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Effects of directionality, signal intensity, and short-wavelength components on iridescent warning signal efficacy

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Abstract

Many bright colors function as visual signals and are produced by structural mechanisms that result in their being iridescent. Iridescent colors differ from other types of animal coloration in that they vary in both hue and intensity, as the angle of viewing and illumination change relative to the color surface. Iridescent colors can serve as warning signals by deterring predation on distasteful animals. Due to their directional and mirror-like reflection, iridescent signals have the potential to (1) vary in appearance with each approach; (2) create a flashing signal; (3) under the right conditions, be more intense than diffusely reflecting signals; (4) display different hues with different positioning of the sun, predator, and prey; and (5) display angle-dependent camouflage, which could all influence warning signal effectiveness. Here, we examine how signal intensity, short-wavelength hue, and variation in appearance affect the response of domestic chickens (Gallus gallus domesticus) to warning signals. Variation did not affect their response. Higher intensity signals were more effective in terms of predator avoidance than were lower intensity, and blue-violet signals were more effective than were blue and blue-green. This could influence the effectiveness of an iridescent warning signal in nature if the prey displays more intense, blue-violet coloration. With the properties we tested, we found no cost to having warning colors that are iridescent and that there may be benefits in the ability to change hue and display a more intense signal. We suggest future research to examine the effects of iridescent flashing and angle-dependent camouflage on warning signal effectiveness.

Significance statement

Iridescent colors have only been recently shown to function as warning signals, but we know very little about how they function to deter predation. Here, we show that birds are more likely to change their behavior in response to higher intensity and blue-violet signals compared to low intensity and blue and blue-green signals. The intensity results are consistent with previous experiments, but the role of short-wavelength hue on warning signal effectiveness has never been tested before. Variation due to the shifting appearance of an iridescent signal did not influence the response to the warning signal, consistent with studies in long-wavelength signals. We found no cost to displaying an iridescent signal and potentially an adaptive benefit as iridescent warning signals have the potential to be brighter and display a different hue based on the arrangement of light source, signaler, and receiver.

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References

  • Aronsson M, Gamberale-Stille G (2008) Domestic chicks primarily attend to colour, not pattern, when learning an aposematic coloration. Anim Behav 75:417–423

    Article  Google Scholar 

  • Aronsson M, Gamberale-Stille G (2009) Importance of internal pattern contrast and contrast against the background in aposematic signals. Behav Ecol 20:1356–1362

    Article  Google Scholar 

  • Barron AB, Hebets EA, Cleland TA, Fitzpatrick CL, Hauber ME, Stevens JR (2015) Embracing multiple definitions of learning. Trends Neurosci 38:405–407

    Article  CAS  PubMed  Google Scholar 

  • Beatty CD, Beirincky K, Sherratt T (2004) The evolution of Müllerian mimicry in multispecies communities. Nature 431:63–67

    Article  CAS  PubMed  Google Scholar 

  • Bradbury JW, Vehrencamp SL (2011) Principles of animal communication, vol 2. Sinauer Associates Inc., Sunderland, MA

    Google Scholar 

  • Cott HB (1940) Adaptive coloration in animals. Methuen, London

    Google Scholar 

  • Crothers LR, Cummings ME (2013) Warning signal brightness variation: sexual selection may work under the radar of natural selection in populations of a polytypic poison frog. Am Nat 181:E116–E124

    Article  PubMed  Google Scholar 

  • Dakin R, Montgomerie R (2009) Peacocks orient their courtship displays towards the sun. Behav Ecol Sociobiol 63:825–834

    Article  Google Scholar 

  • DeCock R, Matthysen E (1999) Aposematism and bioluminescence: experimental evidence from Glow-worm larvae (Coleoptera: Lampyridae). Evol Ecol 13:619–639

    Article  Google Scholar 

  • Doucet SM, Meadows MG (2009) Iridescence: a functional perspective. J R Soc Interface 6:S115–S132

    Article  PubMed  PubMed Central  Google Scholar 

  • Exnerová A, Svádová K, Štys P, Barcalová S, Landová E, Prokopová M, Fuchs R, Socha R (2006) Importance of colour in the reaction of passerine predators to aposematic prey: experiments with mutants of Pyrrhocoris apterus (Heteroptera). Biol J Linn Soc 88:143–153

    Article  Google Scholar 

  • Fabricant SA, Exnerová A, Ježová D, Stys P (2014) Scared by shiny? The value of iridescence in aposematic signaling of the hibiscus harlequin bug. Anim Behav 90:315–325

    Article  Google Scholar 

  • Fan R, Hansson BS (2001) Olfactory discrimination conditioning in the moth Spodoptera littoralis. Physiol Behav 72:159–165

