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ColourPreferencesofUKGardenBirdsatSupplementarySeedFeeders
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LukeRothery1,GrahamW.Scott1&LesleyJ.Morrell1*
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*Correspondingauthor
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email:[email protected](LJM)
SchoolofEnvironmentalSciences,UniversityofHull,Kingston-upon-Hull,UK
1
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Abstract
Supplementaryfeedingofgardenbirdshasbenefitsforbothbirdpopulationsand
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humanwellbeing.Birdshaveexcellentcolourvision,andshowpreferencesforfooditemsof
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particularcolours,butresearchintocolourpreferencesassociatedwithartificialfeedersis
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limitedtohummingbirds.Here,weinvestigatedthecolourpreferencesofcommonUK
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gardenbirdsforagingatseed-dispensingartificialfeederscontainingidenticalfood.We
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presentedbirdssimultaneouslywithanarrayofeightdifferentlycolouredfeeders,and
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recordedthenumberofvisitsmadetoeachcolourover37030-minuteobservationperiods
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inthewinterof2014/15.Inaddition,wesurveyedvisitorstoagardencentreandscience
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festivaltodeterminethecolourpreferencesoflikelypurchasersofseedfeeders.Ourresults
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suggestthatsilverandgreenfeederswerevisitedbyhighernumbersofindividualsof
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severalcommongardenbirdspecies,whileredandyellowfeedersreceivedfewervisits.In
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contrast,peoplepreferredred,yellow,blueandgreenfeeders.
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Introduction
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Ithasbeenestimatedthat20-30%ofpeopleinmoredevelopedareasoftheworld
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providewildbirdswithadditionalfood(supplementaryfeeding)atsomepointintheyear
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(typicallyduringthewintermonths)[1,2].IntheUK,approximately60%ofhouseholdswith
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gardensprovidefoodforbirds[3],estimatedat12.6millionhouseholds[1],7.4millionof
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whichusebirdfeeders[4].AsaresulttheUKwildbirdfeedingindustrywasestimatedas
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beingworth£210mperannum[5],andthewildbirdcaremarketrose15%invalue
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between2014and2015[6].Levelsofbirdfeedingvaryenormouslyacrosssociety[7],but
2
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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theimportanceoftheconnectionbetweenpeopleandnaturetohumanwell-beinginurban
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environmentsiswellestablished[8].Peoplefeedbirdsbecauseitgivesthemasenseof
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personalwellbeing,althoughtheunderpinningemotions,experiencesandpersonal
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perceptionsofthepeoplefeedingbirdsarecertainlymorecomplexthansuchasimplistic
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statementmightsuggest[9].Somepeople(thoseinvolvedinavianmonitoringorresearch)
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feedbirdsinordertoattractthemforcapture,measurementandsubsequentrelease.
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Duringthenorthernhemispherewinternaturalfoodresourcesareattheirlowest
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levelofavailability[10]andabird’sthermodynamiccostsareattheirhighest[11].Over
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wintersurvivalisthushighlydependentuponthecharacteristicsandavailabilityoffood
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supply[10].Gainingenoughenergyeachdaytoensureovernightsurvivalisparticularly
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importantforsmallpasserines:individualsinthetitfamily(Paridae)canloseupto10%of
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theirbodyweightovernightinwinter[12].Supplementaryfeedingmayoff-settheeffectsof
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winterresourcedepletion[13]andinmanycasesawinterfeedingstationmaybethemost
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abundantanddependablefoodsourceinaparticulararea[14].Supplementaryfeedinghas
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beenrecordedashavinganumberofotherbenefitstobirds,includinglargerclutchsizes
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(housesparrowsPasserdomesticus[15]),betterbodyconditionandmorerapidrecovery
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frominjury(CarolinachickadeeParuscarolinensis,tuftedtitmiceParusbicolorandwhite-
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breastednuthatchSittacarolinensis[16]).Supplementaryfeedingincreasesboththerange
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ofspeciesandnumberofindividualsvisitinggardens[1,17]andincreasesabundanceata
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landscapescale[1].IntheUK,forexample,supplementaryfeedinghasbeenimplicatedin
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populationincreasesofbothhousesparrowandstarling(Sturnusvulgaris[18])andmaybe
3
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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importantintheevolutionof‘new’migrationstrategiesamongstover-winteringblackcap
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(Sylviaatricapilla[19]).
