Abstract

Reduced male aggression towards different phenotypes generating negative frequency-dependent intrasexual selection has been suggested as a mechanism to facilitate the invasion and maintenance of novel phenotypes in a population. To date, the best empirical evidence for the phenomenon has been provided by laboratory studies on cichlid fish with different colour polymorphisms. Here we experimentally tested the hypothesis in a natural population of Lake Malawi cichlid fish, in which males build sand-castles (bowers) to attract females during seasonal leks. We predicted that if bower shape plays an important role in male aggressive interactions, aggression among conspecific males should decrease when their bower shape is altered. Accordingly, we allocated randomly chosen bowers in a Nyassachromis cf. microcephalus lek into three treatments: control, manipulated to a different shape, and simulated manipulation. We then measured male behaviours and bower shape before and after these treatments. We found that once bower shape was altered, males were involved in significantly fewer aggressive interactions with conspecific males than before manipulation. Mating success was not affected. Our results support the idea that an extended phenotype, such as bower shape, can be important in maintaining polymorphic populations. Specifically, reduced male conspecific aggression towards males with different extended phenotypes (here, bower shapes) may cause negative frequency-dependent selection, allowing the invasion and establishment of a new phenotype (bower builder). This could help our understanding of mechanisms of diversification within populations, and in particular, the overall diversification of bower shapes within Lake Malawi cichlids.

Highlights

  • The mechanisms which lead to species divergence in secondary sexual characters are relatively poorly understood and their potential to lead to speciation has remained contentious (Arnegard & Kondrashov, 2004; Coyne & Orr, 2004)

  • Verification of experimental design We first confirmed that bowers assigned to different treatment groups were distributed throughout the lek (ANOVA, F2 = 0.336, P = 0.716), that the two Discriminant Functions (DFs) of bower shape did not differ significantly (MANOVA: F4 = 0.599, P = 0.664) and that the eight behaviour observation categories did not differ significantly (MANOVA: F14 = 0.937, p = 0.520)

  • We found that on a lek of bower building cichlid fish from Lake Malawi, males provided with a rare and altered extended phenotype were involved in less aggressive interactions with other males than males holding bowers of the normal extended phenotype

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Summary

Introduction

The mechanisms which lead to species divergence in secondary sexual characters are relatively poorly understood and their potential to lead to speciation has remained contentious (Arnegard & Kondrashov, 2004; Coyne & Orr, 2004). This would allow the stabilization of trait polymorphisms and eventual population divergence (Arnegard & Kondrashov, 2004). Negative frequency-dependent selection has been suggested as a mechanism by which sexual selection can maintain polymorphisms in populations, and eventually lead to speciation (Mikami, Kohda & Kawata, 2004; Seehausen & Schluter, 2004; Van Doorn, Dieckmann & Weissing, 2004). If individuals direct more aggression towards rivals that resemble their own phenotype than towards alternative phenotypes (Pauers et al, 2008), the rare phenotypic forms will enjoy an advantage through reduced aggression This would allow the invasion of a new phenotype into a population, which could increase in number. These processes have been recently reviewed in Qvarnstrom, Vallin & Rudh (2012) and Dijkstra & Groothuis (2011)

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