    Article  CAS  PubMed  Google Scholar 

  • Fernandez PC, Locatelli FE, Person-Rennell N, Deleo D, Smith BH (2009) Associative conditioning tunes transient dynamics of early olfactory processing. J Neurosci 29:10191–10202

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Fleishman LJ, Persons M (2001) The influence of stimulus and background colour on signal visibility in the lizard Anolis cristatellus. J Exp Biol 204:1559–1575

    CAS  PubMed  Google Scholar 

  • Gamberale G, Tullberg BS (1996) Evidence for a peak-shift in predator generalization among aposematic prey. Proc R Soc Lond B 263:1329–1334

    Article  CAS  Google Scholar 

  • Gamberale-Stille G, Tullberg BS (1999) Experienced chicks show biased avoidance of stronger signals, an experiment with natural colour variation in live aposematic prey. Evol Ecol 13:579–589

    Article  Google Scholar 

  • Gellerman LW (1933) Chance orders of alternating stimuli in visual discrimination experiments. J Genet Psychol 42:206–208

    Google Scholar 

  • Gerber B, Geberzahn H, Hellstern F, Klein J, Kowalsky O, Wüstenberg D, Menzel R (1996) Honey bees transfer olfactory memories established during flower visits to a proboscis extension paradigm in the laboratory. Anim Behav 52:1079–1085

    Article  Google Scholar 

  • Ghiradella H (1985) Structure and development of iridescent Lepidopteran scales: the Papilionidae as a showcase family. Ann Entomol Soc Am 78:252–264

    Article  Google Scholar 

  • Ghirlanda S, Enquist M (2003) A century of generalization. Anim Behav 66:15–16

    Article  Google Scholar 

  • Gittleman JL, Harvey PH (1980) Why are distasteful prey not cryptic? Nature 286:149–150

    Article  Google Scholar 

  • Gomez D (2006) AVICOL, a program to analyse spectrometric data. Last update January 2012. http://sites.google.com/site/avicolprogram/

  • Guilford T, Dawkins MS (1993) Are warning colors handicaps? Evolution 47:400–416

    Article  Google Scholar 

  • Hailman JP (1977) Optical signals: animal communication and light. Indiana University Press, Bloomington

    Google Scholar 

  • Ham AD, Ihalainen E, Lindström L, Mappes J (2006) Does colour matter? The importance of colour in avoidance learning, memorability and generalisation. Behav Ecol Sociobiol 60:482–491

    Article  Google Scholar 

  • Hauglund K, Hagen SB, Lampe HM (2006) Responses of domestic chicks (Gallus gallus domesticus) to multimodal aposematic signals. Behav Ecol 17:392–398

    Article  Google Scholar 

  • Hinton HE (1973) Natural deception. In: Gregory RL, Gombrich EE (eds) Nature and art. Charles Scribener’s Sons, New York, pp 97–160

    Google Scholar 

  • Ihalainen E, Lindström L, Mappes J (2007) Investigating Müllerian mimicry: predator learning and variation in prey defences. J Evol Biol 20:780–791

    Article  CAS  PubMed  Google Scholar 

  • Ihalainen E, Lindström L, Mappes J, Puolakkainen S (2008) Can experienced birds select for Müllerian mimicry? Behav Ecol 19:362–368

    Article  Google Scholar 

  • Kazemi B, Gamberale-Stille G, Tullberg BS, Leimar O (2014) Stimulus salience as an explanation for imperfect mimicry. Curr Biol 24:1–5

    Article  Google Scholar 

  • Kelly DJ, Marples NM (2004) The effects of novel odour and colour cues on food acceptance by the zebra finch, Taeniopygia guttata. Anim Behav 68:1049–1054

    Article  Google Scholar 

  • Land MF (1972) The physics and biology of animal reflectors. Prog Biophys Mol Bio 24:75–106

    Article  CAS  Google Scholar 

  • Lindstedt C, Eager H, Ihalainen E, Kahilainen A, Stevens M, Mappes J (2011) Direction and strength of selection by predators for the color of the aposematic wood tiger moth. Behav Ecol 22:580–587

    Article  Google Scholar 

  • Lindström L, Alatalo RV, Mappes J (1999) Reactions of hand-reared and wild-caught predators toward warningly colored, gregarious, and conspicuous prey. Behav Ecol 10:317–322

    Article  Google Scholar 

  • Loyau A, Gomez D, Moureau B, Théry M, Hart NS, Jalme MS, Bennett ATD, Sorci G (2007) Iridescent structurally based coloration of eyespots correlates with mating success in the peacock. Behav Ecol 18:1123–1131

    Article  Google Scholar 

  • Mappes J, Alatalo RV (1997) Batesian mimicry and signal accuracy. Evolution 51:2050–2053

