Behavioral Ecology and Sociobiology

, Volume 67, Issue 9, pp 1425–1439 | Cite as

Why are warning displays multimodal?

  • Candy RoweEmail author
  • Christina Halpin


Multimodal defensive displays are commonplace, with prey combining conspicuous coloration, sounds, odours and other chemical emissions to deter predators. These components can signal to predators in multiple signal modalities to warn them that prey are defended. The aim of our review is to examine the form and function of multimodal warning displays. Data collected from the literature on multimodal insect warning displays show the degree of complexity and diversity that needs to be explained, and we identify patterns in the data that may be worthy of more rigorous investigation. We also provide a theoretical framework for the study of multimodal warning displays, and evaluate the evidence for different functional hypotheses that can explain their widespread evolution. Our review highlights that whilst multimodal warning displays are well documented, particularly in insects, we lack a good understanding of their function in natural predator–prey systems.


Aposematism Mimicry Multimodal warning display Multicomponent signal Receiver psychology Predator Prey 



We would like to James Higham and Eileen Hebets for inviting us to submit a review to this special issue, and to Eileen and two anonymous reviewers for their enormously stimulating and helpful comments on the manuscript. We would also like to thank Melissa Bateson, Ben Brilot, Sue Healy, Domhnall Jennings, John Skelhorn, Martin Stevens and Jeri Wright for helpful discussions (that they may or may not remember) on various aspects of the manuscript. The review was supported by a BBSRC and NERC co-funded project grant (BB/G00188X/1).


  1. Alatalo RV, Mappes J (1996) Tracking the evolution of warning signals. Nature 382:708–710CrossRefGoogle Scholar
  2. Alexander RD (1964) Acoustic communication in arthropods. Ann Rev Ent 12:495–526CrossRefGoogle Scholar
  3. Barber JR, Conner WE (2007) Acoustic mimicry in a predator–prey interaction. Proc Nat Acad Sci 104:9331–9334PubMedCrossRefGoogle Scholar
  4. Barnett CA (2007) The effects of energetic and physiological state on the foraging decisions of birds. Unpublished PhD thesis, Newcastle UniversityGoogle Scholar
  5. Barnett CA, Skelhorn J, Bateson M, Rowe C (2012) Educated predators make strategic decisions to eat defended prey according to their toxin content. Behav Ecol 23:418–424CrossRefGoogle Scholar
  6. Bates HW (1862) Contributions to an insect fauna of the Amazon valley (Lepidoptera: Heliconidae). Trans Linn Soc Lond 23:495–566CrossRefGoogle Scholar
  7. Bates DL, Fenton MB (1990) Aposematism or startle? Predators learn their responses to the defenses of prey. Can J Zool 68:49–52CrossRefGoogle Scholar
  8. Bedford GO, Chinnick LJ (1966) Conspicuous displays in two species of Australian stick insects. Anim Behav 14:518–521PubMedCrossRefGoogle Scholar
  9. Bennett AD, Cuthill IC (1994) Ultraviolet vision in birds: what is its function? Vis Res 34:1471–1478PubMedCrossRefGoogle Scholar
  10. Bezzerides AL, McGraw KJ, Parker RS, Husseini J (2007) Elytra color as a signal of chemical defense in the Asian ladybird beetle Harmonia axyridis. Behav Ecol Sociobiol 61:1401–1408CrossRefGoogle Scholar
  11. Bisset GW, Frazer JFD, Rothschild M, Schachter M (1960) A pharmacologically active choline ester and other substances in the garden tiger moth Arctia caja (L.). Proc R Soc Lond B 152:255–262PubMedCrossRefGoogle Scholar
  12. Blest AD (1964) Protective display and sound production in some New World arctiid and ctenuchid moths. Zoologica 49:161–181Google Scholar