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Inorderthatthebenefitsofsupplementaryfeedingtobothbirdsandthepeople
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whofeedthemarerealiseditisessentialthatfoodbeprovidedinawaythatmakesit
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accessibletobirds.Inthecaseoftheseedbasedfoodsprovidedtopasserines
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supplementaryfeedingmostofteninvolvestheuseofcommerciallyavailabletubularseed
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dispensers.Thesefeederscommonlyconsistofatransparentplastictubethroughwhich
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seedsarevisibletobirdsandcolouredmetalorplasticlids,bases,perchesandfeederports.
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Here,wereportaninvestigationintowhetherthecolourofthesemetalorplasticparts
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affectedthenumberofbirdschoosingtofeedataparticularfeeder.Forafeedertoattract
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largernumbersofbirds,somethinglikelytobeseenaspreferablebythosethatpurchase
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feeders,thecolourshouldbeattractiveorneutraltoeitheraparticulartargetspecies,or
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seedfeedingbirdsmoregenerally.Afeederthecolourofwhichbirdsavoidwouldnotbean
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effectivefeeder.
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Birdshaveexcellentcolourvisionandexhibittheabilitytodistinguishandchoose
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betweendifferentcoloursandshades[e.g.20-22].Here,wefocusoncolourpreferencesin
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relationtoforaging.Multiplestudiesreportpreferencesofbirdsforfooditemsofa
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particularcolour.Greattits(Parusmajor)bluetits(Cyanistescaeruleus)andEurasian
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nuthatches(Sittaeuropaea)allpreferreduncoloured(natural)peanutsoverthosethathad
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beendyedwhite[23].Willsonetal.[24]reviewedanumberofstudiesdemonstratingthat
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frugivorousbirdspreferblackorredgrapesorcherriesoverothercolourssuchasgreenand
4
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yellow,butpointoutthatpreferenceforcolourhereisconfoundedbypreferenceforother
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factorsassociatedwithcolour,suchasripeness,sizeandnutritionalvalue[24].Other
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studieshaveusedartificialornovelfoodsdyeddifferentcoloursandfoundcolour-based
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preferences[24-26].Wilsonetal[24]reportedapreferenceforred,andavoidanceof
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yellowinthreespeciesoffrugivorousbird,whileNorthIslandrobins(Petroicalongipes)
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preferredyellowandavoidblueandbrown[26]forexample.
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Preferencesforcolourassociatedwithsupplementaryfeeders,ratherthanfood,
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haveexclusivelyfocusedonthepreferencesofhummingbirds(Trochillidae)atfeeders
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designedtoprovidesugarsyrup.Whilehummingbird-pollinatedflowerstendtobered
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[27,28],andbirdstendtopreferred-pigmentedflowersoverthoselackingredpigments(e.g.
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[29-31],reviewedin[28]),experimentalstudiesonfeedersdonotshowaconsistent
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preferenceforanyparticularcolour(e.g.[32-34],reviewedin[28]).Instead,factorssuchas
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location[35,36],previousexperience[37-39]andnectarquality[35,39]appeartobemore
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importantindeterminingchoice.Wehavebeenunabletofindanypeer-reviewedstudiesof
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theimpactofseeddispensingfeedercolouronbirdfeedingbehaviour.Oneanecdotal
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report[40]suggestedthatworkcarriedoutbytheBritishTrustforOrnithology
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demonstratedcolour-basedpreferencesforbirdsvisitingseedandpeanutfeeders,namely
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thatblueseedfeedersarepreferredduringthesummer,whilesilverfeedersarepreferred
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inwinter(althoughgoldfinchespreferredgreen),andredpeanutfeedersarepreferredover
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othercolours.Theprimaryaimofourresearchwastoinvestigatetheeffectoffeedercolour
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onthefeedingpreferencesofwildbirds.Asanadditionalaimweinvestigatedthelevelto
5
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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whichbirdsandthehumanswhofeedthemagreedontheirpreferredfeedercolour,an
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importantconsiderationforthosewhomakeandsellfeedersandthosewhousethem.
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Methods
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Birdcolourpreference
Toexploretheeffectofcolouronthenumberofvisitsbybirds,werecordedbird
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visitratesto8differentcolouredfeedersatthreesiteson78samplingdaysduringthe
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winter/springof2014/15(November2014toMay2015).