    Article  Google Scholar 

  • Mäthger LM, Bell GRR, Kuzirian AM, Allen JJ, Hanlon RT (2012) How does the blue-ringed octopus (Hapalochlaena lunulata) flash its blue rings? J Exp Biol 215:3752–3757

    Article  PubMed  Google Scholar 

  • Mery F, Burns JG (2010) Behavioural plasticity: an interaction between evolution and experience. Evol Evol 24:571–583

    Article  Google Scholar 

  • Osorio D, Jones CD, Vorobyev M (1999a) Accurate memory for colour by not pattern contrast in chicks. Curr Biol 9:199–202

    Article  CAS  PubMed  Google Scholar 

  • Osorio D, Vorobyev M, Jones CD (1999b) Colour vision of domestic chicks. J Exp Biol 202:2951–2959

    CAS  PubMed  Google Scholar 

  • Pegram KV, Han HA, Rutowski RL (2012) Overnight perching aggregations of the aposematic Pipevine Swallowtail (Battus philenor: Lepidoptera: Papilionidae): implications for predation risk and warning signal use. J Res Lepidoptera 45:9–16

    Google Scholar 

  • Pegram KV, Han HA, Rutowski RL (2015) Warning signal efficacy: assessing the effects of color, iridescence, and time of day in the field. Ethology 121:1–13

    Article  Google Scholar 

  • Pegram KV, Lillo MJ, Rutowski RL (2013a) Iridescent blue and orange components contribute to the recognition of a multicomponent warning signal. Behaviour 150:321–336

    Google Scholar 

  • Pegram KV, Nahm AC, Rutowski RL (2013b) Warning color changes in response to food deprivation in the pipevine swallowtail butterfly (Battus philenor). J Insect Sci 13:1–16

    Article  Google Scholar 

  • Pegram KV, Rutowski RL (2014) Relative effectiveness of blue and orange warning colours in the contexts of innate avoidance, learning and generalization. Anim Behav 92:1–8

    Article  Google Scholar 

  • Pérez i de Lanuza G, Font E (2014) Now you see me, now you don’t: iridescence increases the efficacy of lizard chromatic signals. Naturwissenschaften 101:831–837

    Article  PubMed  Google Scholar 

  • Pike TW (2015) Interference coloration as an anti-predator defence. Biol Lett 11:20150159

    Article  PubMed  PubMed Central  Google Scholar 

  • Pinheiro CEG (1996) Palatability and escaping ability in Neotropical butterflies: tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae). Biol J Linn Soc 59:351–365

    Article  Google Scholar 

  • Poulton EB (1890) The colours of animals: their meaning and use especially considered in the case of insects. Kegen Paul, Trench, Trubner, and Co. Ltd., London

  • Prudic KL, Skemp AK, Papaj DR (2007) Aposematic coloration, luminance contrast, and the benefits of conspicuousness. Behav Ecol 18:41–46

    Article  Google Scholar 

  • Rescorla RA (1988) Behavioral studies of Pavlovian conditioning. Ann Rev Neurosci 11:329–352

    Article  CAS  PubMed  Google Scholar 

  • Riipi M, Alatalo R, Lindström L, Mappes J (2001) Multiple benefits of gregariousness cover detectability costs in aposematic aggregations. Nature 413:512–514

    Article  CAS  PubMed  Google Scholar 

  • Roper TJ (1990) Responses of domestic chicks to artificially coloured insect prey: effects of previous experience and background colour. Anim Behav 39:466–473

    Article  Google Scholar 

  • Roper TJ, Wistow R (1986) Aposematic colouration and avoidance learning in chicks. Q J Exp Psychol B 38:141–149

    Google Scholar 

  • Rowe C, Guilford T (1996) Hidden colour aversions in domestic chicks triggered by pyrazine odours of insect warning displays. Nature 383:520–522

    Article  CAS  Google Scholar 

  • Rowe C, Lindström L, Lyytinen A (2004) The importance of pattern similarity between Müllerian mimics in predator avoidance learning. Proc R Soc Lond B 271:407–413

    Article  Google Scholar 

  • Rowland HM, Ihalainen E, Lindström L, Mappes J, Speed MP (2007) Co-mimics have a mutualistic relationship despite unequal defences. Nature 448:64–67

    Article  CAS  PubMed  Google Scholar 

  • Rutowski RL, Macedonia JM, Kemp DJ, Taylor-Taft L (2007) Diversity in structural ultraviolet coloration among female sulphur butterflies (Coliadinae, Pieridae). Arthropod Struct Dev 36:280–290

    Article  PubMed  Google Scholar 

  • Rutowski RL, Nahm AC, Macedonia JM (2010) Iridescent hindwing patches in the Pipevine Swallowtail: differences in dorsal and ventral surfaces relate to signal function and context. Funct Ecol 24:767–775