  13. Blount JD, Rowland HM, Drijfhout FP, Endler JA, Inger R, Sloggett JJ, Hurst GDD, Hodgson DJ, Speed MP (2012) How the ladybird got its spots: effects of resource limitation on the honesty of aposematic signals. Funct Ecol 26:334–342CrossRefGoogle Scholar
  14. Boppré M (1986) Insect pharmacophagously utilizing defensive plant chemicals (pyrrolizidine alkaloids). Naturwissenschaften 73:17–26CrossRefGoogle Scholar
  15. Brower LP, Calvert WH (1985) Foraging dynamics of bird predators on overwintering monarch butterflies in Mexico. Evolution 39:852–868CrossRefGoogle Scholar
  16. Brower LP, Fink LS (1985) A natural toxic defense system: cardenolides in butterfly versus birds. Ann N Y Acad Sci 443:171–188PubMedCrossRefGoogle Scholar
  17. Brown SG, Boettner GH, Yack JE (2007) Clicking caterpillars: acoustic aposematism in Antheraea polyphemus and other Bombycoidea. J Exp Biol 210:993–1005PubMedCrossRefGoogle Scholar
  18. Bruum H, Slabbekoorn H (2005) Acoustic communication in noise. Adv Study Behav 35:151–209CrossRefGoogle Scholar
  19. Bura VL, Fleming AJ, Yack JE (2009) What's the buzz? Ultrasonic and sonic warning signals in caterpillars of the great peacock moth (Saturnia pyri). Naturwiss 96:713–718PubMedCrossRefGoogle Scholar
  20. Bura VL, Rohwer VG, Martin PR, Yack JE (2011) Whistling in caterpillars (Amorpha juglandis, Bombycoidea): sound-producing mechanism and function. J Exp Biol 214:30–37Google Scholar
  21. Bura VL, Hnain AK, Hick JN, Yack JE (2012) Defensive sound production in the tobacco hornworm, Manduca sexta (Bombycoidea: Sphingidae). J Insect Behav 25:114–126CrossRefGoogle Scholar
  22. Candolin U (2003) The use of multiple cues in mate choice. Biol Rev 78:559–571CrossRefGoogle Scholar
  23. Carpenter GDH (1938) Audible emission of defensive froth by insects. Proc Zool Soc A 242–251Google Scholar
  24. Carpenter GDH, Ford EB (1933) Mimicry. Methuen, LondonGoogle Scholar
  25. Claridge MF (1974) Stridulation and defensive behaviour in the ground beetle, Cychrus caraboides (L.). J Entomol A 49:7–15Google Scholar
  26. Cohen JA (1985) Differences and similarities in cardenolide contents of Queen and Monarch butterflies in Florida and their ecological and evolutionary implications. J Chem Ecol 11:85–103CrossRefGoogle Scholar
  27. Conner WE, Corcoran AJ (2012) Sound strategies: The 65-million-year-old battle between bats and insects. Annu Rev Entomol 57:21–39PubMedCrossRefGoogle Scholar
  28. Corcoran AJ, Barber JR, Hristov NI, Conner WE (2011) How do tiger moths jam bat sonar? J Exp Biol 214:2416–2425PubMedCrossRefGoogle Scholar
  29. Cott HB (1940) Adaptive coloration in animals. Methuen, LondonGoogle Scholar
  30. Cuthill IC, Stevens M, Sheppard J, Maddocks T, Parraga CA, Troscianko TS (2005) Disruptive coloration and background pattern matching. Nature 434:72–74PubMedCrossRefGoogle Scholar
  31. Darwin C (1887) The life and letters of Charles Darwin: including an autobiographical chapter, edited by his son Francis Darwin. Murray, LondonCrossRefGoogle Scholar
  32. de Jong PW, Holloway GJ, Brakefield PM, Vos H (1991) Chemical defense in the ladybird beetles (Coccinellidae). II. Amount of reflex fluid, the alkaloid adaline and individual variation in defense in 2-spot ladybirds (Adalia bipunctata). Chemoecology 2:15–19CrossRefGoogle Scholar
  33. Dean J (1980) Encounters between bomardier beetles and two species of toads (Bufo americanus, Bufo marinus): speed pf prey capture does not determine success. J Comp Physiol 135:41–50CrossRefGoogle Scholar
  34. Dittrich W, Gilbert F, Green P, McGregor P, Grewcock D (1993) Imperfect mimicry: a pigeon’s perspective. Proc R Soc Lond B 251:195–200CrossRefGoogle Scholar
  35. Dunning DC (1968) Warning sounds of moths. Z Tierpsychol 25:129–138Google Scholar
  36. Edmunds M (1974) Defence in animals. Longman, HarlowGoogle Scholar
