This is a self-archived version of an original article. This version may differ from the original in pagination and typographic details.
Author(s): Henze, Miriam J.; Lind, Olle; Mappes, Johanna; Rojas Zuluaga, Bibiana; Kelber, Almut
Title: An aposematic colour-polymorphic moth seen through the eyes of conspecifics and predators : sensitivity and colour discrimination in a tiger moth
Year: 2018
Version: Accepted version (Final draft)
Copyright: © 2018 The Authors. Functional Ecology © 2018 British Ecological Society
Rights: In Copyright
Rights url: http://rightsstatements.org/page/InC/1.0/?language=en
Please cite the original version: Henze, M. J., Lind, O., Mappes, J., Rojas Zuluaga, B., & Kelber, A. (2018). An aposematic colour- polymorphic moth seen through the eyes of conspecifics and predators : sensitivity and colour discrimination in a tiger moth. Functional Ecology, 32(7), 1797-1809. https://doi.org/10.1111/1365-2435.13100 This article isThis article by protected copyright. All rightsreserved. 10.1111/1365doi: differences to lead between the of thisversion citethisarticle and asVersion Record.Please copyediting, been throughthe typesetti This article acceptedhas been for publication andundergone fullpeer review buthasnot ID: author,Orcid *corresponding Finland 4 Sweden. University, Science,Lund Cognitive ofPhilosophy, 3 Department Australia, QLD 4072, Lucia, St ofQueensland, Institute, University Brain Queensland 2 present address: 1 Miriam J.Henze predator and colour An aposematic Editor: Dr Section: Animal Morphology Colouration and Corresponding author mail id: Article type :
Lund Vision Group, Department of Biology, Lund University, University, Lund of Biology, Group, Department Vision Lund Accepted Interactions, Biological in Centre ofExcellence Article
Christine Miller Research Article Research s
-
1 sensitivity and colour discrimination in in discrimination colour sensitivity and - ,2 2435.13100 , OlleLind - polymorphic moth moth polymorphic Functional Ecology
0000 3
, JohannaMappes [email protected]
- 0003 ng, pagination andproofreading process,may which - 3937 University of Jyväskylä, of Jyväskylä, University
- 2 808 seen
4 , Bibiana Rojas , Bibiana through
Sölvegatan 35, 22362 35,22362 Sölvegatan
Helgonavägen 3, 3, Helgonavägen a tigermoth
the eyes of the
PO Box 35, FI 40001, 40001, FI Box 35, PO 4 , AlmutKelber 22362 Lund, conspecific
Lund, Sweden Lund, 1 *
s
This article isThis article by protected copyright. All rightsreserved. Accepted ArticleAbstract headline: Rolling 7. 6. 5. 4. 3. 2. 1.
Males of the aposematic wood tiger moth ( tigermoth wood aposematic ofthe Males alone. selection bynatural be cannot explained species inaposematic polymorphism sexual species, aposematic colouration in i Although predation predators andtheir species of the system of The evolution further testing. white for preference not the or Whether white males. forces of theopposing is polymorphism colour whereas male conspecifics, T reflectivity w separate colo to According blue A. plantaginis w electroretinograms, Using colouration the question following females W males andyellow Redder females hindwings. mostoftheir range throughout colouration he he hite males are more conspicuous against green against conspicuous more are hite males hite males often have high have often hite males
costly (457 nm)
can morphs, but not orange and red females. andred females. orange butnot morphs, pheromone , and r , and
red colo red Colour discrimination in in discrimination Colour F
is also is discriminate the two male morphs two male the discriminate s , emale hindwing colo hindwing emale
but similar spectral sensitivities but similarspectral whether , andgreen
ed females are slightly more conspicuous than orange females orange than conspicuous are slightly more ed females
polymorphic aposematic aposematic polymorphic u
sexually selected. sexually u r vision models, conspecifics can discriminate discriminate can models, conspecifics r vision ration among females is likely selected bypredator is females among ration s commonly thought thought s commonly
plume males predation pressure favouring yellow males yellow favouring pressure predation
(521 nm) (521 e found
s from a distance s from er mating success than yellow males. Therefore males. than yellow er mating success discriminate u suffer least ration varies continuously from from varies continuously ration wavelength range wavelength
a tiger moth a tiger significan
is taken into consideration is taken into selection may also mayselection female female to exert animals can be better understood when be understood better can animals Arctia plantaginis Arctia . InScandinavia
from bird predatbird from in both sexes in both , tly
but search visually at at visually but search by colour by
andbrown larger whether female turn, and,in colouration
the strongest selective pressure pressure the strongestselective For moth For probably . males is based on visual cues requires requires cues onvisual males based is
retinal response retinal . M
, ) are polymorphic for hindwing hindwing for polymorphic ) are
,
with peaks inthe peaks with ales backgrounds, mostly due toUV mostly due backgrounds, influence colo influence they display s
maintained ion and bird and approach females by by females approach . , white andyellow white
and female female and bright orange . short range short
s
s
pressure
(
, at least partly, , at least in male than female than in male e Cyanistes ither wh u ration. Specifically, ration. ,
we UV preference . . This raises This .
, not by , not ask whether ask
to red. to caeruleus (349 nm) ite or yellow oryellow ite
on
the visual male
s as by a by
) for , ,
This article isThis article by protected copyright. All rightsreserved. Accepted Article Resumen 6. 5. 4. 3. 2. 1.
continua entre naranja y rojo. Machos amarillos y hembras rojas sufren menor depredación menordepredación rojassufren yhembras amarillos Machos y rojo. naranja entre continua variación presentanuna hembras alastraseras;las sus amarilla) en o (blanca polimórfica aposemática lapolilla de losmachos En Escandinavia, úni explicados ser nopueden aposemáticas especies de enlacoloración polimorfismos Los presión importante. puede ejercer una también sexual aunque selección aposemática, coloración la evolución de en la selección presión de eslaprincipal La depredación cuenta lossistemas enteniendo mejor puedeentenderse aposemáticas especies en de polimorfismos La evolución desconoce. aún se en preferencia color dicha papel del El lashembras. de parte por blancos por machos favoreci depredación la presión de balance entre un mediante mantenerse parece losmachos de polimórfica lacoloración de selección sexual; no depredación, de unafuertepresión bajo estáhembras probablemente de las rojo El color anaranjadas. que más conspicuas unpoco rojasson las hembras Asimismo, UV. reflexión de caeruleus entr perono yamarillos, blancos entre machos pueden discriminar ambossexos de individuos que (3)(521 nm); nm), yverde nm),azul (457 (349 UV enlasregiones sexos, con picos en laretina (1)mayorrespuesta encontramos: color, visión a ymodelosde Usando electrorretinogramas selecciónpor sexual. afectada estar podría hembras delas lacoloración y (2)si hembras; tipos entre losdos discriminar pueden hembras si las (1) preguntamos ello,nos Por localizarlas. para visuales pistas utilizan también distancias acortas feromonas, sus siguiendo hembras alas aproximan ambos se éx tener unmayor suelen blancos Los machos aves. por camente por selección natural. selección por camente
e hembras rojas y anaranjadas; y (4) que para las polillas, así como para un ave( para un como así laspolillas, para (4) yque rojasyanaranjadas; e hembras ), los machos blancos son más conspicuos sobre sustratos verdes y cafés por su por y cafés verdes sustratos sobre conspicuos sonmás blancos machos ), los
de los machos que de las hembras; (2) sensibilidad espectral similar en ambos similaren espectral sensibilidad lashembras;(2) de losmachos que de
visuales tanto de la especie en cuestión como de sus depredadores. desus cuestión encomo especie tanto la de visuales
de machos, y los machos entre los diferentes colores de las colores entrelosdiferentes ylosmachos de machos,
ito reproductivo que los amarillos; aunque aunque amarillos; quelos ito reproductivo endo a machos amarillos, y una preferencia preferencia yuna machosamarillos, a endo Arctia plantaginis
tienen una coloración coloración tienen una las hembras las Cyanistes Cyanistes
This article isThis article by protected copyright. All rightsreserved. 2 the doe sun Helsinki (60°), at of the areshort;latitude Europe andnorthern central 2015; al. et Gordon thefield, observations from between 18 is highest whereasmales hours, evening diurnal Theyare of theirrange. the aposematic, moth tiger The wood e systems, the perceptual receivers’ in orlimitations biases receiver exploit might morphs different ( favours another one may favour selection asnatural 2013), &Endler Rojas one ofthe in polymorphism colour Cummings ( of colourvariation maintenance a selection sexual pointed at morph polym population withinagiven colouration withvariable species aposematic in However, 1887 (Poulton organisms aposematic in colouration conspicuous Predation Introduction sensitivity spectral sexual selection, Key words: specially while searchin while specially 2
Acceptedh, Article Holarctic
and orphism) orphism) s
(Mallet &Joron
nocturnal light levels are n levels light nocturnal i
s
2013). This could be particularly relevant for species displaying species displaying relevantfor be particularly could This2013). Arctiid moths, colour polymorphism, colour vision, natural selection, predator pressure, pressure, predator selection, natural vision, colour polymorphism, moths,colour Arctiid colour commonly commonly .
Adults of Adults this principle is challenged by the difficulty bythe difficulty ischallenged principle this Nokelainen et al. 2012; Crothers &Cummi Crothers 2012; etal. Nokelainen - polymorphic h Arctia plantaginis Arctia
and 2 and advocated as the strongest selective pressure pressure strongest selective as the advocated
g for mates (Limeri & Morehouse (Limeri mates g for A. plantaginis 1999; Lindström 1999; s an alternative oradditional s analternative 2 h already already - , crepuscular, meaning that f that meaning crepuscular,
, and
O and sexually dimorphic arctiid moth moth arctiid dimorphic sexually and 'Donald & Majerus 1984; Jiggins etal. Jiggins 1984; &Majerus 'Donald
ever sexes, or both (Maan &C orboth sexes,
female calling calling female are active are
start flying during daytime start flyingduring (formerly (formerly
reached. reached.
et al.
