The Gnatocerus cornutus or Broad-horned flour beetle is a sexually dimorphic species, meaning that both sexes have different phenotypes (appearances). The males display exaggerated mandibles which are used in intrasexual selection (male-male competition). Mandible size is a sex-limited trait among Broad-horned flour beetles which only the males develop. Males with larger mandibles have a higher mating success rate and better fitness but it has been observed that females within populations of males with enlarged mandibles have a lower fitness. This negative intersexual correlation for fitness between the sexes is a sign of an existing intralocus sexual conflict (Harano et al., 2010). This type of sexual conflict arises when the same set of alleles between males and females possess different optima. An example of this is the human hip where females require a larger hip for childbirth while males do not, instead of this trait being sex-limited and expressed differently in each sex a compromise has been reached with both evolving a hip that does not provide optimum fitness for either sex (Arnqvist & Rowe, 2005). It has been noted in an experiment conducted by Harano et al. (2010) that while mandible size is a sex-limited trait females still show signs of decreased fitness.
It has been widely asserted but unsubstantiated that sex-limited trait development will resolve intralocus sexual conflict over time. In the Harano et al. (2010) experiment the claim that sex-limited trait development will resolve intralocus sexual selection is put to the test. They select six beetle populations containing: two populations selected for large male mandible size (L), two populations selected for small male mandible size (S), and two control populations with no selection on male mandible size (C). If the claim is correct then selection on mandible size shouldn’t affect female fitness in any of the populations. In the experiment the fitness of the female beetles was measured by lifetime fecundity (lifetime reproductive success, LRS) and longevity. The results (figure 1) show a strong correlation between female fitness (LRS) and male mandible size. It is apparent from these results that intralocus sexual conflict has not been resolved through the sex-limited trait development of mandibles. Harano et al. (2010) then tested explanations for the decreased female fitness in large mandible populations. The other possible LRS influences which were recorded were female survival, mass, abdomen size and egg size. Of the possible influences only abdomen size showed a connection between mandible size and female fitness (figure 2). Larger abdomen sizes were linked to higher female fitness but in populations with males possessing enlarged mandibles female and male abdomen sizes were much smaller. It is here where the intralocus sexual conflict becomes evident as the alleles for large mandibles also code for reduced abdomen sizes. Males in the need for large mandibles have evolved smaller abdomens and while females do not develop large mandibles the antagonistic selection for large mandibles has had a negative impact nonetheless.
Sex-limited trait development may resolve some instances of intralocus sexual conflict but in the case of the Broad-horned flour beetle the development of sex distinguishable traits has only moved the focus of this sexual conflict from mandible size towards abdomen size (Harano et al., 2010). For sex limited trait development to be a proper resolve for intralocus sexual conflict traits need to be genetically independent from one another, which is usually not true amongst many extant organisms. The Harano et al. (2010) experiment has successfully tested and provided evidence against sex-limited traits being a resolve for intralocus sexual selection and highlighted that in many situations the focus will shift towards a genetically dependant trait.

Figure 1. The correlation between treatment groups and female fitness: (L) Large mandibles, (C) Control group, (S) Small mandible, and (LRS) Lifetime Reproductive Success.

Figure 2. The correlation between (a) female survival and treatment group; (b) female egg size and treatment group; (c) female body mass and treatment group; and (d) female abdomen size and treatment group: (L) Large mandibles, (C) Control group, (S) Small mandible.
References
· Arnqvist G. & Rowe L. (2005) Sexual Conflict. Princeton University Press, Princeton, New Jersey.
· Harano T. Okada K. Nakayama S. Miyatake T. & Hosken D.J. (2010) Intralocus Sexual Conflict Unresolved by Sex-Limited Trait Expression. Current Biology, 20: 2036-2039.
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