  37. Eisner T (2003) For the love of insects. Harvard University Press, CambridgeGoogle Scholar
  38. Eisner T, Grant RP (1980) Toxicity, odor aversion and “olfactory aposematism”. Science 213:416Google Scholar
  39. Eisner T, Aneshansley D, Eisner M, Rutowski R, Chong B, Meinivald J (1974) Chemical defense and sound production in Australian tenebrionid bettles (Adelium spp.). Psyche 81:189–208CrossRefGoogle Scholar
  40. Endler JA (1993) The color of light in forests and its implications. Ecol Monogr 63:1–27CrossRefGoogle Scholar
  41. Exnerová A, Landová E, Stys P, Fuchs R, Prokopová M, Cehláriková P (2003) Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus; Heteroptera). Biol J Lonn Soc 78:517–525CrossRefGoogle Scholar
  42. Fenton MB, Roeder KD (1974) The microtymbals of some Arctiidae. J Lepidopt Soc 28:205–211Google Scholar
  43. Franchina JJ, Moon C, Peters S (1997) Effects of toxin magnitude on taste aversion and taste-potentiated aversion to visual cues in chicks (Gallus domesticus). Physiol Behav 62:605–609PubMedCrossRefGoogle Scholar
  44. Fryday SL, Greig-Smith PW (1994) The effects of social learning on the food choice of the house sparrow (Passer domesticus). Behaviour 128:281–300CrossRefGoogle Scholar
  45. Gaul AT (1952) Audio mimicry: an adjunct to colour mimicry. Psyche 59:82–83CrossRefGoogle Scholar
  46. Ghirlanda S, Enquist M (2003) A century of generalization. Anim Behav 66:15–36CrossRefGoogle Scholar
  47. Gilbert FS (2005) The evolution of imperfect mimicry. In: Fellowes MDE, Holloway GJ, Rolff J (eds) Insect evolutionary ecology. CABI, Wallingford, pp 231–288Google Scholar
  48. Gittleman JL, Harvey PH (1980) Why are distasteful prey not cryptic? Nature 286:149–150CrossRefGoogle Scholar
  49. Guilford T (1994) Go-slow signaling and the problem of automimicry. J Theor Biol 170:311–316CrossRefGoogle Scholar
  50. Guilford T, Dawkins MS (1991) Receiver psychology and the evolution of animal signals. Anim Behav 42:1–14CrossRefGoogle Scholar
  51. Guilford T, Rowe C (2000) Aposematism: to be red or dead. Trends Ecol Evol 15:261–262CrossRefGoogle Scholar
  52. Guilford T, Nicol C, Rothschild M, Moore BP (1987) The biological roles of pyrazines: evidence for a warning odour function. Biol J Linn Soc 31:113–128CrossRefGoogle Scholar
  53. Halpin CG, Skelhorn J, Rowe C (2008a) Being conspicuous and defended: selective benefits for the individual. Behav Ecol 19:1012–1017CrossRefGoogle Scholar
  54. Halpin CG, Skelhorn J, Rowe C (2008b) Naïve predators and selection for rare conspicuous defended prey: the initial evolution of aposematism revisited. Anim Behav 75:771–781CrossRefGoogle Scholar
  55. Halpin CG, Skelhorn J, Rowe C (2012) The relationship between sympatric defended species depends upon predators’ discriminatory behaviour. PLoS One 7:e44895PubMedCrossRefGoogle Scholar
  56. Haskell PT (1956) tape recording of sounds emitted by the peacock butterfly. Proc R Ent Soc C 21:20–22Google Scholar
  57. Haskell PT (1961) Insect sounds. HF&G Witherby, LondonGoogle Scholar
  58. Hatle JD, Salazar BA, Whitman DW (2002) Survival advantage of sluggish individuals in aggregations of aposematic prey during encounters with ambush predators. Evol Ecol 16:415–431CrossRefGoogle Scholar
  59. Hauglund K, Snorre HB, Lampe H (2006) Responses of domestic chicks Gallus gallus domesticus to multimodal aposematic signals. Behav Ecol 17:392–398CrossRefGoogle Scholar
  60. Hebets EA, Papaj DR (2005) Complex signal function: developing a framework of testable hypotheses. Behav Ecol Sociobiol 57:197–214CrossRefGoogle Scholar
  61. Hill SA (2007) Sound generation in Mantis religiosa (Mantodea: Mantidae): stridulatory structures and acoustic signal. J Orthopt Res 16:35–49CrossRefGoogle Scholar