2001; Endler & Mappes 2004). 2004). &Mappes Endler 2001; and mate and Parasemia Parasemia Rojas et al. 2015). al.2015). Rojas et
and mating mating and
selective pressure influencing the influencing selectivepressure from
plantaginis 2014). 2014). colour morph colour emales emales for predators to learn several colo tolearn predators for ngs ummings mid activity activity
2013 ;
(Conner 2009)
Ruxton release pheromone release At this time of the year, nights in year,in nights of the timeAt this - ; June
Rönkä on ). Moreover, each of theeach of ). Moreover, with a wide distribution awide with peak
2009; Nokelainen 2009; the evolution of of the evolution
until the end of July in most most ofJulyin the end until
, Sheratt while sexual selection selection sexual while 2001; 2001; sexual dichromatism, sexual dichromatism,
et al. at 20 Previous . Crothers & Crothers
Male h 2016
& Speed& ( s not set before notbefore set s own own (colour
towards thetowards ) isan flight activity activity flight
studies have studies
et al. et
2004). 2004). u
r across across 2012; 2012; This article isThis article by protected copyright. All rightsreserved. anddiscriminate backgrounds, model vision a color of theretina spectral thesensitivity determined (ERGs). We we study, In thepresent perceive role in Altogether al. 2015). et attracted (Gordon and to approach, about whichfemale(s) at range short males approaching pheromones, byafar from attractmales While females on choice stress males experience yellow successthan higher mating (e.g. communication intraspecific signal receivers andpredator aposematism of colouration Although the signal 2016) Rojas 2012; etal. (Nokelainen males adult Both obse personal Rojas, B. al.2015; Hegnaet 2011; al. et (Lindstedt are red allfemales where andGeorgia, areyellow, whereall females Scotland, except for populations, within red orange to 25:75 2015). Mappes E partsof butinlarge are monomorphic, males populations, In some hindwings. yellow, orwhite black aconspicuous have moths woodtiger Adult
Accepted Article co hindwings yellow or either white morphs with male colour two urope
( are unpalatable to bird predators, which tobird are unpalatable (Lindstedt et al (Lindstedt Nokelainen 2013 mate their own wing colo own wing their
colour colour
detection i n
In Finnish populations, f populations, In Finnish (e the vegetation or .g. Nokelainen et al. .g. Nokelainen on on to estima to . 2011; Nokelainen et Nokelainen . 2011;
and choice and . other properties other ) However . investigated the eyes of theeyes investigated
The hindwing colour of females, however, ranges continuously from from continuously ranges however, females, of colour The hindwing - prey interactions (i.e. natural selection) natural (i.e. prey interactions A. plantaginis u te how well the moths can detect conspecifics against natural against detectconspecifics can the well how moths te rs.
between (B. Rojas & J. Mappes, personal observation) personal &J.Mappes, Rojas (B.
. , these experiments d experiments , these sexual selection). E sexual Addressing this question requires that we know how tiger moths tigerhow moths know that we requires thisquestion Addressing mal
2012, 2014) 2012, es (No es
their colour morphs. morphs. their colour requencies of yellow and white morphs vary from 60:40 to to from60:40 vary whitemorphs and ofyellow requencies female related tothecolo related
, has mostly been studied within the context of the within context been studied has mostly
this raises the question this the raises al. 2012 kelainen et al.2012 et kelainen learn s
et al. may or may not mate with a malewith a not mate may or may A. plantaginis A. , wing colouration may colouration , wing - and ;
Nokelainen et al. et Nokelainen the moths’ hindwin the
xperiments xperiments 2017 - id notid reveal white pattern on the forewings andred, on thewhite pattern forewings ) and females (Lindstedt ) andfemales u r morph ,
measured measured
W using histology electroretinograms electroretinograms histology using e ;
suggest suggest G also
, whether
considering birds considering
ordon whether female .
2014 assess the g colouration as g colouration that white males have a a have males that white also also
et al. et - visually
; Rönkä etal.2018 ; Rönkä occur (Hegna, Galar (Hegna, occur wing colo wing wing ed . M have a function in afunction in have
how how 2015) ales might be choosy mightbechoosy ales reflectance
search for females search forfemales
they have et al. s base well a
u , particular as the main as the main ration plays a ration plays
a 2010 d rvation). rvation). warning warning natural natural
their
and
). ;
Brain Brain za ly
used used
if &
This article isThis article by protected copyright. All rightsreserved. microscope. with hot plate, stained ona dried a slide, onplaced 3 rinsing h. Afterrepeated glutaraldehyde 2% Three Eye the investigation. tosubjected experiments After paper towels. with until cleaned daily sp. approximately avoided systematically being andsisters between brothers crosses season and thefield every during year has been We investigated Animals Methods andmaintain moth the bluetit predator,
Accepted) Article m thickness were m thickness leaves leaves 9th histology .
male and and male Our results allow predictions on the opposing selective forces that act on the colo act on that forces selective the opposing on allowpredictions results Our , and thosefor , and .
The frequency of The kept at the University of Jyväskylä, Finland ofJyväskylä, kept attheUniversity
ad libitum ad libitum
Rhabdom l Rhabdom colour M 50:50.
oths used for oths for used three animals animals , the
2% formaldehyde 2% (for more details more (for
polymorphism within population within polymorphism
Cyanistes caeruleus taken
Larvae were raised under greenhouse conditions greenhouse under wereLarvae raised
female female reflectance measurements measurements reflectance larvae pupated larvae engt from the moths
in buffer, thebuffer, in th
Fin with a Reichert Ultracut microtome using glass knives. The sections were The sections glass knives. using microtome Ultracut Reichert with a e hs were measured hs weremeasured
A. plantaginis same day
electrophysiology a nish populations. Frequencies of yellow and white morphs andwhite yellow of Frequencies populations. nish stock of stock
had eclosed, straightened their wings and hardened, they were andhardened, wings their straightened eclosed, had . see
, can detect can , and Pupae were kept at 24 °C in transparent plastic boxes lined lined plastic boxes transparent °C at24 in kept were Pupae
head or wood tiger moth woodtiger Lindstedt et al. et Lindstedt
transferred to a dark chamber and stored at andstored chamber a dark to transferred 2% sucrosein 2% were s wereembedd from micrographs using Image J150 (NIH, Bethesda, (NIH, Bethesda, J150 Image using from micrographs
decapitated, and their heads andtheir decapitated, originated originated from from
the different colour m colour the different toluidine blue toluidine , since 2013 , since s the .
20 0.15 M sodium cacodylate buffer cacodylate sodium M 0.15 s 11
ed in epoxy resin. ed inepoxy from the2nd 10 from south th ).
R generation , with wild individuals being added being added individuals wild , with
earing containers werecontainers earing ,
and photographed under a light under alight photographed and
ern and central Finland Finland central and ern and orphs of the orphs of ,
moths
of the laboratory stock thelaboratory of fed dandelion ( fed dandelion dissected andfixated dissected Vertical Vertical for anatomy anatomy for
woodtiger sections of sections 8°C checked and checked u is kept tois kept r morphs morphs r Taraxacum Taraxacum
for later for later for 1 for from from that 2 – .
in in
24 This article isThis article by protected copyright. All rightsreserved. USA). Natick, MA, ERG analyses ERG repeatedly recording experiments. started, M Supplementary fibre peripheral the through co protocol with thewe repeated recording tothe ERG, receptors of shortwavelength contribution theTo isolate recorde animals most for data sets 10ormore with individual, foreach 3 to20times were investigated withincreasingintensity of white light flashes response a voltage wedetermined series, spectral each following as desc flux equal photon S (see times up tofour was measured sensitivity thespectral for30minutes, adaptation dark separat duration of40ms to flashes responses maximize to was adjusted anddistance position whoseangular ofalightguide, thecentral fibre via was illuminated The eye tungsten recording in alight placed For electrophysiology, (ERG Electroretinogram membrane basement USA) Maryland, Accepted Article upplementary s
were analysed bycustom analysed were d at different times of the day at times d different and operated to produce increasing increasing produce operated to
In between sets of experiments and in the end inthe and ofexperiments betweensets In
M - , following the course of the rhabdom from the base of the crystalline cone to the the to of thecrystallinecone base the the from of rhabdom thecourse , following tight, dark Faraday cage. cage. Faraday tight, dark ntinuous adaptation light adaptation light ntinuous ethods) ethods
electrode either in the ventral ( eitherinthe electrode The r . spectral spectral
a male ( a male
in Supporting Information in . Each LED was switched on 5 min onswitched 5 was LED . Each esponse amplitude esponse ribed previously (Te previously ribed ) s of the light guide in the order red, amber red, intheorder s ofthelight guide
recordings control n -
made ( or female = 8) series series .