DatawerecollectedatTophillLowNatureReserve(Driffield,EastYorkshireTA
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075,492),TheUniversityofHullBotanicGarden(Cottingham,EastYorkshireTA050,329)
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andasuburbangardeninOtley(WestYorkshireSE195,472).Thesesiteswerechosendue
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toaccessibilityandthepresenceofexistingartificialfeederswithregularavianvisitors.The
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feedersused(NaturesFeastRoyalSeedFeeders,WestlandHorticulture)wereof
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transparenttubulardesignwithmetallids,twometalportsandtwostraightmetalperches.
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ThemetalpartsofeachfeederwerepaintedasinglecolourusingHammeriteMetalPaint.
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TheproprietarycoloursusedwereSmoothBlack,SmoothBlue,SmoothDarkGreen,Smooth
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Red,SmoothWhite,SmoothYellow,HammeredSilverandPurple(achievedbymixing
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SmoothBlue,SmoothRedandSmoothWhiteataratioof3:2:1).Analysisofthefeeder
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colourscanbefoundinthesectionbelow.Throughouttheexperimentthefeederswere
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filledwith“Nature’sFeastHighenergyNoMess12SeedBlend”(WestlandHorticulture,UK).
6
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Ateachsitethefeedersweresuspendedinalinefromametalcross-bar,30cm
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apartfromoneanotherand1.5mabovetheground.Atanytime,onefeederofeachofthe
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8coloursusedwasavailable(seesupportinginformation:FigS1).Theorderofthefeeders
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alongthecross-barwaschangedafterevery30minuteobservationperiodaccordingtoa
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pre-determinedrandompatterntocontrolforanypreferencesbasedonfeederposition
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ratherthancolour.Feederswerefilledatthebeginningofeachobservationperiodand
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cleanedthoroughlyevery14days.Duringeachdatacollectionsessionthenumbersof
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feedingvisitsbybirdstoeachofthefeedersinthearraywererecordedover30minutes.A
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feedingvisitwasdefinedasabirdlandingontheperchandtakingfoodfromthefeederport.
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Birdswereidentifiedtospecieslevel,butasitwasnotpossibletodistinguishbetween
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individualsofthesamespecies,eachvisittothefeederswascountedasanindependent
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datapoint.Allobservationsperiodswerevideorecorded(SonyHandycamHDR-CX240E)
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mountedonatripodapproximately10mfromthefeeders.Identificationandcountingof
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birdseithertookplaceinreal-timeinthefieldorlaterusingthevideorecordings(wherethe
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numberofvisitswastoohightoallowforaccuratereal-timerecording).
Datawerecollectedacrossatotalof370observationperiods(Otley:208;Tophill;26
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BotanicGardens:136),andatotalof7535visitstothefeederswererecorded(table1).
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Table1:Summaryofdata,showingthetotalnumberofvisitsbyeachspeciesateachsite,
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andthenumberofsampleperiodsinwhichthatspecieswasobservedatleastonce.
Species
Thwaite
Tophill
Gardens
Low
Otley
Total
Sample
periods
7
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BluetitCyanistescaeruleus
810
1824
629
3263
108
GreattitParusmajor
833
1564
13
2410
38
HousesparrowPasser
-
-
701
701
58
CoaltitPeriparusater
311
116
109
536
48
RobinErithacusrubecula
171
12
105
288
75
StarlingSturnusvulgaris
-
-
172
172
13
GreenfinchChlorischloris
-
1
135
136
21
MarshtitPoecilepalustris
-
16
-
16
3
LongtailedtitAegithalos
2
3
-
5
BullfinchPyrrhulapyrrhula
5
-
-
5
3
GoldfinchCardueliscarduelis
-
3
-
3
1
2132
3539
1864
7535
370
domesticus
2
caudatus
Total
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143
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Humancolourpreference
Toassessthepreferencesoflikelypurchasersofbirdfeeders,wecollecteddataina
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gardencentre(HornseaGardenCentre,Sigglesthorne,Hornsea,UK)wheresimilarfeeders
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weresold(3days,82-hoursampleperiods)andattheUniversityofHullScienceFestival(1
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dayasasinglesampleperiod).Ateachvenueweexplainedtoadultvolunteersthatwe
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wereinvestigatingthechoicesmadebybirdsandpeoplebutwedidnotprovideany
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informationonactualbirdpreferences(supportinginformation:FigS2).Peoplewereshown
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thecolouredfeedersusedinthestudyandaskedsimplytoindicate(byplacingatokeninan
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appropriatelycolouredcontainer)whichtheywouldbemostlikelytobuyfortheirown
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garden.Containerswereemptiedandtokenscountedattheendofeachsampleperiod.In
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total,587‘votes’werecastduringthepoll.