  62. Ingalls V (1993) Startle and habituation responses of blue jays (Cyanocitta cristata) in a laboratory simulation of anti-predator defenses of Catocala moths (Lepidoptera: Noctuidae). Behaviour 126:77–96CrossRefGoogle Scholar
  63. Järrvi T, Sillen-Tullberg D, Wiklund C (1981) The cost of being aposematic: an experimental study of predation on larvae of Papilio machaon by the great tit Parus major. Oikos 36:267–272CrossRefGoogle Scholar
  64. Jetz W, Rowe C, Guilford T (2001) Non-warning odors trigger innate color aversions—as long as they are novel. Behav Ecol 12:134–139CrossRefGoogle Scholar
  65. Johnstone RA (1996) Multiple displays in animal communication: “back-up signals” and “multiple messages”. Phil Trans R Soc Lond B 352:329–338CrossRefGoogle Scholar
  66. Jones CG, Whitman DW, Compton SJ, Silk PJ, Blum MS (1989) Reduction in diet breadth results in sequestration of plant chemicals and increases efficacy of chemical defense in a generalist grasshopper. J Chem Ecol 15:1811–1822CrossRefGoogle Scholar
  67. Kaye H, Mackintosh NJ, Rothschild M, Moore BP (1989) Odour of pyrazine potentiates an association between environmental cues and unpalatable taste. Anim Behav 37:563–568CrossRefGoogle Scholar
  68. Kelly DJ, Marples NM (2004) The effects of novel odour and colour cues on food acceptance by the zebra finch (Taeniopygia guttata). Anim Behav 68:1049–1054CrossRefGoogle Scholar
  69. Kirchner WH, Röschard J (1999) Hissing in bumblebees: an interspecific defence signal. Insect Soc 46:239–243CrossRefGoogle Scholar
  70. Laurent P, Braekman JC, Daloze D (2005) Insect chemical defence. Top Curr Chem 240:167–229Google Scholar
  71. Lindström L, Alatalo RV, Mappes J, Riipi M, Vertainen L (1999) Can aposematic signals evolve by gradual change? Nature 397:249–251CrossRefGoogle Scholar
  72. Lindström L, Rowe C, Guilford T (2001) Pyrazine odour makes visually conspicuous prey aversive. Proc R Soc Lond B 268:1–4CrossRefGoogle Scholar
  73. Mackintosh NJ (1974) The psychology of animal learning. Academic, LondonGoogle Scholar
  74. Marples NM, Roper TJ (1996) Effects of colour and smell on the response of naive chicks towards food and water. Anim Behav 51:1417–1424CrossRefGoogle Scholar
  75. Marples NM, van Veelen W, Brakefield PM (1994) The relative importance of colour, taste and smell in the protection of an aposematic insect Coccinella septempunctata. Anim Behav 48:967–974CrossRefGoogle Scholar
  76. Mason JR, Reidinger RF (1982) Observational learning of food aversions in red-winged blackbirds (Agelaius phoeniceus). Auk 99:548–554Google Scholar
  77. Masters WM (1979) Insect disturbance stridulation: its defensive role. Behav Ecol Sociobiol 5:187–200CrossRefGoogle Scholar
  78. Møller AP, Pomiankowski A (1993) Why have birds got multiple sexual ornaments? Behav Ecol Sociobiol 32:167–176Google Scholar
  79. Moore BP, Brown MV, Rothschild M (1990) Methylalkylpyrazines in aposematic insects, their hostplants and their mimics. Chemoecology 1:43–51CrossRefGoogle Scholar
  80. Nickle DA, Castner JL, Smedley SR, Attygalle AB, Meinwald J, Eisner T (1996) Pyrazine emission by a tropical katydid: an example of chemical aposematism? (Orthoptera Tettigoniidae: Copiphorinae: Vestra Stål). J Orthopt Res 5:221–223CrossRefGoogle Scholar
  81. Olofsson M, Vallin A, Jakobsson S, Wiklund C (2011) Winter predation on two species of hibernating butterflies: monitoring rodent attacks with infrared cameras. Anim Behav 81:529–534CrossRefGoogle Scholar
  82. Pasteels JM, Grégoire J-C, Rowell-Rahier M (1983) The chemical ecology of defense in arthropods. Ann Rev Ent 28:263–289CrossRefGoogle Scholar
  83. Pearson DL (1989) What is the adaptive significance of multicomponent defensive repertoires? Oikos 54:251–253CrossRefGoogle Scholar