Matlab Matlab
To explore regional differences in the retina, we inserted a the retina, weinserted in differences To regional explore from from thedark intensities V lles et al. et lles I scripts (R2013b or R2015b, The MathWorks, 160 The MathWorks, orR2015b, (R2013b scripts
. Response characteristics ofthedark characteristics Response . was
one of four light emitting diodes diodes emitting light oneof four n
= 19) = 19) n
calculated ) presenting narrow ) presenting = 9) or dorsal ( ordorsal = 9)
at the position of theeye positionat the 2014; J 2014; a - dult moth was moth dult adapted retina. retina. adapted , utes we monitored the state of the animal by of theanimal the state we monitored akobsson et al. akobsson as - value the absolute
intensity ( intensity before a series of recordings of recordings aseries before n
= 5) half of one compound eye. eye. compound of one 5)half = , green -
band spectral flashes of flashes spectral band mounted on a holder and and holderon a mounted ed by pauses of 5 s. After of5 pauses ed by
V
- , 2017). Preceding and and Preceding 2017).
log and blue and
in subsequent in subsequent (LED I ) relationship using relationshipusing )
of - adapted the potential the potential
s (see )
presen
retina retina ted ted This article isThis article by protected copyright. All rightsreserved. w UK) Congleton, (Labsphere, standard white reflectance X spectrometer Optics USB4000 anusing Ocean weremeasured asred classified females and10 Spectral reflectance measurements Reflectance curves selected type,we receptor specific non ( maxima sensitivity described in spectral series, Naka the Basedon 2017). of day dependence design ( nested witha effects models thee half thata elicited For thedark Rushton function ofresponses, level saturation rangeand thedynamic andtodocument the recordings change
Accepted see details (forfurther enon lamp Article - linear least squares approach ( approach squares least linear ffect of
in hours after midnight. midnight. after hours in
from the baseline at stimulus ons stimulus at the baseline from , w ,
- Telles et al. Telles etal. sex adapted retina, we calculated the asymptotic maximal response response maximal asymptotic thewe calculated adapted retina, e we converted we converted
and fitted - was fitted was maximal response, and the slope of theNaka the slope and response, maximal - λ Rushton function fitted to fitted function Rushton
s measurement
α ( ) of receptor types contributing to the averaged sensitivity curves sensitivity the to averaged types contributing receptor ) of
linear linear ) ( of 16 white and 10 yellow males, as well as 8 females classified as orange orange as classified females aswell8 yellow10 males, whiteand16 of 2014 to each regression regression response
All statistical models were implemented in R v3.3.3. (R Core Team Core (R in Rv3.3.3. implemented were models All statistical ) . Template
number see S see Lindstedt et al. 2011; Nokelainen et al.2012 Nokelainen etLindstedt al.2011; V measurement number measurement - amplitudes for spectral flashes forspectral amplitudes
( log log models models upplementary repetitions et to maximal hyperpolarization tomaximal et s with minimal
I (Govardovskii
dataset based on anon dataset based the to the data, in the range of 300 to 700 nm, with 1 nm resolution, nm resolution, nm, with1 700 to 300 in theof range (Dunedin, FL, USA) FL,USA) (Dunedin, V )
on these three parameters using linear mixed linear parameters using these three on -
log log M
nested within nested within contribution ethods I
as used for calibration for as used
et al. datasets obtained before and after each andafter each before datasets obtained with
) 2000) - . To get the best estimate for for get estimate the best . To the effect of the effect Rushton function at at function Rushton
- s linear linear connected to
individual into into from were used to estimate theestimate to were used . To assess the stability of of assess theTo stability normalized normalized least other receptors. V sex ). ). max - squares solution squares ). A Spectralon TM TM Spectralon ). A .
To test for To
adjusted forthe adjusted , the intensity a PX K a Naka sensitivities
. We tested . Wetested - ( n 2 pulsed 2 pulsed
= 4) time time
- λ α by a by
K . of a -
time time as This article isThis article by protected copyright. All rightsreserved. Acceptedhold conclusions photoreceptors moth noise in 2017 & Kristensen Kelber, that channel green receptor retina where a Weberfraction limitedby is mechanism Each receptor that thecontrast assume alog Article mechanisms. retinal opponent via neuronal levels intohigher thatpropagates receptor noise Osorio, 1998). and (Vorobyev areceptor using contrast visual We estimated spectrum,and illumination where catch The quantum calculations Model each ommatidium has 1 UV, 1 blue and 7 green receptors blue and7green 1 1 UV, has ommatidium each ) , . As r . As
e we
is the noise in each single receptor and single receptor each noise in is the
is wavelength, is wavelength, set receptor set eceptor noise and spatial summation and noise eceptor - linear relationship between receptor quantum catch and receptor signals, signals, receptor catchand quantum receptor between linear relationship
even for a flat receptor abundance ratio of1:1:1. ratio abundance flatreceptor a even for
f
between two stimuli two between
Q
i
2003
of photoreceptor photoreceptor of the
R to 0.1, to
is receptor sensitivity, is receptor
proportion
, Briscoe 2008 k
is a scaling factor given by adaptation to the background to the byadaptation given factor is ascaling
, we
the In
this model, we model, this
level level focus
to
measured measured
i on relative chromatic contrast and contrast relativechromatic on s
1:1: ; 1
(
Jakobsson et al. 2017, al. 2017, et Jakobsson i and
= 7
-
noise limited noise limited
UV, blue, greenUV, receptor (UV s
2 s
are , is
suppose suppose
is thereflectance spectrum is the is the in honeybee :blue:
un known proportion green receptor)
ω that colour discrimination discrimination that colour
given by receptor noise: bygiven receptor (RNL) (RNL)
s for , as in various other as in , ( Vorobyev but see
A. plantaginis A.
model of colour discrimination discrimination ofcolour model of of each ) isgiven by: .
Beluši Given the Given
et al. et receptor also test whether also test
of thespecimen of , w ç
, Šporar, & Megliç , Šporar, 2001 e set noise e Eq 2 Eq 1 Eq 4 Eq 3
moth uncertainty
channel limits ) .
spectrum . , , . f Assuming Assuming
s = ln(
( Warrant are set by are set
in the in the
W in the in the , Q our I
i e
of of ), is the is the
s so so b
: ,
This article isThis article by protected copyright. All rightsreserved. (Eq. 5): limited ( thecoefficients where axes: to the following diagram according of colourloci position the We determine S 2005) the illumination assuming when not change brown and to green corresponding spectra background twousing We M ( at contrast (just noticeable jnds is contrast colour The unitfor contrast Colour Cyanistes caeruleus 3
ethods )
Accepted . Article
calculated ,
daylight daylight
model s model
).
detection threshold
colour contrast colour contrast d uch
S
75, or light spectra typical for sunset or twilight (Johnsen et al., 2006) al., 2006) et twilight (Johnsen or forsunset spectra typical 75, orlight )
, weused
is given by: is given
that the Euclidean distance of1between Euclidean distance that the
A
,
B
themodel for
,
a
, daylight under standard
. For estimating the contrast in thecontrast . Forestimating
b
) scale the chromaticity diagram according to the receptor according diagram chromaticity the ) scale
(
tetrachromatic tetrachromatic position of position
in
a deciduous forest a deciduous
differences)
colours in the colour space) colour inthe colours
illumination illumination
bird
two
the visual system of system the visual vision (
, and
colour locicorrespond colour (Håstad
vegetation
(d65
1 jnd is defined asthe is 1 jnd
given in
;
Wyszecki and Stiles, 2000) Stiles,2000) and Wyszecki
, Victorsson
.
Eq 6 O S upplementary upplementary ur conclusions do do conclusions ur
Eq 9, Eq 9, Eq 8, Eq 7, in a chromaticity achromaticity in the ,
Eq 5.
& Ödeen tit blue instead
s
to 1 jnd 1jnd to
- noise noise colour
( F
ig.
This article isThis article by protected copyright. All rightsreserved. difference of linearmixed a on are based twiceas large dark 2 in striking differences We observed neurons. downstream lamina retina and 2A) eyes of thecompound ERGs from E plantaginis is thelamina, opticneuropil, that thesecond eyes. asapposition likely function theeyes could bedetected, in rhabdoms of ≈18 cornea plantaginis ofthe sections vertical We inspected anatomy Eye Results contrast Achromatic ; lectroretinograms (E lectroretinograms Acceptedd Article , indicating that we recorded photoreceptor responses only. responses photoreceptor that we recorded , indicating ata available from the Dryad Digital Repository: https://doi.org/10.5061/dryad. Repository: Digital the Dryad from ata available - adapted retina, t adapted
(Fig light The microscope. light under the
females 12 mV was highly significant ( significant mV washighly 12 in males (mean ± standard error: 18.6 ± 1.0 mV) as in females (6.5 ± 1.1 mV; values 1.1 (6.5± females 1.0 mV)as in error: 18.6± ±standard (mean in males . 1
). m thickness, a crystalline cone of about twice this length, and and twice length, about this cone of acrystalline m thickness,
(
see see
wa he (
n Fig
s calculated as Michelson contrast (see Supplementary (seeSupplementary contrast as Michelson s calculated =
estimated estimated
RGs
. 1
10
-
) effects model accounting for repeated measurements per measurements repeated for accounting effects model
)
)
is probably the reason why our electrode did not pick up signals of notupsignals of did pick electrode our why the reason is probably , but100
maximal maximal
A
response
compound compound .
±11 plantaginis
t response response (12) m long rhabdoms inmales rhabdoms m long
spatially strength between males betweenstrength males
- = 8.86, = 8.86, adapted eyes adapted
consisted of monophasic hyperpolari ofmonophasic consisted eyes and optic of lobes eyes and V not
p max
<
direct
to flashes of white light toof white light flashes
0.001, Table1). 0.001, (Fig. The sections of the heads oftheheads The sections ly
The rather long distance between distance long The rather
adjacent 1 ) are close to spherical, to close spherical, are ( n
(
= male and female female and male n
= 8) theretinain to K
10 , the intensity that elicits that elicits , the intensity and females ( females and 76±8 76±8 M ) .
ethods). As no clear zone noclearzone As Eq 10. was more than than was more
s46t627 individual z m long m long ations also
Arctia Arctia A. n have a have
= revealed revealed )
. (Fig.
