The protective value of conspicuous signals is not impaired by shape, size, or position asymmetry

Various conspicuous signals in nature promote initial and learned avoidance by predators. It is widely thought that such signals are most effective when highly symmetrical in features such as size and shape, supported by recent laboratory experiments with domestic chicks and artificial prey. However, no study has investigated the effect of asymmetry on conspicuous signals in a natural setting, where viewing distances, angles, predator species, and light conditions vary and where predators encounter prey sequentially rather than simultaneously. We undertook 2 field experiments with artificial gray-scale prey, marked with a pair of white markings presented to wild avian predators, to test the effect of asymmetry on the survival value of conspicuous signals in the field. Experiment 1 had treatments with symmetrical spots or with spots asymmetrical in area between 5 and 50%. All marked treatments survived better than unmarked controls, but there was no benefit of being symmetrical. Experiment 2 tested the effect of possessing markings asymmetrical for shape or position and any additive effect of these 2 features. Again, symmetry conferred no benefit and targets with markings asymmetrical for position and/or shape survived equally well as those with symmetrical arrangements. These findings indicate that asymmetry in warning signals may not be costly to prey in nature or be of less importance compared with other features of the signal, such as color and overall size. Copyright 2009, Oxford University Press.

[1]  J. Lawless Statistical Models and Methods for Lifetime Data , 2002 .

[2]  Martin Stevens,et al.  Predator perception and the interrelation between different forms of protective coloration , 2007, Proceedings of the Royal Society B: Biological Sciences.

[3]  Innes C Cuthill,et al.  The predation costs of symmetrical cryptic coloration , 2006, Proceedings of the Royal Society B: Biological Sciences.

[4]  G. Beauchamp,et al.  Time for some a priori thinking about post hoc testing , 2008 .

[5]  P. Brakefield Structure of a character and the evolution of butterfly eyespot patterns. , 2001, The Journal of experimental zoology.

[6]  Alicia M. Frame,et al.  A test of receiver perceptual performance: European starlings' ability to detect asymmetry in a naturalistic trait , 2008, Animal Behaviour.

[7]  H. B. Cott,et al.  Adaptive Coloration in Animals , 1940 .

[8]  A. Møller,et al.  Asymmetry, Developmental Stability, and Evolution , 1998 .

[9]  S. Merilaita,et al.  Fearful symmetry: pattern size and asymmetry affects aposematic signal efficacy , 1999, Evolutionary Ecology.

[10]  I. Cuthill,et al.  Disruptive coloration, crypsis and edge detection in early visual processing , 2006, Proceedings of the Royal Society B: Biological Sciences.

[11]  Asymmetry in size, shape, and color impairs the protective value of conspicuous color patterns , 2004 .

[12]  J. Swaddle Limits to length asymmetry detection in starlings: implications for biological signalling , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[13]  Gil G. Rosenthal,et al.  Symmetry without fear , 1994, Nature.

[14]  V. French,et al.  The relationship between eyespot shape and wing shape in the butterfly Bicyclus anynana: A genetic and morphometrical approach , 1997 .

[15]  M. Stevens,et al.  The anti-predator function of ‘eyespots’ on camouflaged and conspicuous prey , 2008, Behavioral Ecology and Sociobiology.

[16]  T. Sherratt,et al.  Empirical tests of the role of disruptive coloration in reducing detectability , 2007, Proceedings of the Royal Society B: Biological Sciences.

[17]  S. Merilaita,et al.  Fearful symmetry? Intra-individual comparisons of asymmetry in cryptic vs. signalling colour patterns in butterflies , 2003, Evolutionary Ecology.

[18]  J. Swaddle,et al.  European starlings are capable of discriminating subtle size asymmetries in paired stimuli. , 2007, Journal of the experimental analysis of behavior.

[19]  M. Stevens,et al.  Conspicuousness, not eye mimicry, makes "eyespots" effective antipredator signals , 2008 .

[20]  I. Cuthill,et al.  The effects of pattern symmetry on detection of disruptive and background matching coloration , 2006 .

[21]  D. Kelly,et al.  Neophobia and Dietary Conservatism:Two Distinct Processes? , 1999, Evolutionary Ecology.

[22]  Daniel Osorio,et al.  Symmetry detection by categorization of spatial phase, a model , 1996, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[23]  I. Cuthill,et al.  Field experiments on the effectiveness of ‘eyespots’ as predator deterrents , 2007, Animal Behaviour.

[24]  Jerald F. Lawless,et al.  Statistical Models and Methods for Lifetime Data: Lawless/Statistical , 2002 .

[25]  E. Hancock,et al.  Asymmetrical colour and wing-folding in Tithrone roseipennis (Saussure 1870) a neotropical praying mantis (Mantodea Hymenopodidae) , 1999 .

[26]  Nina Stobbe,et al.  Disruptive coloration provides camouflage independent of background matching , 2006, Proceedings of the Royal Society B: Biological Sciences.

[27]  V. French,et al.  The relationship between eyespot shape and wing shape in the butterfly , 1997 .

[28]  D. Rubin,et al.  Contrasts and Effect Sizes in Behavioral Research , 1999 .

[29]  D. Cox Regression Models and Life-Tables , 1972 .

[30]  J. Endler,et al.  The complex business of survival by aposematism. , 2005, Trends in ecology & evolution.

[31]  J. Klein,et al.  Survival Analysis: Techniques for Censored and Truncated Data , 1997 .

[32]  Douglas A. Ruff,et al.  STARLINGS HAVE DIFFICULTY IN DETECTING DOT SYMMETRY: IMPLICATIONS FOR STUDYING FLUCTUATING ASYMMETRY , 2004 .

[33]  S. Merilaita,et al.  Great Tits (parus Major) Searching for Artificial Prey: Implications for Cryptic Coloration and Symmetry , 2005 .

[34]  M. Huynen,et al.  Disruptive coloration and background pattern matching , 2005, Nature.