  84. Pomini AM, Machado G, Pinto-da-Rocha R, Macías-Ordóñez R, Marsaioli AJ (2010) Lines of defense in the harvestman Hoplobunus mexicanus (Arachnid: Opiliones): aposematism, stridulation, thanatosis, and irritant chemicals. Biochem Syst Ecol 38:300–308CrossRefGoogle Scholar
  85. Poulton EB (1890) The colours of animals: their meaning and use especially considered in the case of insects. Kegan Paul, Trench, Trübner and Co, LondonGoogle Scholar
  86. Prudic KL, Skemp AK, Papaj DR (2007) Aposematic coloration, luminance contrast and the benefits of conspicuousness. Behav Ecol 18:41–46CrossRefGoogle Scholar
  87. Raab DH (1962) Statistical facilitation of simple reaction times. Trans NY Acad Sci 24:574–590CrossRefGoogle Scholar
  88. Rashed A, Khan MI, Dawson JW, Yack JE, Sherratt TN (2009) Do hoverflies (Diptera: Syrphidae) sound like the Hymenoptera they morphologically resemble? Behav Ecol 20:396–402CrossRefGoogle Scholar
  89. Ratcliffe JM, Nydam ML (2008) Multimodal warning signals for a multiple predator world. Nature 455:96–99PubMedCrossRefGoogle Scholar
  90. Ritland DB (1994) Variation in palatability of queen butterflies (Danaus gilippus) and implications regarding mimicry. Ecology 75:732–746CrossRefGoogle Scholar
  91. Roper TJ, Cook SE (1989) Responses of chicks to brightly coloured insect prey. Behaviour 110:276–293CrossRefGoogle Scholar
  92. Roper TJ, Marples NM (1997) Odour and colour as cues for taste-avoidance learning in domestic chicks. Anim Behav 53:1241–1250PubMedCrossRefGoogle Scholar
  93. Roper TJ, Redston S (1987) Conspicuousness of distasteful prey affects the strength and durability of one-trial aversive learning. Anim Behav 35:739–747CrossRefGoogle Scholar
  94. Roth LM, Eisner T (1962) Chemical defenses of arthropods. Annu Rev Entomol 7:107–136CrossRefGoogle Scholar
  95. Rothschild M (1961) Defensive odours and Mullerian mimicry among insects. Trans R Ent Soc Lond 113:101–121CrossRefGoogle Scholar
  96. Rothschild M (1965) Proc R Ent Soc Lond C 30:3Google Scholar
  97. Rothschild M, Haskell PT (1966) Stridulation of the garden tiger moth Arctia caja L. audible to the human ear. Proc R Ent Soc Lond A 41:167–170Google Scholar
  98. Rothschild M, Moore BP (1987) Pyrazines as alerting signals in toxic plants and insects. In: Labeyrie V, Fabres G, Fachaise D (eds) Insect–plants. W. Junk, DordrechtGoogle Scholar
  99. Rothschild M, Moore BP, Brown WV (1984) Pyrazines as warning odour components in the monarch butterfly, Danaus plexippus, and in moths of the genera Zygaena and Amata (Lepidoptera). Biol J Linn Soc 23:372–380CrossRefGoogle Scholar
  100. Rowe C (1998) Multicomponent signals. Unpubl. DPhil thesis, University of OxfordGoogle Scholar
  101. Rowe C (1999) Receiver psychology and the evolution of multicomponent signals. Anim Behav 58:921–931PubMedCrossRefGoogle Scholar
  102. Rowe C (2002) Sound improves visual discrimination learning in avian predators. Proc R Soc Lond B 269:1353–1357CrossRefGoogle Scholar
  103. Rowe C, Guilford T (1996) Hidden colour aversions in domestic chicks triggered by pyrazine odours of insect warning displays. Nature 383:520–522CrossRefGoogle Scholar
  104. Rowe C, Guilford T (1999) Novelty effects in a multimodal warning signal. Anim Behav 57:341–346PubMedCrossRefGoogle Scholar
  105. Rowe C, Guilford T (2000) Aposematism: to be red or dead. Trends Ecol Evol 15:261–262CrossRefGoogle Scholar
  106. Rowe C, Guilford T (2001) The evolution of multimodal warning displays. Evol Ecol 13:655–671CrossRefGoogle Scholar
  107. Rowe C, Skelhorn J (2005) Colour biases are a question of taste. Anim Behav 69:587–594CrossRefGoogle Scholar
  108. Ruxton GD, Kennedy MW (2006) Peppers and poisons: the evolutionary ecology of bad taste. J Anim Ecol 75:1224–1226PubMedCrossRefGoogle Scholar