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Dataanalysis
AllanalysiswascarriedoutusingRv3.2.3[41].Thetotalnumberofvisits(acrossall
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species,togiveameasureoftheoverallpreferenceforparticularcolours)tothefeeders
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wereanalysedusingageneralisedlinearmixedeffectsmodelwithaPoissonerror
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distribution(asappropriateforcountdata).Observationperiodandsitewereaddedas
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randomeffectstoaccountfornon-independenceofvisitstofeedersdisplayedatthesame
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time,andoveralldifferencesinbirdpopulationsatagivensite.Anobservation-level
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randomeffectwasincludedtoaccountforoverdispersioninthedata[42].Re-levelingthe
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datawithinthemodelallowedforallpairwisecomparisonsbetweencolourstobemade,
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andp-valueswerecorrectedformultipletestingacrosspairwisecomparisonsusingthefalse
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discoveryratecontrolmethod[43].Thesameanalysiswasusedforthenumberofvisitsfor
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eachspecieswithmorethan100totalvisitstothefeeders(seesupplementarytablesS1-S5),
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toevaluatewhetherdifferentspecieshaddifferentcolourpreferences.Preferences
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expressedbyvisitorstothegardencentreandsciencefestivalwerealsoanalysedusingthe
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samemethodology.
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Feedercolouranalysis
9
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Toobjectivelydescribethecolourofthefeeders,photographsofthefeederlids
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weretakeninRAWformatusingaCanonPowershotG12camera.Lidswereplacedintoa
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lighttent(EZCube,Ventura,CA,USA)underdaylightspectrumilluminationwithawhite
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reflectancestandard(OceanOptics,Dunedin,FL,USA).
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forImageJ1.50i[45].Afterusingthetoolboxtolineariseandstandardisetheimageagainst
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thewhitestandard,apatchoneachfeederthatwasapproximatelythesamedistanceand
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orientationasthereflectancestandardandfreefromspecularreflections,wasselected,and
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themeancamera-specificRGBvaluesofthepatchwererecorded(16-bitcolourdepth).
ImageswereprocessedusingtheImageCalibrationandAnalysisToolbox[44]plugin
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Tosummarisetheluminanceindependentcolourmeasures,RGandBYratioswere
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calculated(RG=(R-G)/(R+G);BY=B–((R+G)/2)/B+((R+G)/2);Fig1A).Theseratiosdescribe
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therednessversusgreenness(RG),andbluenessversusyellowness(BY)ofastimulusand
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approximatehumanandpotentialavianopponentcolourchannels[46].Additionally,
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luminance((R+G+B)/3)isshowninFig1B.AsthecamerawasnotUVsensitiveandhadnot
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beencharacterised(i.e.thespectralsensitivityofeachsensormeasured),itwasnot
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possibletomeasurereflectanceintheUVrangeortransformtheRGBvaluesintoavian
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colourspace[44].
10
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Fig1:Analysisoffeedercolour.(A)RGandBYratios,and(B)luminanceforthe8different
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feedercolours
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Ethicalstatement
ExperimentswereapprovedbytheUniversityofHull'sSchoolofBiological,
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BiomedicalandEnvironmentalSciencesandFacultyofScienceandEngineeringethical
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reviewcommitteesbeforecommencement.Allavianworkwasobservational,andcarried
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outatlocationswheresupplementaryfeedingofbirdsalreadyoccurredandwouldcontinue
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afterdatacollectionwascompleted.Permissiontocarryoutfieldworkwasgrantedbythe
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UniversityofHull(ThwaiteGardens),RichardHampshire(TophillLowReserveWarden)and
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MarkRothery(Otleysiteowner).Thefieldstudiesdidnotinvolveendangeredorprotected
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species.Allparticipationinthehumancolourpreferencewasentirelyvoluntaryandthe
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purposeoftheexperimentwasexplainedtotheparticipantseitherverballyorviaanA4
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posterdisplayednearthestand(FigS2).Writtenconsentwasnotobtainedtoensure
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participationwassimpleandtomaximisethenumberofparticipants,andapprovedbythe
11
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institutionalreviewboardsabove.Nodataontheparticipants(otherthantheirchoiceof
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colour)wascollected.
