Evolutionary Ecology

, Volume 12, Issue 6, pp 667–679 | Cite as

Plant defence signals and Batesian mimicry

  • Magnus Augner
  • Elizabeth a. Bernays


In a game theory context, we investigated conditions for an evolutionarily stable equilibrium of defended, signalling plants, and plants mimicking these signals – that is, conditions for a stable mimicry complex. We modelled this in three steps. First, we analysed conditions for selection for defended, signalling plants, in a population of undefended plants. Second, we analysed conditions for when mimicking plants can invade a population of defended, signalling plants, leading to a stable equilibrium between the two strategies. Third, we analysed how sampling of signalling plants by herbivores affects the equilibrium between the strategies. The predictions show that mimicry of plant defence signals may be common, and even imperfect mimics could invade a population of defended, signalling plants. Whether the latter prediction holds or not depends on how herbivores generalize over signals, and on the length of their ’avoidance sequence'. The length of the avoidance sequence is the number of signalling plants that a herbivore avoids to attack, after attacking a defended plant. If herbivores always sample signalling plants, then mimicry cannot evolve, whereas if herbivores have a long avoidance sequence, this may allow selection even for imperfect mimics.

foraging game theory herbivory mimicry plant defences plant signals 


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  1. Augner, M. (1994) Should a plant always signal its defence against herbivores? Oikos 70, 322–332.Google Scholar
  2. Augner, M. (1995) Low nutritive quality as a plant defense: Effects of herbivore mediated interactions. Evol. Ecol. 9, 605–616.Google Scholar
  3. Augner, M., Fagerström, T. and Tuomi, J. (1991) Competition, defense and games between plants. Behav. Ecol. Sociobiol. 29, 231–234.Google Scholar
  4. Avery, M.L. (1985) Application of mimicry theory to bird damage control. J. Wildl. Manage. 49, 1116–1121.Google Scholar
  5. Bannister, P. (1989) Nitrogen concentration and mimicry in some New Zealand mistletoes. Oecologia 79, 128–132.Google Scholar
  6. Barlow, B.A. and Wiens, D. (1977) Host-parasite resemblance in Australian mistletoes: The case for cryptic mimicry. Evolution 31, 69–84.Google Scholar
  7. Barrett, S.C.H. (1987) Mimicry in plants. Sci. Am. 257, 68–75.Google Scholar
  8. Bates, H.W. (1862) Contributions to an insect fauna of the Amazon Valley. Lepidoptera: Heliconidae. Trans. Linn. Soc. Lond. 23, 495–566.Google Scholar
  9. Brower, J.vz. (1958) Experimental studies of mimicry in some North American butterflies. Part I. The Monarch, Danaus plexippus, and the Viceroy, Limenitis archippus archippus. Evolution 12, 32–47.Google Scholar
  10. Brower, J.vz. (1960) Experimental studies of mimicry. IV. The reactions of starlings to different proportions of models and mimics. Am. Nat. 877, 271–282.Google Scholar
  11. Carpenter, G.D.H. (1949) Pseudacraea eurytus (L.) (Lep. Nymphalidae): A study of polymorphic mimic in various degrees of speciation. Trans. R. Ent. Soc. Lond. 100, 71–133.Google Scholar
  12. Dukas, R. and Clark, C.W. (1995) Sustained vigilance and animal performance. Anim. Behav. 49, 1259–1267.Google Scholar
  13. Ehleringer, J.R., Ullmann, I., Lange, O.L., Cowan, I.R., Schultze, E.-D. and Ziegler, H. (1986) Mistletoes: A hypothesis concerning morphological and chemical avoidance of herbivory. Oecologia 70, 234–237.Google Scholar
  14. Eisner, T. and Grant, R.P. (1981) Toxicity, odor aversion and ‘olfactory aposematism’. Science 213, 476.PubMedGoogle Scholar
  15. Faegri, K. and van der Pijl, L. (1979) The Principles of Pollination Ecology. Pergamon Press, New York.Google Scholar
  16. Ford, E.B. (1936) The genetics of Papilio dardanus (Lep.). Trans. R. Ent. Soc. Lond. 85, 435–466.Google Scholar
  17. Fowler, M.E. (1983) Plant poisoning in free-living wild animals: A review. J. Wildl. Dis. 19, 34–43.PubMedGoogle Scholar
  18. Getty, T. (1985) Discriminability and the sigmoid functional response: How optimal foragers could stabilize model-mimic complexes. Am. Nat. 125, 239–256.Google Scholar
  19. Glendinning, J.I. (1993) Preference and aversion for deterrent chemicals in two species of Peromyscus mouse. Physiol. Behav. 54, 141–150.PubMedGoogle Scholar
  20. Hay, M.E. (1986) Associational plant defenses and the maintenance of species diversity: Turning competitors into accomplices. Am. Nat. 128, 617–641.Google Scholar