  109. Ruxton GD, Sherratt TN, Speed MP (2004) Avoiding attack: the evolutionary ecology of crypsis, warning signals and mimicry. OUP, OxfordCrossRefGoogle Scholar
  110. Schilman PE, Lazzari CR, Manrique G (2001) Comparison of disturbance stridulations in five species of triatominae bugs. Acta Trop 79:171–178PubMedCrossRefGoogle Scholar
  111. Schlenoff DH (1985) The startle responses of blue jays (Cyanocitta cristata) to Catocala (Lepidoptera: Noctuidae) prey models. Anim Behav 33:1057–1067CrossRefGoogle Scholar
  112. Schuler W, Hesse E (1985) On the function of warning coloration: a black and yellow pattern inhibits prey-attack by naive domestic chicks. Behav Ecol Sociobiol 16:249–255CrossRefGoogle Scholar
  113. Sen-Sarma M, Fuchs S, Werber C, Tautz J (2002) Worker piping triggers hissing for coordinated colony defence in the dwarf honeybee Apis florae. Zoology 105:215–223PubMedCrossRefGoogle Scholar
  114. Sherratt TN, Speed MP, Ruxton GD (2004) Natural selection on unpalatable species imposed by state-dependent foraging behaviour. J Theor Biol 228:217–226PubMedCrossRefGoogle Scholar
  115. Siddall EC, Marples NM (2008) Better to be bimodal: the interaction of color and odor on learning and memory. Behav Ecol 19:425–432CrossRefGoogle Scholar
  116. Siddall EC, Marples NM (2011a) The effect of pyrazine odor on avoidance learning and memory in wild robins Erithacus rubecula. Curr Zool 57:208–214Google Scholar
  117. Siddall EC, Marples NM (2011b) Hear no evil: the effect of auditory warning signals on avian innate avoidance, learned avoidance and memory. Curr Zool 57:197–207Google Scholar
  118. Skelhorn J (2011) Colour biases are a question of conspecifics’ taste. Anim Behav 81:825–829CrossRefGoogle Scholar
  119. Skelhorn J, Rowe C (2006) Avian predators taste-reject aposematic prey on their level of chemical defence. Biol Lett 2:348–350PubMedCrossRefGoogle Scholar
  120. Skelhorn J, Rowe C (2007) Predators’ toxin burdens influence their strategic decisions to eat toxic prey. Curr Biol 17:1479–1483PubMedCrossRefGoogle Scholar
  121. Skelhorn J, Rowe C (2009) Distastefulness as an antipredator defence strategy. Anim Behav 78:761–766CrossRefGoogle Scholar
  122. Skelhorn J, Rowe C (2010) Birds learn to use distastefulness as a signal of toxicity. Proc R Soc Lond B 277:1729–1734CrossRefGoogle Scholar
  123. Skelhorn J, Griksaitis D, Rowe C (2008) Colour biases are more than a question of taste. Anim Behav 75:827–835CrossRefGoogle Scholar
  124. Smith SM (1975) Innate recognition of coral snake pattern by a possible avian predator. Science 187:759–760PubMedCrossRefGoogle Scholar
  125. Speed MP, Alderson NJ, Hardman C, Ruxton GD (2000) Testing Müllerian mimicry: an experiment with wild birds. Proc R Soc Lond B 267:725–731CrossRefGoogle Scholar
  126. Srygley RB (2004) The aerodynamic costs of warning signals in palatable mimetic butterflies and their distasteful models. Proc R Soc Lond B 271:589–594CrossRefGoogle Scholar
  127. Vallin A, Jakobsson S, Lind J, Wiklund C (2005) Prey survival by predator intimidation: an experimental study of peacock butterfly defense against blue tits. Proc R Soc Lond B 272:1203–1207CrossRefGoogle Scholar
  128. Wallace AR (1867) Journal of Proceeding of the Entomological Society of London meeting March 4th, 5:lxxx-lxxxi, In Transactions of the Entomological Society of London 3rd series, vol III, London 1864–1869Google Scholar
  129. Wheeler JW, Chung RH, Oh SK, Benfield EF, Neef SE (1970) Defensive secretions of cychrine beetles (Coleoptera: Carabidae). Ann Ent Soc Am 63:469–471Google Scholar

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© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  1. 1.Centre for Behaviour and Evolution, Institute of NeuroscienceNewcastle UniversityNewcastle upon Tyne,UK

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