). In the 6;
This Fig. This article isThis article by protected copyright. All rightsreserved. rhabdoms. UV peak sensitivity wavelengths toshifted longer receptor is three receptors 349 receptor, and deviation, (mean ±standard to leading a type foreach receptor all results therefore averaged We accuracy. and dorsal and females, males and sensitivity maximal the wavelengthsof in Differences thefit. improve not considerably htt example peak with pigments visual types expressing of re The log mV/ 0.02 oneffect significant hours ofday the time or of measurements of thenumber was independent andS1). This model log mV / difference 3.1 However, to the response a half
Acceptedps://doi.org/10.5061/dryad. Article
and spectral sensitivity curves sensitivitycurves spectral d, yellow, yellow, d, - , maximal response ( response maximal between
sex s, see the blue re the blue
the
had Fig. 3 slope of the curve at of thecurve slope I
green, or blue light green,or blue , a robust effect on a robust effect 10 - are not of the green receptor of thegreen intensity ( intensity t (130)
± 2 nm ( ± 2 h ceptor type, indicating a type,indicating ceptor
;
and K = d , but the , but ata available from the Dryad Digital Repository: Repository: Digital the Dryad from ata available
- equally equally
2.10, I 2 , V
V t h (12) n max -
( = 13) for the UV receptor. UV = 13)forthe n log the nextthe day p
= 7.69, = 7.69, /2) and marks the turning point of the Naka the pointthe of andturning marks /2)
obtained = 14)forthere green spaced across theacross spectrum spaced p = 0.04, Table1). = 0.04,
- s46t627 I value was high and the size of the effect minimal (mean difference difference (mean minimal sizeofthe effect the highand was value ) relationship, did not differ significantly between the sexes differsignificantly not did ) relationship, V
were ventral eye regions were within the limits of experimental experimental thelimitsof were within eye regions ventral K max
was significantly higher for males than females (mean thanfemales males for higher significantly was type p
andon the
all 2 andFig. Table1 < 0.001, under ). ( )
i. e., i. , for which we have recordings. recordings. we have which , for Adding another Adding was 11 to was 11 in agreement with the assumption of three photoreceptor photoreceptor of three assumptionwith thein agreement
sensitivities sensitivities larger
dark adaptation adaptation dark red
- number of green receptors, ofgreen receptors, number S shifted) shifted) lope
ceptor, 456 ± 7 nm ( ± 7 456 ceptor, 12 times as high as the sensitivity peak of the the of peak as thesensitivity high times12 as
in the at Note that t Note that K
. spectral spectral
by nm. 21 , but no , but Instead, UV, blue UV, and and
). adaptation to different intensities intensities toadaptation different receptor t on t he he the sensitivity peak of the blue of theblue peakthe sensitivity
Regardless of the of Regardless
For thedark peak sensitivities ,
K andgreen n
itself (compare Table (compare itself Measurement number
- = 14) for theblue = 14)for Rushton function fitted fitted function Rushton type type λ likely α - value of 521± 2 nm of521± value in - , range
adapted eye, the the adapted eye,
at least the also with statistical
models models λ (
λ α ( for for
α between ) for the 16 for
of the .
bigger had a had did did s
1
- This article isThis article by protected copyright. All rightsreserved. morph between hindwing from (Figvariable the case for This the not is clusters contrast morphs theRNLmodel by described space thecolour We areas coloured for contrasts contrast only the values, input contr of chromatic measure discrimina colour model of wingpatches white andcoloured theWe used https:/ Repository: Digital the Dryad available from t very similar orred. aseitherorange andfemales yellowmorph, or the the to white belonging on w the studies species, in previous As colouration Wing Accepted Article reflect ofwhite males hindwings he first address the discrimination between wing colours of wingcolours between the discrimination first address give a rough estimate arough give
the s (T .
, which by colour by between colours with between colours
oblique able able reflectance
6 receptor receptor black wing regions, assuming regions,assuming black wing B
are mostly
2 ) is unaffected by this is unaffected ), and - t differ vertical distribution o vertical distribution he ,
colour
if the
noise level in each photoreceptor channel photoreceptor levelineach noise sensitivities
ing -
curves
with the exception of two outliers oftwo outliers theexception with average contrast between contrast average s
detected by detected
most strongly strongly most of of white and yellow yellow of white and tion (Vorobyev & Osorio, 1998). &Osorio, tion (Vorobyev forewing forewing ast
absolute discrimination thresholds. Instead, we Instead, thresholds. discrimination absolute
in from while achromatic (orluminance achromatic while each
of determined by ERGs by ERGs determined
ed A. the uncertainty colours ( colours
morph the achromatic channel. channel. achromatic the
f colour loci in colour space (F space incolour loci f colour more UV than UV more plantaginis e classified males after inspection by the human eye as eye thehuman by after inspection males classified e forewings in thein that , s T . uch stimulation ofthe stimulation male W able able the
(
Table S2, Fig. S2, Table in daylight illumination (Fig. 5C) (Fig. illumination in daylight e assume that e assume
of
the that morph that contrasts between black and blackbetween and contrasts s 2 /doi.org/10.5061/dryad. s46t627 /doi.org/10.5061/dryad.
, all morphs all is higher than the contrast within each withineach the contrast than is higher
F those of those colours colours ig. (Fig.
6 -
A).
the Th different different 5A) ,
of different of different
)
colours any other morph any other
is S is the moths can discriminate discriminate can the moths Hindwing colouration is more is colouration Hindwing , contrast hindwing
2
hindwings differed hindwings UV to estimate the estimate to model provides a quantitative a provides model un ) .
We known for - ig. receptor, which is which receptor, morphs, and morphs, and
form do
is ignored. is ignored. 6
B). colour of the white morph morph theof white colour
morphs exceeds the morphs exceeds not
non focus focus The
predict A. either - (Fig. colour colour While overlapping
average with plantaginis on relative colour on relative plot all colours in plot allcolours Because .
In particular, ) 4 .
white
colour ;
we measured we measured the
contrast of contrast evident data contrast contrast
distinct distinct RNL
, of these of these one of one or
we can we
This article isThis article by protected copyright. All rightsreserved. contrast. achromatic than rather chromatic in h of sensitivities F illumination ifweassu hold trueeven (F backgrounds brown between ( stellatarum, w detect conspecifics, whether To understand achromati and backgrounds, brown 5C) Next, we morphs. two tobelonging as visually them categorize to be not theyable will individuals, andred orange certain discriminate to will beable conspecifics form we do that cluster forms adistinct white symbols in white symbols indwing inally, weask Fig Accepted and background, brown than the the against green higher is hindwings thewhite all except Article . A. plantaginis plantaginis A. For
distinct . ,
7 based on human vision. onhuman based ). A
these two these
the moths themselves the
investigate colour c contrasts are higher than 0.1 higher than are c contrasts a moth with a
chromatic contrasts are contrasts chromatic see Fig.5B) see spectra
clusters, but largely overlap clusters, butlargely ed s are
whether likely discriminate F
visual systems, neither in the in neither systems, visual
ig. d (Fig.
more conspicuous than forewing than conspicuous more e compared colour discrimination discrimination colour e compared
thecolour 7
ig. in colour space, sep space, in colour equally spaced equally , can detect specific morphs detect specific can , ). the red the me different relative numbers of receptor types (Fig. S ofreceptor numbers different relative me
S conspicuous more are hindwings
7 Assuming the same noise levels, w noise thesamelevels, Assuming the blue tit,the blue 3 ), nor in the discrimination between wing patchesbetween ( wing thediscrimination norin ), ).
,
- T shift of the blue receptor of of thebluereceptor shift wing colours have very similar havevery wing colours contrasts ofthewing contrasts he orange and red hindwing colours colours andred hindwing orange he s also
white and yellow males as two different different astwo males andyellow white
a predator receptor sensitivities receptor
high (Fig.S1 high and thus easily discriminable thuseasily and
( Table 2,
arated from all othermorph from arated
discrimination of wing colours again ofwing discrimination
with Fig
better )
, s tetrachromatic colour vision tetrachromatic
6
but birds are generally thought to use thought touse generally are but birds colour against against B of ) and build a continuum a build ) and
than others than
A. plantaginis than forewi ( Fig. 5A, data from data from Fig. 5A, e found no significant differences nosignificant e found A. plantaginis A. s on both backgrounds both s on chromatic chromatic different natural backgrounds different
(Fig. S1) of females . For this bird, bird, . Forthis ngs s
with that of of thatwith (Fig. 6 (Fig.
enhances enhances contrasts against greenand against contrasts
( red symbols in in symbols red colour Table S3 Table .
2 B) Telles etal. , in contrast, , in ) or differ or ) . .
Thus While (for receptor (for receptor its ability to its ability to morphs, just as as morphs, just Macroglossum Macroglossum
st green or st green colour contrast colour contrast ( blue symbols blue symbols ). ,
All results All results we assume we ent
Fig 2014)
do (Fig. (Fig. all .
7).
not
All This article isThis article by protected copyright. All rightsreserved. Accepted intensity thesame centredon is andfemale eyes of male range linear working analy Our understand to required are studies (Fig.2).Further females than larger inmales 100% than more retina were sensitive slightly comes asasurprise moths ( eyes superposition typical for thephotoreceptors Woodtiger sensitivity and anatomy Eye below. findings of our the implications discuss females.We thanorange conspicuous slightly more females are whereasred backgrounds, andbrown green against conspicuous more are white males cons eyes both of in the Furthermore, females. red from orange distinguish Articleto unable are morphs, they two colour themale discriminate sexescan both while but, sensitivities polymorphism. conspecifics, visually whethermoths can the we investigated present study females. by be to preferred seems other morph the while predators favoured by be to appears T tospecies coexist. anof aposematic morphs multiple allowing thus predation, counterbalancing of pressure capable a suggestedas been has selection prey. Sexual aposematic of avoidance thesubsequent and learning asthis aposematic species, ofsome in thecolouration theintra explain to fallsshort thisapproach However, colouration. aposematic favouring pressure selective representative themost as predation have invoked approaches Classical Discussion
(Warrant et al.2003) et (Warrant longer rhabdoms than females, females, than rhabdoms longer .
