Abstract

Genetic polymorphism can be maintained over time by negative frequency-dependent (FD) selection induced by Rock-paper-scissors (RPS) social systems. RPS games produce cyclic dynamics, and have been suggested to exist in lizards, insects, isopods, plants, and bacteria. Sexual selection is predicted to accentuate the survival of the future progeny during negative FD survival selection. More specifically, females are predicted to select mates that produce progeny genotypes that exhibit highest survival during survival selection imposed by adult males. However, no empirical evidence demonstrates the existence of FD sexual selection with respect to fitness payoffs of genetic polymorphisms. Here we tested this prediction using the common lizard Zootoca vivipara, a species with three male color morphs (orange, white, yellow) that exhibit morph frequency cycles. In a first step we tested the congruence of the morph frequency change with the predicted change in three independent populations, differing in male color morph frequency and state of the FD morph cycle. Thereafter we ran standardized sexual selection assays in which we excluded alternative mechanisms that potentially induce negative FD selection, and we quantified inter-sexual behavior. The patterns of sexual selection and the observed behavior were in line with context-dependent female mate choice and male behavior played a minor role. Moreover, the strength of the sexual selection was within the magnitude of selection required to produce the observed 3-4-year and 6-8 year morph frequency cycles at low and high altitudes, respectively. In summary, the study provides the first experimental evidence that underpins the crucial assumption of the RPS games suggested to exist in lizards, insects, isopods, and plants; namely, that sexual selection produces negative-FD selection. This indicates that sexual selection, in our study exert by females, might be a crucial driver of the maintenance of genetic polymorphisms.

Highlights

  • Rock-paper-scissors (RPS) games can explain the maintenance of genetic polymorphisms over time and they give rise to evolutionary stable three-morph systems through negative-frequencydependent (FD) selection (Maynard Smith, 1982; Nowak, 2006; Sinervo and Calsbeek, 2006)

  • RPS games have been described in lizards (Sinervo and Lively, 1996; Sinervo et al, 2007), insects (Zhang et al, 2013), and bacteria (E. coli, Kerr et al, 2002), and it has been suggested by Sinervo and Calsbeek (2006), that they exist in isopods

  • RPS dynamics can explain how genetic polymorphisms are maintained over time (Maynard Smith, 1982; Nowak, 2006; Sinervo and Calsbeek, 2006)

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Summary

Introduction

Rock-paper-scissors (RPS) games can explain the maintenance of genetic polymorphisms over time and they give rise to evolutionary stable three-morph systems through negative-frequencydependent (FD) selection (Maynard Smith, 1982; Nowak, 2006; Sinervo and Calsbeek, 2006). RPS dynamics are characterized by fitness intransitivity whereby, each morph, when being at high frequency, is vulnerable to invasion by another morph. In a true RPS game driven by male strategies (Sinervo and Calsbeek, 2006), one rare strategy has high fitness while the other rare strategy has low fitness relative to the common strategy. RPS games have been described in lizards (Sinervo and Lively, 1996; Sinervo et al, 2007), insects (Zhang et al, 2013), and bacteria (E. coli, Kerr et al, 2002), and it has been suggested by Sinervo and Calsbeek (2006), that they exist in isopods

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