title: What makes eyespots intimidating-the importance of pairedness.
abstract: BACKGROUND
  Many butterflies possess striking structures called eyespots on their wings, and several studies have sought to understand the selective forces that have shaped their evolution. Work over the last decade has shown that a major function of eyespots is their ability to reduce predation by being intimidating to attacking predators. Two competing hypotheses seek to explain the cause of intimidation, one suggesting 'eye-mimicry' and the other their 'conspicuousness' as the reason. There is an on-going debate about which of these better explains the effectiveness of eyespots against predation. We undertook a series of indoor experiments to understand the relative importance of conspicuousness and eye-mimicry, and therefore how predator perception may have influenced the evolution of eyespots. We conducted choice tests where artificial paper models mimicking Junonia almana butterflies were presented to chickens and their preference of attack recorded.
  
  
  RESULTS
  We first established that birds avoided models with a pair of eyespots. However, contrary to previous, outdoor experiments, we found that the total area of eyespots did not affect their effectiveness. Non-eye-like, fan shaped patterns derived from eyespots were found to be just as effective as eye-like circular patterns. Furthermore, we did not find a significant effect of symmetry of patterns, again in discordance with previous work. However, across all experiments, models with a pair of patterns, symmetric or asymmetric, eyelike or non-eye-like, suffered from fewer attacks compared with other models.
  
  
  CONCLUSIONS
  The study highlights the importance of pairedness of eyespots, and supports the hypothesis that two is a biologically significant number that is important in prey-predator signalling. We discuss the implications of our results for the understanding of eyespot evolution.
authors.affiliation: School of Biology, Indian Institute of Science Education and Research Thiruvananthapuram, CET campus, Trivandrum, 695016, India. ritwika@iisertvm.ac.in.
authors.forename: Ritwika
authors.initials: R
authors.lastname: Mukherjee
authors.affiliation: School of Biology, Indian Institute of Science Education and Research Thiruvananthapuram, CET campus, Trivandrum, 695016, India. ullasa@iisertvm.ac.in.
authors.forename: Ullasa
authors.initials: U
authors.lastname: Kodandaramaiah
mesh_terms: D000818:Animals; D005075:Biological Evolution; D001717:Birds; D002080:Butterflies / Q000033:anatomy & histology* / Q000235:genetics; D005260:Female; D008297:Male; D010858:Pigmentation / Q000235:genetics; D011235:Predatory Behavior; D014921:Wings, Animal / Q000033:anatomy & histology*