  21. Hay, M.E. and Fenical, W. (1992) Chemical mediation of seaweed-herbivore interactions. In Plant-Animal Interactions in the Marine Benthos (D.M. John, S.J. Hawkins and J.H. Price, eds), pp. 319–337. Systematics Association Special Volume No. 46. Clarendon Press, Oxford.Google Scholar
  22. Huheey, J.E. (1964) Studies of warning coloration and mimicry. IV. A mathematical model of model-mimic frequencies. Ecology 45, 185–188.Google Scholar
  23. Huheey, J.E. (1980) Studies in warning coloration and mimicry. VIII. Further evidence for a frequency-dependent model of predation. J. Herpet. 14, 223–230.Google Scholar
  24. Huheey, J.E. (1988) Mathematical models of mimicry. Am. Nat. 131, S22–S41.Google Scholar
  25. Launchbaugh, K.L. and Provenza, F.D. (1993) Can plants practice mimicry to avoid grazing by mammalian herbivores? Oikos 66, 501–504.Google Scholar
  26. Launchbaugh, K.L., Provenza, F.D. and Burritt, E.A. (1993) How herbivores track variable environments: Response to variability of phytotoxins. J. Chem. Ecol. 19, 1047–1056.Google Scholar
  27. Leimar, O., Enquist, M. and Sillén-Tullberg, B. (1986) Evolutionary stability of aposematic coloration and prey unprofitability: A theoretical analysis. Am. Nat. 128, 469–490.Google Scholar
  28. Luedeman, J.K., McMorris, F.R. and Warner, D.D. (1981) Predators encountering a model-mimic system with alternative prey. Am. Nat. 117, 1040–1048.Google Scholar
  29. Malcolm, S.B. (1990) Mimicry: Status of classical evolutionary paradigm. Trends Ecol. Evol. 5, 57–62.Google Scholar
  30. Maynard Smith, J. (1982) Evolution and the Theory of Games. Cambridge University Press, Cambridge.Google Scholar
  31. Mead, R.J., Oliver, A.J., King, D.R. and Kubach, P.H. (1985) The co-evolutionary role of fluoroacetate in plant-animal interactions in Australia. Oikos 44, 55–60.Google Scholar
  32. McNamara, J. and Houston, A. (1980) The application of statistical decision theory to animal behaviour. J. Theor. Biol. 85, 673–690.PubMedGoogle Scholar
  33. Nonacs, P. (1985) Foraging in a dynamic mimicry complex. Am. Nat. 126, 165–180.Google Scholar
  34. Owen, D.F. (1974) Exploring mimetic diversity in West African forest butterflies. Oikos 25, 227–237.Google Scholar
  35. Owen, R.E. and Owen, A.R.G. (1984) Mathematical paradigms for mimicry: Recurrent sampling. J. Theor. Biol. 109, 217–247.Google Scholar
  36. Pough, F.H. (1988) Mimicry of vertebrates: Are the rules different? Am. Nat. 131, S67–S102.Google Scholar
  37. Provenza, F.D., Pfister, J.A. and Cheney, C.D. (1992) Mechanisms of learning in diet selection with reference to phytotoxicosis in herbivores. J. Range. Manage. 45, 36–45.Google Scholar
  38. Rhoades, D.F. (1979) Evolution of plant chemical defense against herbivores. In Herbivores: Their Interaction with Secondary Plant Metabolites (G.A. Rosenthal and D.H. Janzen, eds), pp. 3–54. Academic Press, Orlando, FL.Google Scholar
  39. Smith, S.M. (1977) Coral-snake pattern recognition and stimulus generalisation by naive great kiskadees (Aces: Tyrannidae). Nature 265, 535–536.Google Scholar
  40. Spence, K.W. (1937) The differential response in animals to stimuli varying within a single dimension. Psychol. Rev. 44, 430–444.Google Scholar
  41. Stephens, D.W. (1987) On economically tracking a variable environment. Theor. Pop. Biol. 32, 15–25.Google Scholar
  42. Till-Bottraud, I. and Gouyon, P.-H. (1992) Intra-versus interplant Batesian mimicry? A model on cyano-genesis and herbivory in clonal plants. Am. Nat. 139, 509–520.Google Scholar
  43. Tuomi, J.M. and Augner, M. (1993) Synergistic selection of unpalatability in plants. Evolution 47, 668–672.Google Scholar
  44. Vet, L.E.M. and Dicke, M. (1992) Ecology of infochemical use by natural enemies in a tritrophic context. Ann. Rev. Entomol. 37, 141–172.Google Scholar
  45. Wiens, D. (1978) Mimicry in plants. Evol. Biol. 11, 365–403.Google Scholar
  46. Wiklund, C. (1975) The evolutionary relationship between adult oviposition preferences and larval host plant range in Papilio machaon L. Oecologia 18, 185–197.Google Scholar
  47. Wrubel, R.P. and Bernays, E.A. (1990) Effects of ingestion of some non-host plant secondary compounds on larvae of Manduca sexta. Ent. Exp. Appl. 54, 125–130.Google Scholar
  48. Zalucki, M.P. and Brower, L.P. (1992) Survival of first instar larvae of Danaus plexippus (Lepidoptera: Danainae) in relation to cardiac glycoside and latex context of Asclepias humistrata (Asclepiadaceae). Chemoecology 3, 81–93.Google Scholar

Copyright information

© Kluwer Academic Publishers 1998

Authors and Affiliations

  • Magnus Augner
    • 1
  • Elizabeth a. Bernays
    • 1
  1. 1.Department of EntomologyUniversity of ArizonaTucsonUSA

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