However, this However, ses suggest that the eyes of both sexes are optimized for for optimized sexesare thatboth suggest of the eyes ses
the moths
a precondition a
physiological basis of the basis physiological Our results indicate that male and female wood tiger moths have similar spectral similarspectral mothshave tiger wood female malethat and indicate results Our
appear to have functional apposition compound eyes compound apposition havefunctional to appear , are in are in his could be the case in the polymorphic wood tiger morph, where one morph onemorph where morph, tiger wood thepolymorphic be thecasein his could although can only partly explain why the signals recorded from the dark fromthe the signalsrecorded why explain partly can only direct contact including for
similar cases similar cases Land
sexual selection sexual selection &
Arctiidae ( Arctiidae
Nilsson 2002) Nilsson which with observed sex observed have been reported (Warrant et al.2003 et (Warrant been reported have the cr
should make should to to D. - y play a role in the maintenance of this of maintenance roleinthe play a E. stalline cones, without the clear zone thattheclear zone is without cones, stalline . As superposition . As variation is expected to hinder predator tohinder expected is variation - Nilsson, pers Nilsson, specific differences specific
their photoreceptors their photoreceptors discriminate thecolo discriminate
similar onal
eyes
(Fig. 1) comm
light intensities light .
pecifics and predators, andpredators, pecifics
are the general rule for general rulefor are the K - unication
population variation variation population , ( ≈ Table 1,Fig.2 Table as 20% ur morphsof ur )
. Males have Males . the rhabdoms of the rhabdoms -
more adapted ), this result ), thisresult , since In the )
t . he
This article isThis article by protected copyright. All rightsreserved. male See Wing receptor differences 1982) (e colour stimuli general of discrimination optimal for allow sensitivities photoreceptor is the bluereceptor of sensitivity ( moths in sphingid W ERG. tothe substantially sensitivity haveaeither spectral present, missed that we cannot exclude &Meinecke Hamann in been described have photoreceptors light O control flight For dense vegetation. amate,and fly to instead Males find reproduction. or for survival emit pheromones h a which carry Higher sensitivity increasecontrast might differences,which response intolarger differences translate However, ur results show sex differences in the response strength of the photoreceptors to flashes of white flashesto photoreceptors ofthe strength the response in sexdifferences results show ur Acceptedhile the Article n s but but with the eyes of thewith .
have clearly clearly have colouration in the eyes of of eyes the in colouration However, However, (
absolute andcontrast absolute in no
spectral spectral the slope of the curves is of thecurves slopethe
A. plantaginis A. in discriminability ofcolours, discriminability in t in
eavy , with their our models comparing comparing our models
until they mate, lay eggs and die. Their flight activity is limited and not important not important andis limited flightactivity Their eggs anddie. mate, lay they until distinct distinct sensitivity of the UV and green receptors receptors green and oftheUV sensitivity Manduca sexta Manduca
weight of developed eggs when they eclose, mostly sit mostly eclose, eggs whenthey of developed weight spectral tuning spectral
sensitivit A. plantaginis 1979; 1979; many many
(
Fig. 7 Fig. hindwing colouration hindwing Warrant et al. 2003; 2003; et al. Warrant butterflies (Arikawa & Stavenga 2014) & Stavenga (Arikawa butterflies ) y peak
sensitivity ) additional additional .
: red
White et al.2003 White et , forewings do not differ between morphs, but betweenmorphs, notdiffer do , forewings conspecifics
and steeper in males steeper inmales (Fig. 3). We (Fig. - s shifted
for
for for A. plantaginis close close
and thus and of male eyes of male receptor types receptor UV, blue UV, detecting
( Fig. to those to
Belušiç ( have found evidence for evidence found have
Table 2, Table 5
no obvious obvious no ; , A
Macroglossum Macroglossum and greenlight and than females, females, than )
. It is normally assumed thatequally assumed isnormally . It females with
fit c
et al. haracterized in s
Fig well M. A. plantaginis of
, .
2017
stellatarum
males 7 effect to the to A. plantaginis A. ) . Mostly . Mostly
stellatarum have have ; indicat
. More than three More than . Jakobsson et al. 2017 al. etJakobsson ,
need sensitive need sensitive yet of of here or do not contribute not here ordo natural natural to , but these receptors, if , butthese the revealed no significant significant no revealed rarely in moth in rarely
ing because
three three detect the detect : o
red Telles et al.201 Telles et
n the vegetation and the vegetation n
is similar to those found those found to is similar that intensity that intensity behaviour - spectral spectral shift of theblue shift
white and yellow and white of theof stronger eyes. m
receptor
visually s . . Females, . Females, g. Barlow g. Barlow types of of types (Langer
). - spaced spaced We 4 ), the
.
type in ,
s
This article isThis article by protected copyright. All rightsreserved. Mappes andevenattra 2001), can avoidance learning. contribute to to unlikely is andbirds, to females both themconspicuous making though white males, of hindwings moths. yellow when attacking birds of hesitation malesyellow aposematic function perhaps depends on al.2012) et (Nokelainen a greenbackground on presented ( signals a bird,white To intraspecific vs Aposematic and conspecifics, to appearance their Obviously backgrounds thehigh that tonote interesting It is mate (T that results alsoshow s implications for a b of white bark spectrum background against UV blue symbols in blue symbols
Accepted Articleable - reflection of their hindwings, white males have a higher chromatic contrast contrast higher chromatic a males have white of theirhindwings, reflection
selection. selection. 2 green andbrown green
, Fig 2004 the black ,
our anthropomorphic the the . , which is more effective than that of white males (Rojas et al. et males(Rojas thatofwhite effectivethan more which is , ,
;
7
and to the hind andto Olofsson )
. local and yellow yellow and irch tree, but a lower contrast alower tree,but irch Fig w Therefore, exual selection in the species inthespecies exual selection - and . 7 of bird . - orange andred females orange ). yellow pattern, and pattern, and yellow Thus, y Thus,
the hindwings the
et al. et B ct birds rather than birds ct backgrounds. backgrounds. community lue males
wi tits 2010 e ellow males hav ellow males
ng colours of other morphs ofother ng colours suggest that suggest
hesitate much hesitate UV view and view and of of ). Nevertheless, our results results our Nevertheless, contrast of thewhite hindwings of contrast A. plantaginis
can be can is not an efficient warning signal on its own (Lyytinen onitsown signal warning efficient an is not . (Nokelainen et al. 2014) al. et (Nokelainen
Alternatively, the recently observed strong strong observed recently Alternatively, the Nokelainen et al. (2012) al. et Nokelainen not naming of colour morphs does not does not morphs of colour naming
female misleading when searching for communication signals. signals. forcommunication when searching misleading stop them from attacking (Lyytinen attacking from stop them
the cannot be discriminated discriminated be cannot e a higher chromatic contrast chromatic higher e a
(Nokelainen et al. 2012; Gordon et al. 2015) al. etGordon al.2012; et (Nokelainen on green or brown orbrown on green longer before attacking yellow than yellow before attacking longer By
contrast against the background against the contrast have have
hind
contrast a similar wing colouration wing colouration ,
but
mostly results from high UV reflection. reflection. UV fromhigh results mostly , the strong UV reflection ofthe reflection the UV strong . We suggest that . Wesuggest support support
the predation on both mor both on predation the showed
contrast against a against contrast
vegetation
against the as distinct colour morphs as distinct colour
theopposite is of little importance for for importance is oflittle
role of role
2017 than white males than green and brown brown and green . This could have could This
always always colour colour ), than yellow males males yellow than , Lindström
natural
may explain the explain may pyrazine odour of odour pyrazine determines the determines green green
white male for conform with with conform per se
background background a flat white a flatwhite selection in selection p .
hs
, Our
et al. et & and and on s
the
This article isThis article by protected copyright. All rightsreserved. req as this would O Tri of known case 2010) Rutowski also are These males white population studied andmate work choice morphs males taxa. ways across indifferent signals manifests of visual the evolution in selection sexual and of natural The interplay the readily avoid with forewings (Zylinski much faster former the to avoid patterns contrast the on Forewings, paler depends the pigmentation red et al.2011 (Lindstedt females than orange seems to continuum O role in colour of hindwing theevolution ur Accepted and pposing sexual Article ,
model calculations too (lessred)
butterflies ( butterflies and
(Finkbeiner . combining black and between white on larval diet larval on
represent represent
deterred predators deterred
by
conspecific wing pigmentation (Lindstedt et(Lindstedt wing pigmentation red or ,
a situation uire forewing pattern forewing B Pieris rapae nidadian guppies (Endler 1983 guppies (Endler nidadian other hand, rightly coloured models of models coloured rightly a of this Heliconius species species Heliconius thisof
, Briscoe , yellow or red or yellow
selecti
mos exactly equal strength of both selective pressures both selective of equalstrength exactly warn natural selection alone cannot selection natural , such that l , such
yellow hindwings may thus be an an be maythus hindwings yellow in
indicate indicate
the the t conspicuous to birds to t conspicuous s s ; it is ; on pressure
) resembling birds about unprofitability (Lindstedt et al. 2011 al. et (Lindstedt about unprofitability birds ,
same direction
more efficient & Reed f
emales prefer males that are more contrasting against theb against are more contrasting that males prefer emales are
thus that
arvae reared on a on reared arvae
with
patches of highly conspicuous conspicuous highly A. plantaginis
the un A. 2014)
black the balance between natural and sexual selection inthewell selection sexual natural and between balance the on likely likely
plantaginis variation in female hindwing colouration colouration hindwing infemale variation
), and learn the ), and learn
& (Fig. S1) female colouration . ly ,
-
This suggests thatsuggests pr This Osorio Osorio which may explain why only one morph occurs occurs onemorph whyonly explain which may and
to (Finkbeiner et al. 2014). al.2014). et (Finkbeiner than red postman butterflies ( butterflies red postman
, a contribut - ).
andthus l white males males .