    Article  Google Scholar 

  • Ruxton GD, Franks DW, Balogh ACV, Leimar O (2008) Evolutionary implications of the form of predator generalization for aposematic signals and mimicry in prey. Evolution 62:2913–2921

    Article  PubMed  Google Scholar 

  • Sandre SL, Stevens M, Mappes J (2010) The effect of predator appetite, prey warning coloration, and luminance on predator foraging decisions. Behaviour 147:1121–1143

    Article  Google Scholar 

  • Sandre SL, Tammaru T, Esperk T, Julkunen-Tiitto R, Mappes J (2007) Carotenoid-based colour polyphenism in a moth species: search for fitness correlates. Entomol Exp Appl 124:269–277

    Article  CAS  Google Scholar 

  • Sargent TD (1990) Startle as an anti-predator mechanism, with special reference to the underwing moths (Catocala). In: Evans DL, Schmidt JO (eds) Insect defenses: adaptive mechanisms and strategies of prey and predators. SUNY Press, Albany

    Google Scholar 

  • Schultz TD, Finke OM (2009) Structural colours create a flashing cue for sexual recognition and male quality in a Neotropical giant damselfly. Funct Ecol 23:724–732

    Article  Google Scholar 

  • Sicsú P, Manica LT, Maia R, Macedo RH (2013) Here comes the sun: multimodal displays are associated with sunlight incidence. Behav Ecol Sociobiol 67:1633–1642

    Article  Google Scholar 

  • Smith BH, Abramson CI, Tobin TR (1991) Conditional withholding of proboscis extension in honeybees (Apis mellifera) during discriminative punishment. J Comp Psychol 105:345–356

    Article  CAS  PubMed  Google Scholar 

  • Smith SM (1975) Innate recognition of coral snake pattern by a possible avian predator. Science 187:759–760

    Article  CAS  PubMed  Google Scholar 

  • Svádová K, Exnerová A, Stys P, Landová E, Valenta J, Fučíková A, Sochad R (2009) Role of different colours of aposematic insects in learning, memory and generalization of naïve bird predators. Anim Behav 77:327–336

    Article  Google Scholar 

  • Umbers KDL, Lehtonen J, Mappes J (2015) Deimatic displays. Curr Biol 25:R58–R59

    Article  CAS  PubMed  Google Scholar 

  • Uy JAC, Endler JA (2004) Modification of the visual background increases the conspicuousness of golden-collared manakin displays. Behav Ecol 15:1003–1010

    Article  Google Scholar 

  • Vorobyev M, Osorio D (1998) Receptor noise as a determinant of colour thresholds. Proc R Soc Lond B 265:351–358

    Article  CAS  Google Scholar 

  • Vorobyev M, Osorio D, Bennett ATD, Marshall NJ, Cuthill IC (1998) Tetrachromacy, oil droplets and bird plumage colours. J Comp Physiol A 183:621–633

    Article  CAS  PubMed  Google Scholar 

  • Vukusic P, Sambles JR, Lawrence CR, Wooton RJ (2001) Structural color: now you see it—now you don’t. Nature 410:36

    Article  CAS  PubMed  Google Scholar 

  • White TE, Zeil J, Kemp DJ (2015) Signal design and courtship presentation coincide for highly biased delivery of an iridescent butterfly mating signal. Evolution 69:14–25

    Article  PubMed  Google Scholar 

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Acknowledgments

We thank Kaci Fankhauser, Aly Galt, Sean Hannam, Drew Hensley, Ashley Lewis, Julea Shaw, and Mike Shillingburg for the assistance on this project; Brett Seymoure, Nikos Lessios, Martin Bergman, Kevin McGraw, and Brian Smith for the helpful discussions on the methodology and for comments on the manuscript. We also thank the anonymous reviewers for their helpful contributions. This work was supported by the National Science Foundation (IOS 1145654 to RLR).

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Correspondence to Kimberly V. Pegram.

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All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. All procedures performed involving animals were in accordance with the ethical standards of the institution or practice at which the studies were conducted. All procedures used in this study were approved by the Arizona State University Institutional Animal Care and Use Committee (Protocol no. 14-1349R). At the end of the experiment, most of the chickens were adopted for personal use but some males were humanely euthanized. This article does not contain any studies with human participants performed by any of the authors.

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The authors declare that they have no competing interest.

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Pegram, K.V., Rutowski, R.L. Effects of directionality, signal intensity, and short-wavelength components on iridescent warning signal efficacy. Behav Ecol Sociobiol 70, 1331–1343 (2016). https://doi.org/10.1007/s00265-016-2141-z

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