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Results
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Birdcolourpreference
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Therewasasignificanteffectoffeedercolouronthenumberofvisitstothefeeders
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(F7,875=6.120,p<0.001;Fig2A).Birdsmadesignificantlymorevisitstothesilverfeederand
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significantlyfewervisitstotheredandyellowfeedersthananyothercolour(allp<0.05;
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table2).Greenwasvisitedsignificantlymoreoftenthananyothercolourexceptsilver,but
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therewasnodifferenceinthenumberofvisitstoblue,purple,whiteandblack.
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Table2:Pairwisecomparisonsofvisitstofeeders.
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Pleaseseeendofdocumentfortable2(landscapeformat)
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Thecellsabovethediagonalshowthez-andp-values,whiletheestimate±standarderroris
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belowthediagonal.Significantp-valuesarehighlightedinbold.Adjustedp-valuesfollowing
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falsediscoveryratecontrolarepresented.
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Forbluetits(Fig2B,supportingtableS1),therewasasignificanteffectofcolouron
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numberofvisits(F7,749.73=4.3575,P<0.001).Yellowandredweretheleastvisitedcolours,
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andwerevisitedwithsimilarregularity(tableS1,p>0.05).Yellowwasvisitedsignificantly
12
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lessthanallothercoloursexceptwhite(p>0.050),whilevisitstoredwerenotdifferent
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fromwhiteorgreen(p>0.05).Therewerenodifferencesinthenumberofvisitsbetween
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theothercolours(p>0.05;tableS1).
Forgreattits(Fig2C,tableS2),therewasasignificanteffectofcolouronnumberof
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visits(F7,259=2.671,p=0.011).Thereweresignificantlyfewervisitstoyellowthantoall
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othercoloursexceptred(p<0.05inallcases),whileredwasvisitedsignificantlylessoften
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thangreen(p=0.017).Therewerenoothersignificantpairwisedifferences(tableS2).
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Forcoaltits,therewasasignificantoveralleffectofcolouronvisits(F7,329=3.796,p<0.001),
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butnosignificantpairwisecomparisonswerefoundaftercorrectionformultipletesting(Fig
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2D;tableS3).
Forhousesparrows,therewasasignificanteffectofcolouronvisits(F7,399=11.139,
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237
P<0.001).Theyellowfeederwasvisitedsignificantlylessoftenthanallothercolours(Fig
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2E,tableS4,p<0.05inallcases),andredwasvisitedlessoftenthanblue,green,silverand
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black(p<0.05).Whiteandpurplewerevisitedlessoftenthanblue,greenandblack(p<
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0.05)whichwerethecoloursvisitedmost(althoughnotsignificantlymorethansilver;table
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S4)
Forrobins,therewasasignificanteffectofcolouronvisits(F7,518=3.1033,p=0.003;
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Fig2F).Black,themostvisitedcolour,wasvisitedsignificantlymoreoftenthanpurpleand
244
white(p<0.05,tableS5),theleastvisitedcolours,butnootherpairwisecomparisonswere
245
significant.
13
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Therewasnosignificanteffectofcolouronvisitsforgreenfinch(F7,140=1.3.383,p=0.217)
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orstarling(F7,84=1.232,P=0.294),andnootherspecieswasrecordedmorethan100times
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duringthesampleperiod,sotheirpreferenceshavenotbeenanalysed.
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Fig2:Birdcolourpreferences.Meannumbersofvisitsperobservationperiodtofeedersof
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eachcolour,for(A)allspeciescombined,(B)BluetitCyanistescaeruleus(C)GreattitParus
14
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major(D)CoaltitPeriparusater(E)HousesparrowPasserdomesticusand(F)Robin
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Erithacusrubecula.Errorbarsrepresent+/-1S.E.