(Hegna & Mappes 2014). 2014). &Mappes (Hegna .
achromatic models andmodels models achromatic 2010).
diet high diet
N explain explain 2013). The combination of of The combination 2013). aïve birds are birds aïve because
than with than and efficient efficient
red colour faster (Rönkä et al. et (Rönkä faster red colour e
in greater risk of predation in greaterrisk
to ma to the suggest that suggest in in , as birds avoid red females more often more red females avoid birds , as
of the high achromatic achromatic the high of iridoid glycosides iridoid maintenance of local polymorphism oflocal maintenance cuefor learning t
edator grey e choice.
more more
In con . T patche H
deterrence sexual hus, additional selective or selective hus, additional eliconius strongly strongly P ),
trast, redation s . it However selection may selection
black
is costly and is costlyand
erato bird become
is in attracted t attracted of
(aposematism
perceived as a perceived as small
, however, ) s. I
nonlocal (Morehouse & (Morehouse
and (luminance) attracted more attracted more
2018). n fact ,
ackground. ackground.
they adults with adults white cabbage cabbage o
in play a play
, birds birds ,
learn learn While While
the the ) - ,
This article isThis article by protected copyright. All rightsreserved. Accepted figures preparing discussed All authors models. vision calculatedcolour OL ERGs, andanalysed measured MJH eyehistology, AKstudied the species, on study knowledge andecological wingspectra animals, provided andBR JM theplanned study. andAK BR, JM, MJH contributions Author Kelber (2100000256) Interactions of earlier versions W with statistics. on analyses data advice for to Thomas Labhart equipment, eye types lepidopteran plantaginis of measurements reflectance providing forkindly C.Lindstedt and Nokelainen O. to animals, the for rearing ofJyväskylä Article theUniversity at staff andtheto greenhouse K.Suisto We are grateful Acknowledgements investigation selection her approached colo Galarza 2014; variable against predators advantage areusually genetic factors u ration is unlikely is ration (2012 selection by by selection ,
, to Carina Rasmussen for excellenthelp for Carina Rasmussen , to or
. whether other traitswhether other -
2212) and O. Lind (637 Lind andO. 2212) et al. , favour e thank ,
and the manuscript.
2014). 2014). predat
, , to Thomas Labhart, Eric Warrant andDan EricWarrant Labhart, Thomas , to approved
but but ing Alan Goldizen Alan involved
O to J. Mappes and B. Rojas, B.Rojas, and to J.Mappes the UV ors a female a while ur results ur
seem to offer an explanation for polymorphism (Nokelainen et al. al. et (Nokelainen polymorphism for explanation an toseem offer
publication. explain
white - Funding from Funding . reflecting reflecting
I may use chromatic cues formate cues usechromatic may n - 2013 on A. A.
and three anonymous reviewers reviewers threeanonymous and
male mating success of white males ofwhitemales mating success
A. plantaginis plantaginis - 388)
No author has any conflict ofinterest. has conflict any author No s white males. Whetherfemales white males. tend totend
is gratefully acknowledged. acknowledged. is gratefully the Finnish Centre of Excellence in Biological in Biological ofExcellence Centre the Finnish
with histology with , where , where and and
have have the vision confirm that male selection for female female selectionfor that male confirm vision results, c from the Swedish Research Research thefrom Swedish
and yellow yellow a mating a mating
to Alan Ho fordili Ho Alan to , to Dan to ,
males, on males, - ontributed E. Nilsson for accessto Nilsson for E. advantage choice better, - for critical E. Nilsson for insights into insights for E. Nilsson
use visual cues in mate cuesinmate use visual
average once a male hasmale a once
to writing text and to and text writing requires requires , gene flow , gene gent assistance gentassistance
comments on comments ,
have an an have Council to A. to Council further
and and A.
This article isThis article by protected copyright. All rightsreserved. AcceptedFinkbeiner Endler, J. Endler, J. L Crothers W. Conner, (2008) Briscoe, A.D. Brain Beluši Article(1982) Barlow, H.B. K., St Arikawa, References found All accessibility Data original original ,
ç M o , G., Šporar, K., & Megli & K., Šporar, , G., butterflies. attraction in andmate avoidance predator to pattern of colorand contributions American Naturalist of Biology Naturalist poison frog. ofapolytypic inpopulations selection natural radarof the work under Arctiidae. Biology Experimental Finland. ofJyväskylä, University Thesis, MSc tigermoth. wood female defence inthe mothborer Vision Research 1 andnon ofvisual Evolution & Collin P. S. Hankins, W. Hunt,M. D. M. adaptations. In visual n . - A. A. & A. A. the 4614 , .
R E. (Ed.) E. data used in this study study in this used data
S
( (1983) 2016 . & . . Dryad Digital Repository: https://doi.org/10.5061/dryad. s46t627. https://doi.org/10.5061/dryad. Repository: Dryad Digital D
- Mappes, J.(2004) Mappes, avenga, D.G. (2014) D.G. avenga, 4355 .
, Briscoe Cummings Fish
181 ) Oxford
Evolution
Can warning signals be honest? Wing colouration and the strength of chemical chemical of andthe strength colouration honest? Wing be signals Can warning Natural and sexual selecti sexual and Natural
Ostrinia ( -
What causes trichromacy? A theoretical analysis using comb using analysis Atheoretical trichromacy? What causes es, 2009 , Reconstructing the ancestral butterfly eye: focus on the opsins on the eye: focusbutterfly theancestral Reconstructing 1. , E116
22 9 , 173 ,
: Oxford Univers : Oxford , 635 ) A ,
,
Tiger Moths and Wolly Bears. Wolly and Tiger Moths
. 163 - , , 211 .
E124. D Biology of Experimental Journal M 68 – . & – 190. , 532 . , 1805 , 3410 E. ç - 6
visual pigments visual , A. (2017) Extreme polarisation sensitivity in the retina of the corn thecorn retina of inthe sensitivity polarisation Extreme , A.(2017) Predator mixes and the conspicuousness of aposematic signals. ofaposematic conspicuousness mixes andthe Predator 43.
Reed
that arenot (
2013
– Insect photopigments: photoreceptor spectral sensitivities and sensitivities spectral photoreceptor photopigments: Insect
547. – – 1813 3420. ,
) R
Warning signal brightness variation: sexual selection may selection may sexual brightness variation: signal Warning . D ity Press .
. on on color patterns in poeciliid fishes. fishes. inpoeciliid patterns color on on
(2014) Warning signals are seductive: relative seductive: relative are signals (2014) Warning
(pp. 137 (pp. presented in the main textorthe supplement in themain presented .
– 162). New York: Springer. doi:10.1007/978 162). NewYork:Springer. ,
220 Behaviour, Ecology and Evolution Ecology and Behaviour, , 2047
– 2056.
N. J. Marshall (Eds), Marshall (Eds), N. J.
-
filtered spectra. spectra. filtered . Environmental Environmental Journal Heliconius
of American s , can be be , can
of the of
- This article isThis article by protected copyright. All rightsreserved. AcceptedLanger Land Jiggins Jakobsson, A. Kelber,B. & J.,Rojas, O., Mappes, Lind, J., Henze, M. Hegna, R. ArticleHegna, J. & J. O.,Victorsson, Håstad, Govardovskii S. Gordon, Galarza , M.F. & Nilsson, D. & , M.F. , , H.&
, C Spodoptera exempta Spodoptera a tiger inmoth colour discrimination predators and of conspecifics theeyes through moth seen polymorphic 1469 ( moth woodtiger ofthe coloration in warning signal o America United States of eyesoftheirpredators. inthe conspicuous template. pigment visual Animal Ecology frequency positive apart byopposing plantaginis ( moth tiger wood the aposematic in signal variation between andwarning genetic
pattern mimicry. pattern mimicry. 161 of sensitivity andspectral sites oviposition J R. & Mappes, J.(201 Mappes, R. & . . H., D A
P., Kokko, H., Rojas, B., Nokelainen, B., Nokelainen, H.,Rojas, Kokko, P., J., Henze, M. J., Svensson, J., Henze, M. J., – . . , Naisbit , , Nokelainen , – 168 , Galarza, J. A. & J.A. Galarza, Hamann 1481.
V f the wood tiger moth in Alaska. inAlaska. moth the woodtiger f . . I ).
. , Fyhrquist Molecular Ecology Molecular
,
, R ,
84 B. . E
, Nature - , (Insecta: Noctuidae) (Insecta:
. E. (2002) E. & O , Coe , 1555
. Ödeen, A. (2005) Ödeen, A. Meinecke C Meinecke , Ashrafi ,
, 4 Mappes, J. (2015) J. Mappes, N
102 ) –
,
411
. Influences of geographic differentiation in the forewing warning warning intheforewing differentiation ofgeographic Influences Visual Neuroscience Visual , Reuter 1564 R , 6391 , . , L
,
23
Animal eyes. Oxford: Oxford University P University Oxford eyes.Animal Oxford: . 302 , Mallet ,
G. , R 4939 . – – , . P., Lind, O., Anderbrant, O. ( Anderbrant, O., P., Lind, , Hegna -
305. 6394. T C. . , Kuzmin
. – , - (1979)
dependent selection, yet maintained inspace selection,yetmaintained dependent Dryad Digital Repository. doi:10.5061/dryad.s46t627 doi:10.5061/dryad.s46t627 Repository. Digital Dryad J.
4957.
. Differences in color vision make passerines less less passerines make vision incolor Differences
J
( ournal of ournal Global phylogeography and geographical variation variation geographical and phylogeography Global Evolutionary Ecology 2001 , O. & Mappes, J. (2015) & Mappes,J. O.
R , Proceedings of the National Academ oftheNational Proceedings Cydia strobilella Cydia
. 17 Tetrachromatic visual s visual Tetrachromatic H ) , ,
. & .
