectstocarpaintworkexplainedbyreflection-polarizationsignalsProceedings of the Royal Society of London B2731667ndash1671httpsdoiorg101098rspb20063500
KriskaGHorvaacutethGampAndrikovics S (1998)Why domayflies laytheir eggs en masse on dry asphalt roads Water-imitating polar-izedlightreflectedfromasphaltattractsEphemeropteraJournal of Experimental Biology2012273ndash2286
KriskaGMalikPSzivaacutekIampHorvaacutethG(2008)Glassbuildingsonriver banks as lsquopolarized light trapsrsquo for mass-swarming polarot-actic caddis flies Naturwissenschaften 95 461ndash467 httpsdoiorg101007s00114-008-0345-4[accessed2012Jun8]
Labhart T (2016) Can invertebrates see the e-vector of polarizationasaseparatemodalityoflightJournal of Experimental Biology2193844ndash3856httpsdoiorg101242jeb139899
LampharHASampKocifajM(2013)LightpollutioninultravioletandvisiblespectrumEffectondifferentvisualperceptionsPLoS ONE8e56563httpsdoiorg101371journalpone0056563
vanLangeveldeFEttemaJADonnersMWallis-DeVriesMFampGroenendijk D (2011) Effect of spectral composition of artificiallightontheattractionofmothsBiological Conservation1442274ndash2281httpsdoiorg101016jbiocon201106004
Lerner A Meltser N Sapir N Erlick C Shashar N amp Broza M(2008)Reflectedpolarizationguideschironomidfemalestoovipo-sitionsitesJournal of Experimental Biology2113536ndash3543httpsdoiorg101242jeb022277
Lerner A Sapir N Erlick C Meltser N Broza M amp Shashar N(2011)Habitatavailabilitymediateschironomiddensity-dependentoviposition Oecologia 165 905ndash914 httpsdoiorg101007s00442-010-1893-9
LongcoreTAldernHLEggersJFFloresSFrancoLHirshfield-YamanishiEhellipBarrosoAM(2015)Tuningthewhitelightspec-trumoflightemittingdiodelampstoreduceattractionofnocturnalarthropodsPhilosophical Transactions of the Royal Society of London B37020140125httpsdoiorg101098rstb20140125
Longcore T amp Rich C (2004) Ecological light pollution Frontiers in Ecology and the Environment 2 191ndash198 httpsdoiorg1018901540-9295(2004)002[0191ELP]20CO2
LorneJKampSalmonM(2007)EffectsofexposuretoartificiallightingonorientationofhatchlingseaturtlesonthebeachandintheoceanEndangered Species Research 3 23ndash30 httpsdoiorg103354esr003023
Maacutelnaacutes K Polyaacutek L Prill E Hegeduumls R Kriska G Deacutevai G hellipLengyelS(2011)Bridgesasopticalbarriersandpopulationdisrup-tors for themayflyPalingenia longicauda An overlooked threat tofreshwaterbiodiversityJournal of Insect Conservation15823ndash832httpsdoiorg101007s10841-011-9380-0
MuheimR SjoumlbergSampPinzon-RodriguezA (2016)Polarized lightmodulates light-dependentmagnetic compass orientation in birdsProceedings of the National Academy of Sciences 113 1654ndash1659httpsdoiorg101073pnas1513391113
Nowinszky L (2003) The Handbook of Light Trapping (p 276)SzombathelyHungarySavariaUniversityPress
NowinszkyLKissOSzentkiraacutelyiFPuskaacutesJampLadaacutenyiM(2012)Influenceofilluminationandpolarizedmoonlightonlight-trapcatchofcaddisflies(Trichoptera)Research Journal of Biology279ndash90
PawsonSMampBaderMKF(2014)LEDlightingincreasestheecologicalimpactoflightpollutionirrespectiveofcolortemperatureEcological Applications241561ndash1568httpsdoiorg10189014-04681
Perkin E K Houmllker F amp Tockner K (2014) The effects of artificiallightingonadult aquaticand terrestrial insectsFreshwater Biology59368ndash377httpsdoiorg101111fwb12270
RadwellAJampCampNB(2009)ComparingchemiluminescentandLEDlightfortrappingwatermitesandaquaticinsectsSoutheastern Naturalist8733ndash738httpsdoiorg1016560580080414
RandASBridarolliMEDriesLampRyanMJ(1997)LightlevelsinfluencefemalechoiceinTungarafrogspredationriskassessmentCopeia1997447ndash450httpsdoiorg1023071447770
RundloumlfMAnderssonGKBommarcoRFries IHederstroumlmVHerbertssonLhellipSmithHG(2015)Seedcoatingwithaneonico-tinoidinsecticidenegativelyaffectswildbeesNature521(7550)77
RobertsonBACampbellDRDurovichCHetterich ILesJampHorvaacutethG (2017)The interfaceofecologicalnoveltyandbehav-ioralcontextintheformationofecologicaltrapsBehavioral Ecology281166ndash1175httpsdoiorg101093behecoarx081
RobertsonBAampHuttoRL(2006)Aframeworkforunderstandingecological traps and an evaluation of existing evidenceEcology871075ndash1085
RobertsonBAKeddy-Hector IAShresthaSDSilverbergLYWoolner C E Hetterich I amp Horvaacuteth G (2018) Susceptibilityto ecological traps is similar among closely related taxa but sensi-tive to spatial isolationAnimal Behaviour 135 77ndash84 httpsdoiorg101016janbehav201710023
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
eva12690
186emsp |emsp emspensp ROBERTSON aNd HORVAacuteTH
RobertsonBAOstfeldRSampKeesingF(2017)TrojanfemalesandJudas goats Evolutionary traps as tools in wildlifemanagementBioScience67983ndash994httpsdoiorg101093bioscibix116
RobertsonBARehageJSampSihA(2013)Ecologicalnoveltyandtheemergenceofevolutionary trapsTrends in Ecology amp Evolution28552ndash560httpsdoiorg101016jtree201304004
Schlaepfer M A Runge M C amp Sherman PW (2002) EcologicalandevolutionarytrapsTrends in Ecology and Evolution17474ndash480httpsdoiorg101016S0169-5347(02)02580-6
SchlaepferMA ShermanPWBlosseyBampRungeMC (2005)IntroducedspeciesasevolutionarytrapsEcology Letters8241ndash246
SchwindR (1991)Polarizationvision inwater insectsand insects liv-ing on amoist substrate Journal of Comparative Physiology A169531ndash540
Schwind R (1995) Spectral regions in which aquatic insects see re-flected polarized light Journal of Comparative Physiology A 177439ndash448
Somers-Yeates R Hodgson D McGregor P K Spalding A ampffrench-Constant R H (2013) Shedding light on moths Shorter
wavelengthsattractnoctuidsmorethangeometridsBiology Letters920130376httpsdoiorg101098rsbl20130376
Szaacutez D Horvaacuteth G Barta A Robertson B A Farkas A Egri Aacutehellip Kriska G (2015) Lamp-lit bridges as dual light-traps for thenight-swarmingmayflyEphoron virgo Interaction of polarized andunpolarized light pollution PLoS ONE 10 e0121194 httpsdoiorg101371journalpone0121194
WarrantEampDackeM(2011)Visionandvisualnavigationinnoctur-nal insectsAnnual Review of Entomology56 239ndash254 httpsdoiorg101146annurev-ento-120709-144852
How to cite this articleRobertsonBAHorvaacutethGColorpolarizationvisionmediatesthestrengthofanevolutionarytrapEvol Appl 201912175ndash186 httpsdoiorg101111
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