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Humancolourpreference
Therewasasignificanteffectofcolouronthepreferencesobservedinoursurvey
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(F1,7=10.485,P=<0.001;Fig3a).Pairwisecomparisonsrevealedthatred,yellow,greenand
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bluewerepreferredoverpurple,white,silverandblack(table3).Fig3Bshowsthemean
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numberofvisitsbybirdsplottedagainstthemeannumberofvotesfromvisitors,and
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suggeststhathumanandbirdpreferencesdonotnecessarilyalign.Coloursinthetopright
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ofFig3Barethosethatreceivedhighvisitratesfrombirdsandhighnumbersofvotesfrom
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visitors,andwesuggestthosecolours(greenandtoalesserextent,blue)maybe
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simultaneouslymarketableandwell-visitedbybirds.Whileredandyellowreceivedhigh
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numbersofvotesfromvisitors,thesearethecoloursthatreceivedthelowestnumbersof
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visitsfrombirds.
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Table3:Pairwisecomparisonsofnumbersofvotesforeachcolour.
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Pleaseseeendofdocumentfortable3(landscapeformat)
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Thecellsabovethediagonalshowthez-andp-values,whiletheestimate±standarderroris
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belowthediagonal.Significantp-valuesarehighlightedinbold.
15
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
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Fig3:Humancolourpreferences.(A)Meannumberoftokensplacedintothecontainer
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correspondingtoeachcolouredfeederbypotentialpurchasersofbirdfeeders.(B)The
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combinedpreferencesofpotentialpurchasers(xaxis)andvisitsbyallbirds(yaxis)foreach
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colourfeeder.Errorbarsrepresent+/-1S.E.
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Discussion
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Overall,birdspreferentiallyvisitedthesilverfeeders,followedbygreen,andmade
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fewervisitstotheredandyellowfeederswhenallfeedersweredisplayedsimultaneously.
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Thesepatternsarelikelydrivenbythepreferencesofthemostabundantspeciesatthe
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feeders(bluetitsandgreattits),whichshowedsimilarpreferencestotheoverallpatterns.
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Thesepatternscontrastwiththepreferencesexpressedbythepotentialpurchasersof
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feeders,whopreferredred,yellow,greenandblue,butrarelyvotedforsilver.Intermsof
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feederdesign,thissuggeststhatgreen(andtoalesserextent,blue)maybesimultaneously
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marketableandwellvisitedbybirds.Ourfindingsalsosuggestthatdifferentspeciesof
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birdsmayhavedifferentcolourpreferences,althoughthetotalnumberofvisitsbysome
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specieswastoolowtoevaluatethis.
16
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
288
Silverandgreenfeedersmaybepreferredoverredandyellowforavarietyof
289
290
reasons.Greenandsilverarecommoncoloursforbirdfeeders,andfamiliaritywith
291
particularcoloursmayhaveplayedaroleindeterminingpreferences.Inhummingbirds,
292
previousexperienceofparticularcoloursplaysaroleincolourchoice.Anna’s(Calypteanna)
293
andrufous(Salasphorusrufus)hummingbirdspreferentiallychooseredfeedersifcaptured
294
fromred-floweredRibesspeciousmplants,butpreferyellowifcapturednearyellow-
295
floweredNicostianaglauca[38].Hummingbirdscanalsobetrainedtopreferparticular
296
colourswhenthatcolourisassociatedwithhigherqualityrewards[31,35,39].Astheseed
297
qualityinourfeederswasidentical,thepreferencesexhibitedbythebirdscouldhavebeen
298
duetoourchoiceoflocationswherebirdswereregularlyfed,andthusfamiliarwith
299
commonlycolouredfeeders.
300
301
Incontrast,redandyellowareuncommoncoloursforseeddispensingbirdfeeders.
302
Neophobiainrelationtofoodcolourhasbeenwelldocumentedinbothbirds(e.g.[47-51])
303
andotherspecies[52,53].Redandyellowarealsoassociatedwithwarningcolourationand
304
aposematism,andmaybeavoidedbyforagingbirds[54,55].Redandyellowfeedersmay
305
alsobemoreconspicuousagainstthebackground(whilegreenandsilveraremorecryptic),
306
whichmayincreaseperceivedpredationrisk[56].However,thesecoloursmayalsomake
307
theresourcemoreconspicuousfromadistance[57]andthusbrightlycolouredfeedersmay
308
beeffectiveinattractingbirdstonewforagingsitesmorerapidly:someevidencefrom
309
Anna’shummingbirdssuggeststhatredfeedersplacedinnovellocationsinitiallyattract
310
morebirdsthanothercolours[36],butredisacommoncolourofthenectarresourcefor
17
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
311
thisspecies,andsomaynotbeapplicabletoseed-feedingbirds.Ourexperimentdoesnot
312
allowustospeculateonwhetherparticularcolourswouldbemoreorlessattractivetobirds
313
ifputoutalone.