Parasemia plantaginis Parasemia D Reproductive isolation caused by colour bycolour caused isolation Reproductive 509
Comp .
G Mappes (2018) Data from: An aposematic colour Anaposematic from: (2018) Data – . & 528 arative Physiology, arative
Donner
L. ,
, J . 28 Journ
(2014) , 1003 2017 ,
K
Colour Colour al of Insect Physiology, al ofInsect ). .
(2000) In search of the Insearch of (2000) ystem in themothystem in
) Visual cuesof ) Visual Journal of Biogeography of Journal Temporal relationship relationship Temporal –
1017 129 ress. -
polymorphism torn polymorphism sensitivity and sensitivity ,
y of Sciences of the of Sciences y of 235 .
– 239 Parasemia Parasemia . Journal of Journal . .
10 1 , ,
42 - ,
This article isThis article by protected copyright. All rightsreserved. Accepted J.(2014) C. &Mappes, J.,Lindstedt, O., Valkonen, Nokelainen, R O., Hegna, Nokelainen, Nokelainen N. Morehouse, (1999) & Joron, M. Mallet, J. (2009) M.E. Cummings, & Maan, M.E. Lyytinen ArticleAlatalo Lyytinen, A., Alatalo, L., Lindström, Limeri A., Kahilainen, E., H., Ihalainen, Eager, Lindstedt, C., J.H. ReudlerTalsma Lindstedt, C., , L.B. & Morehouse, N.H.(2014) &Morehouse, , L.B. 83 moths signals inArctiid warning of two the efficacy structure shifts B Society tigermoth. inthe success wood mating efficacyand signal warning Thesis, signals.PhD warning favours polymorphic heterogeneity 768 predat speciation. and balance, shifting America States of a poison frog in aposematic signals Lepidoptera andnocturnal aposematic signals America prey aposematic novel rare Colias of male systemthe visual through polymorphism female viewingthe 'alba' polymorphisms: Ecology tigermoth. wood of theaposematic color for the by predators of selection strength 64:68 a gener costs in excretion coloration indirectly: affects warning ,
A., Li A., , 598 , O. (2013) , – 784. -
butterflies. butterflies. 78. ion risk associated with an exaggerated male butterfly color. color. butterfly male exaggerated an associated with risk ion , I. &
– , ndström, L. & Mappes, J. (2004) J. &Mappes, L. ndström, 22 -
Biological Sciences 605. 98
, 580 , 9181 Rutowski, R.L. (2010) Rutowski, , R. ,
Many forms of the wood tiger moth( tiger the wood of forms Many – R.
V., Lindstr V., 106, 587. – Func V., Lyytinen, A. & Mappes, J. (2001) J. Lyytinen,A.&Mappes, V., .H., Reudler,J. .H., ? Behavioral Ecology ? Behavioral 9184.
19072 Evolution of diversity in warning color and mimicry: polymorphisms, polymorphisms, mimicry: and color inwarning diversity of Evolution tional
,
279 .
ö . Proceedings of the National Academy of Sciences of the United Statesof of the United Sciences of Academy National ofthe Proceedings . – , Ihalainen
Ecol Behavioral Ecology m, L. & Mappes, &Mappes, m, L. 19077. , 257
ogy, Sexual dimorphism and directional sexual selection onselection directional sexual and dimorphism Sexual
Annual Review of Ecology and Systematics Review ofEcology Annual Sensory limitations Sensory H., Lindstedt, C.& H., Lindstedt,
– In the eye of the beholders: female choice and avian andavian choice female of thebeholders: In theeye
. 265. 28 Proceedings of the National Academy of Sciences of the United United ofthe of Sciences Academy National ofthe Proceedings ,
, , 12
E 1197
, 65 ,
. Ultraviolet reflection and predation risk in diurnal in diurnal risk andpredation Ultraviolet reflection , Lindstr – – , 70.
1207. 15 J. (2001 J. Stevens, M. & Mappes, J.(2011) Mappes, & Stevens, M. ,
982 ö
m
and the maintenance of colour maintenance andthe – Mappes, J. Mappes, Parasemia plantaginis Parasemia , )
987. L
Strong antiapostatic selection against against selection antiapostatic Strong Can ultraviolet cues function as cuesfunction Can ultraviolet .
, Mappes Changes in predator community community predator in Changes
alist herbivore. Evolution herbivore. alist ( 2012 , J. University ofJyväskylä. University American Naturalist American . J Proceedings of the Royal the Proceedings of
( 2010 ournal of ournal ) ,
Trade 30 ): selective selective ): , 201 )
Diet quality Diet quality -
off between off Anim –
233. Direction and Direction al
Behavioral
, Ecol
176
ogy, ,
This article isThis article by protected copyright. All rightsreserved. Accepted Brandt M., Vorobyev, Telles, F. T. Ruxton, G.,Sherratt, Rönk Rönkä B.,Rojas, Go J.A. & Endler, B. Rojas, Rojas Article (201 R CoreTeam Poulton, Osorio Olofsson M.E.N. Majerus, P., O'Donald, ä ,
, , B ,
K., K receptor noise: behaviour and physiology compared. physiology compared. and behaviour receptor noise: A sensitivi theblue: spectral (Lepidoptera). ofthesubtribeArctiina molecular analysis 163. moth. an aposematic of morphs among predator generalization polymorphic frog theaposematic B Royal Society ofthe Proceedings How tof Austria. Vienna, Computing, Statistical 191 vertebrate enemies. totheir insects inreference 1256. chicks bypoultry colours novel e10798. d selection dependent sexual D .
, Burdfield , J., Lind, O.,Henz J., Lind, E. , eflect .
, , De Pasqual , De . M
, Ham 200 Mappes, Mappes, B. –
. 274. , Vallin , rdon, S.P. & Mappes, J.(2015) S.P.&Mappes, rdon,
( , 537 1887 b
ird ird ,
ight multiple enemies: target enemies: multiple ight
A 7 - – ) ) . a Steel , J.,
D
546 ttacks R: A language and environment for statistical for andenvironment R: Alanguage The experimental proof of the protective value of colour and markings in in markings and colourof value of theprotective proof The experimental wood tiger moth. woodtiger A ,
. , Kaila , Gonda R., Peitsch, Peitsch, R., .
N. , Jakobsson C . , (2013) . , Mappes , E e, M.J., Rodriguez M.J., e, & u . , L. Dendrobates tinctorius , Pakkanen nder
Speed, M. Speed,
,
(
Sexual dimorphism and intra and dimorphism Sexual & Z 1984 l
. & . ow ow . Wahlberg D., Laughlin, S. D., Laughlin,
, ,
Biological Journal of the Linnean Society the Linnean Journalof Biological
J S - . ty of hummingbird hawkmoths. hawkmoths. hummingbird ty of ) Biological Sciences Biological . & , Gordon Andrew
l
Polymorphism of melanic ladybirds maintained by frequency maintained ladybirds melanic of Polymorphism ight . Quarterly Journal of Experimental Psychology Experimental of . Quarterly Journal P. Current Zoology Current ,
H
Wiklund ( . 2004 , Suisto i
ntensities with UV with ntensities , N.( Frequency - - Girones, M. Girones, , specific chemical defences in an aposematic moth. aposematic defences inan chemical specific
R , URL http://www.R URL . Evolutionary Ecology 27, Evolutionary . )
S . Avoiding Attack Avoiding J 2016 . , , Rojas . , B. & B.
C (2009) Sensory generalization and learning about about learning and generalization Sensory (2009) K . . ,
.
, Maczka , (2010) Marginal Marginal (2010) In press. In ) 61 -
P Putting Parasemia in its phylogenetic place: a place: itsphylogenetic in Putting Parasemia inthecolour activity dependent flight Menzel, R. Menzel, , 765 , roceedings ofthe roceedings
A., Goyret, J. & Kelber, A. (2014) Out of A. (2014)Out J.&Kelber, A., Goyret, B. (2018) - Vision Res Vision – populational colour pattern variation in pattern variation colour populational
772. , w
. Oxford University . Oxford M
avelengths. avelengths. Systematic Entomology Systematic
. computing. R Foundation for RFoundation computing. , Schulz ,
(2001)
- Colour alone matters: nomatters: alone Colour project.org/. Journal of Comparative Physiology Physiology of Comparative Journal e
earch, Zool 23 yespots on on yespots Animal Behaviour Animal
, Colour thresholds and and thresholds Colour
, 101 739 ogical
S 41, PLoS ONE PLoS .,
– – &
639 111. 753
Soc (Hove)
Mappes b iety of London, iety of –
. Press utterfly
653.
,
41 ,
62
135 , 844 5
, . (5): (5):
, 1249 J. (2017) , w
153 – ings ings 853 – – 55 .
, -
This article isThis article by protected copyright. All rightsreserved. Accepted references Supplementary Figure S3 Figure S2 Figure S1 S3 Table S2 Table S1 Table methods Supplementary ofthisarticle. the onlineversion foundin be may information Additional supporting INFORMATION SUPPORTING Article Zylinski & Stiles,W. G. Wyszecki, H. Xu, White, R.H., E. Warrant, Vorobyev Journal of Experimental Biology ofExperimental Journal f regional specialization ofgreen mosaic the expression, rhodopsin Handbook of the Royal ,
ormulae, 2nd Edn. New York: Wiley. New Edn. York: 2nd ormulae, S , Sex and time effects andtime Sex Chromatic contrasts for equally spacedreceptors forequally Chromatic contrasts proportions receptor for1:1:1 Chromatic contrasts .