314
Duringdatacollection,weobserved(butdidnotrecord)multipleeventswherea
315
316
competitordisplacedfeedingindividualsfromonefeedertoanother.Thismaymaskfeeding
317
preferencesasindividualsarethenrecordedatlesspreferredfeedercolours,alimitationof
318
presentingallcolourstogether.Thewaysinwhichdifferentoptionsarepresentedoften
319
affectsthechoicesthatanimalsmake.Hummingbirdsofferedachoicebetweenredand
320
yellow-floweredMimulusaurantiacusprefertofeedattheredmorph[58],butinahybrid
321
populationwhereorangemorphsoccur,visitorangeflowersmoreoftenthanexpectedby
322
chance,giventheirprevalenceinthepopulation[28].Preferencesbetweentwooptionmay
323
alsobeaffectedbytheadditionofathirdoption(e.g.ifAispreferredoverB,andBoverC,
324
thenAisnotnecessarilypreferredoverC),violatingtheprinciplethatchoicesare‘rational’
325
andpreferencesaretransitive[59,60].Evidencesuggeststhattheprincipleofrational
326
choiceisviolatedbyarangeofspecies,includinghumans(e.g.[61-63]),honeybees(Apis
327
mellifera[64]),rufoushummingbirds[59,65],starlings[66]andgreyjays(Perisoreus
328
canadensis[64]).Bypresentingallcolourstogether(andcoveringawiderangeofcolour
329
options)wewereabletoovercomesomeoftheseissuesassociatedwithanimaldecision-
330
making.
331
Furtherworkisneededtoexploreinterspecificdifferencesincolourpreference:for
332
333
thebirdfeederindustry,itmaybedesirabletodesignandmarketfeedersforparticular
18
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
334
targetspeciesorgroupsofspecies-thosethatareseenasdesirablebythepeoplethatfeed
335
birds.Furtherquestionsincludewhetherfeedercolourisimportantforattractingbirdstoa
336
newfeedingsite,increasingavianvisitornumbersatexistingfeedingsites,andwhether
337
differenttypesoffeeders,suchasthosedesignedtodispenseseeds,peanutsornyjerseeds,
338
wouldattractmorebirdsiftheyweredifferentcolours.Finally,otherfactors,suchas
339
distancetocover,orthetypeandqualityoffoodprovided,maybemoreimportantin
340
determiningthe‘success’ofaparticularfeederthanthecolour,asinhummingbirds(e.g.
341
[35-39])orthesefactors,andothers,maytradeoffwithcolourindeterminingthenumber
342
ofvisitsbybirds,astheyforageoptimally[67].
343
344
Acknowledgements
WethankRoseBullandAdamLeaforassistancewithdatacollection,WilliamAllen
345
346
forassistancewiththefeedercolouranalysis,MartinMcDaidandLorronBrightforuseful
347
discussions,andHornseaGardenCentreforallowingustocollectdataonhumanchoices.
348
RolandEnnos,SueHullandKatherineJonesprovidedusefulfeedbackonearlierversionsof
349
thismanuscript.
350
19
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
351
Supportinginformation
352
353
FigS1:Birdfeedersinthefield.Anexamplearrayoffilledbirdfeedersreadyfor
354
observationsinthefield.Thecolourorder(fromlefttoright)is:red,yellow,blue,silver,
355
green,purple,white,black.Colourorderwasrandomisedbetweentrials.
356
357
FigS2:Posterexplainingtheproject.Acopyoftheposterexplainingtheproject,as
358
displayedattheScienceFestivalandinthegardencentre.
359
360
TableS1:Pairwisecomparisonsofvisitstofeedersbybluetits.Thecellsabovethe
361
diagonalshowthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.
362
Significantp-valuesarehighlightedinbold.
363
20
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not peer-reviewed) is the author/funder. All rights reserved. No reuse allowed without permission.
364
TableS2:Pairwisecomparisonsofvisitstofeedersbygreattits.Thecellsabovethe
365
diagonalshowthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.
366
Significantp-valuesarehighlightedinbold.
367
368
TableS3:Pairwisecomparisonsofvisitstofeedersbycoaltits.Thecellsabovethediagonal
369
showthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.