Achromatic (Michelson) contrasts of wing colours ofwing (Michelson)contrasts Achromatic , Osorio Effect of i Effect proportions for1:1:1receptor contrasts Chromatic M J., Kelber, A. & Kristensen, N & Kristensen, Kelber, A. J., . , Osorio Society B Society ,
H. Z D llumina oology Vol. IV 36, Lepidoptera 2 Lepidoptera IV36, Vol. oology , .
Munch, T Munch, D. (2013) Visual contrast and color in rapid learning of novel patterns by chicks. by chicks. patternsnovel of learning in rapid color and contrast Visual (2013) - Biological Sciences Biological S.
(1998) Receptor noise as a determinant of colour thresholds. thresholds. colour of asadeterminant noise Receptor (1998) for ERGs and visual models visual and ERGs for (2000) tion
.
on ERGs on ERGs Journal of Experimental Biology ofExperimental Journal
. on chromatic contrasts chromatic on A
Color Color
., ., ,
216 Bennett, R Bennett, , 4184 ,
s
.P. (2003) 265
cience: ,
– 351 . 4189. R c . - –
oncepts and oncepts and Grable, E Grable, , blue ,
358 Eyes and vision. andIn Eyes (p
p. 325 .
-
and UV and
206, 3337 206, – .
A 360
. (2003) m
) ethods, ethods, . - sensitive photoreceptors, and photoreceptors, sensitive Berlin: De Gruyter De Berlin:
– 3348.
The retina of The retina N. P. Kristensen (E N. P.Kristensen q uantitative uantitative
Manduca sexta Manduca d .
ataand Proceedings of Proceedings d.)
: Yellow White Yellow White This article isThis article by protected copyright. All rightsreserved. brackets. percentiles in woodtiger 2 Table Measurement Sex Slope Measurement Sex K Measurement Sex V of of eyes thecompound retina in 1. Table
max [
V log
Accepted Article
max at [
mV I ] , K
.
K
Influence C [ ]
mV/
0.9 and olour contrast olour moths Male hindwing colour hindwing Male 0. Male forewing colour forewing Male 1
no. no. no. White White (0. (0. S
log lope 3
1 .
- -
of sex and measurement number on number measurement and of sex For each com For each 0
I 1.6 at
] . 3 )
K )
difference see Fig.2. see
(
jnds Mean - 12. - 0.0 0.1 3. 0 0.1 .
1 1 ) 0 2
A. plantaginis
between
2. 0.7 1.0 0
parison
.9
6
Yellow Yellow
( ( ( ( 0.5 1.8 0.3 0.7
Standard Standard
- - - - , patches of 1.5 3.3 1.2 1.4 we give error 0.04 0.01
0.4 0. 0.1 1.4
) ) ) )
2 tested
the
by linear mixed by linear Orange Orange forewing
Red Red median ( - confidence ( ( 0. ( ( -
(2. 9.1 response properties of the dark ofthe response properties - - 0.03 interval 0. 0.2 1
95% 95% 2 , 5 ,
, - , 0. , 0. 15 0.0 , 0. 4.0 s colour contrast withthe contrast colour
and hindwing and . 3 3 0
01 1 )
) ) )
)
)
- 1.4 1.1 effects models. models. effects Orange Orange
- - ( ( Female hindwing colour hindwing Female 1.23 7.69 0.54 8.87 2.10 0.72 0.6 0.5 Female t
- -
2.7 1.8
s in male andfemale s inmale
) )
forewing colour forewing 130 130 130 12 12 12 df
For a
25 - adapted th definition definition
and75 0.5 1.7 0.5 1.1 <0.001 <0.001 p
0 0. 0.4 0. value
.22 (0. ( (0. ( 04 59
Red Red
1 0.6
8 .
2 1 2 th - - - -
0 2 0 1.4 . . . 8 5 9 ) ) ) )
This article isThis article by protected copyright. All rightsreserved. andgreen receptors. UV ofthe theresponses isolating bybluelight, suppressed receptors were with with thereceptors UV Exampl ofphotoreceptor. spectral types sensitivity( ofmaximal the wavelength toestimate selected stateswere adaptation ERGs. Fig. at all from of data K response maximal theasymptotic andestimated each dataset to Rushton function hyperpolarization shown to inresponse moth afemale and ofamale eyes compound the from (ERGs) (A) Electroretinograms Fig. 2. 100 scale bar bas plantaginis Fig F
igure captions igure thata half elicited K 40 ms 40
Accepted Articleement .
(mean +/ (mean λ 3 1 . α . . (B) Amplitudes . (B) Amplitudes
Vertical Spectral sensitivities of male (left) and female (right) female and (left) male of sensitivities Spectral Sex of the green and UV receptors predetermined (E, F). (G, H) Examples, in which the blu inwhich H) Examples, F). (G,(E, predetermined receptors andUV green of the For each individual, averaged measurements ( measurements averaged individual, For each flashes of white light with increasing intensity intensity increasing with ofwhite light flashes
membrane, scale bar scale50 membrane,
differences in response strength ofthedark strength in response differences .
Left, enlarged region of the eye with c cornea, cc crystalline cone, cone, cc crystalline eye withccornea, ofthe region Left, enlarged - m.
standard error) tested standard error) eight sections of
)
plotted against intensity intensity against plotted - maximal response, and the and response, maximal
males and males λ V α
of the green receptors predetermined (C, D), and D), (C, predetermined receptors theof green
( potential changes from the baseline at stimulus onset to maximal maximal to onset stimulus the baseline at from potential changes the compound the compound six
females. (C) Sex differences of the parameters the of (C) differences females. Sex m. Right, eyes by linear mixed linearby e curves used to assess to assess curves used e I
for the datasets partially shown in A. We fittedaNaka shown inA. partially for thedatasets eyes S lope ( of R retina
of thecurve at of female ( female - n effects models. effects
= 4) from one compound eye under different underdifferent eye compound one = 4)from (see ) and first optic neuropile, optic neuropile, first ) and -
B adapted retina in adapted ) top . For clarity, . For λ
α Arctia Arctia
) and male ( male ) and of the green receptors (A, B), (A, of thegreen receptors K
. The inset illustratesthe . Theinset plantaginis
only every second ERG is ERG second every only λ α
of the blue receptors the bluereceptors of bottom Arctia plantaginis Arctia rh rhabdom, rh rhabdom, λ V V α ) forthree ) max max
derived from derived from lamina ) , the intensity ,
K Arctia Arctia , and
b (L)
range range S
e , λ lope α
. of -
This article isThis article by protected copyright. All rightsreserved. not wellkno however in set to detection is 1jnd, ( countedas outliers interval are this outside 25th).Data points ±1.5 of a maximum to of the distributions therange spanning red 10 colour hindwing and forewing calculatedfor M Fig s of andindependent noise set by receptor diagram inthe chromaticity axes opponent specific onvisualized equal between loci (brown background brown locus ofthe by illuminated Arctia Fig Stiles 2000 R middle MWS, wavelength sensitive, ( theblue tit theblue receptors in sensitivity of but the species,both same in arethe 2000) etal. (Govardovskii thetemplate basedon ( tigermoth( ofthewood eyes compound Fig. females red orange females,10 8 yellowmales, white males,10 female Fig. 4 Macroglossum stellatarum Macroglossum eflectance spectr eflectance Acceptedacroglossum Article 7 6 5. . . .
The loci offorewi The loci Colour contrast between wing patches and and patches wing contrastbetween Colour
female plantaginis
Data R Arctia eflectance of ).
Cyanistes caeruleus wn (see wn stellatarum used s.
daylight d65 plantaginis B oxes show the 25th, 50th, and 75th percentiles of colour contrast with whiskers with whiskers contrast colour of percentiles and75th the 50th, 25th, oxes show a of green and brown vegetation andbrown of green a s
in model calculations. model in .
their contrast expressed in jnds (see M (see in jnds expressed their contrast M Each data point represents the colour locus given by a reflectance spectrum spectrum a reflectance by locusgiven thecolour point represents Each data A.
ethods) (white), ; data from Telles al fromet ; data
forewing p lantaginis ng (A) and hindwing (B) colours colours (B) hindwing and (A) ng . Receptors are adapted to a green background (green background a green toadapted Receptors are . . ; data from ; data from L ines give averages and ines giveaverages .
A rctia
( is dashed lines dashed - wavelength sensitive, LWS, long LWS, wavelength sensitive, the moths red
plantaginis - triangle Hart et al. Hart et shifted. (B) shifted. Arctia plantaginis pecific colour
(A) S s
.
from 16 white and from 16white , this limit is uncertain asthe uncertain limitis , this 2014)
(red) comparison. for ) isshown ) and hindwing ) and pectral sensitivities of photoreceptors in the in of photoreceptors sensitivities pectral
2000). and . The sensitivities for UV and green receptors greenreceptors and for UV sensitivities . The S grey and and pectral sensitivities of the four single cones in cones single ofthefour sensitivities pectral (A)
i
rradiance of daylight (d65 daylight rradiance of opponency (Vorobyev and Osorio and (Vorobyev opponency ) and ) and
C area UVS, ultraviolet sensitive, SWS, short SWS, ultraviolet UVS, sensitive, a green, green, a ethods for details). C details). for ethods yanistes s in the chromaticity diagram chromaticity in the . the hummingbird hawkmoth hawkmoth the hummingbird crosses
standard deviations standard times the
10 yellow 10
but
( c aeruleus solid lines solid or -
the contrast between colours is is betweencolours thecontrast wav ). The (B)
interquartile range (75th range interquartile el
(blue). Contrast was (blue). was Contrast
a brown background brown background a ength sensitive. (C) sensitive. ength The males receptor
absolute threshold of of threshold absolute ) colours colours olour loci are lociare olour Eu , triangle ; Wyszecki and and ; Wyszecki and 8 orange and . Data from 16 . Datafrom clidean distance clidean distance noise levels are levels noise of male male of ) ,
the colour the colour
1998). 1998).
of
- and and a - in nd
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article
This article isThis article by protected copyright. All rightsreserved. Accepted Article