370
Significantp-valuesarehighlightedinbold.
371
372
TableS4:Pairwisecomparisonsofvisitstofeedersbyhousesparrows.Thecellsabovethe
373
diagonalshowthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.
374
Significantp-valuesarehighlightedinbold.
375
376
TableS5:Pairwisecomparisonsofvisitstofeedersbyrobins.Thecellsabovethediagonal
377
showthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.
378
Significantp-valuesarehighlightedinbold.
379
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Table2:Pairwisecomparisonsofvisitstofeeders.
Red
Red
-
Yellow
z=-2.042
p=0.052
-0.114±0.056 -
Green
z=-10.765
p<0.001
Blue
z=-7.383
p<0.001
Purple
z=-7.129
p<0.001
White
z=5.793
p<0.001
Silver
z=12.751
p<0.001
Black
z=-7.614
p<0.001
Yellow
z=-9.440
p<0.001
z=-9.093
p<0.001
z=-7.779
p<0.001
z=-3.774
p<0.001
z=-5.128
p<0.001
Blue
-0.372±0.050 -0.485±0.051 0.148±0.043
z=3.413
p=0.001
-
z=-14.586
p<0.001
z=2.115
p=0.046
z=-9.569
p<0.001
Green
z=-12.650
p<0.001
-0.519±0.048 -0.633±0.050 -
z=-1.728
p=0.098
z=5.512
p<0.001
z=-0.135
p=0.892
Purple
z=-0.363
p=0.743
-0.355±0.050 -0.469±0.052 -0.164±0.043 -0.016±0.045 -
z=5.870
p<0.001
z=-0.499
p=0.666
White
0.292±0.050
z=-1.365
p=0.193
-0.406±0.052 -0.277±0.044 -0.079±0.046 -0.063±0.046 -
z=-1.863
p=0.076
Silver
0.606±0.047
-0.719±0.049 0.086±0.041
z=-7.212
p<0.001
-0.313±0.043 -
Black
-0.378±0.005 -0.491±0.051 0.142±0.043
0.234±0.042
0.250±0.043
z=3.278
p=0.001
z=5.378
p<0.001
-
554
-0.006±0.045 -0.022±0.045 -0.085±0.046 0.228±0.042
Thecellsabovethediagonalshowthez-andp-values,whiletheestimate±standarderrorisbelowthediagonal.Significantp-valuesare
555
highlightedinbold.
27
556
Table3:Pairwisecomparisonsofvotesbyvisitorstothegardencentreandsciencefestival.
Red
Red
-
Yellow
Yellow
z=-0.474
p=0.810
-0.722±1.522 -
Green
z=0.620
p=0.715
Blue
z=-0.182
p=0.0.887
Purple
z=3.285
p=0.003
White
z=-4.014
p<0.001
Silver
z=-4.234
p<0.001
Black
z=3.759
p=0.001
z=-0.657
p=0.718
z=2.810
p=0.011
z=3.540
p=0.002
z=3.759
p=0.001
z=3.285
p=0.003
0.222±1.522
z=0.146
p=0.884
-
Green
0.944±1.522
z=-2.664
p=0.015
z=-3.394
p=0.002
z=-3.613
p=0.001
z=3.139
p=0.004
Blue
z=-0.803
p=0.659
-0.278±1.522 -1.000±1.522 -1.222±1.522 -
5.000±1.522
4.278±1.522
z=-4.416
p<0.001
z=-0.949
p=0.568
White
-6.111±1.522 5.389±1.522
z=-4.197
p<0.001
z=-0.730
p=0.688
-5.167±1.522 -6.389±1.522 -1.111±1.522 -
z=3.942
p<0.001
Purple
z=-3.467
p=0.002
-4.056±1.522 -5.278±1.522 -
z=-0.255
p=0.895
Silver
-6.444±1.522 5.722±1.522
-5.500±1.522 -6.722±1.522 1.444±1.522
z=0.219
p=0.891
-
557
z=-0.474
p=0.810
Black
5.722±1.522 5.000±1.522 4.778±1.522 6.000±1.522 0.722±1.522 -0.389±1.522 -0.722±1.522 -
Thecellsabovethediagonalshowthet-andp-values,whiletheestimate±standarderrorisbelowthediagonal.Significantp-valuesare
558
highlightedinbold.
0.333±1.522
z=0.474
p=0.810
28