The effects of weaponry and mating experience on the level and outcome of agonistic interactions in male field crickets, gryllus bimaculatus (orthoptera: gryllidae)

Abstract
A wide variety of factors are predicted to influence the intensity and outcome of agonistic interactions in animals, including the resource holding potential of the opponents and the nature and value of the resource over which the individuals are competing. Field crickets (Orthoptera: Gryllidae) have been used extensively as model organisms with which to study animal contests, but relatively few studies have examined the effect of mandible size or structure, or the level of contact with females on the intensity and outcome of agonistic interactions. To do so was the aim of the present thesis, using Gryllus bimaculatus as the study species. The first finding of this study was that there is a significant degree of sexual dimorphism for anterior components of the anatomy in G. bimaculatus. The mandibles, head and pronotum of male crickets were all relatively larger than those of females. This indicates that these traits may be acted upon by intrasexual selection. In many animal species that show sexual dimorphism, a trade-off in development sees enhanced weapon growth at the expense of testes size, but no such relationship was seen in this species. A comparison of the mandible structure of males that either won or lost at flaring and or wrestling showed that a relatively wider mandible span was a significant predictor of success during mandible displays. It was also found that specific components of tooth structure, namely the length of the incisor and length to distal tip, were significantly associated with victory at the jaw flaring stage. This is the first time that mandible shape has been shown to affect fight outcome in the Gryllidae, and also the first confirmed identification of a visual cue component of fighting behaviour. Despite the effectiveness of their weapons in fighting, body mass is a primary predictor of victory in combat between G. bimaculatus males, with the greater the degree of asymmetry in weight the more likely the heavier fighter will win. However, a study of fighting behaviour between asymmetrically matched opponents found that even males who were out-weighed by 40% were still likely to escalate the fight to grappling. Furthermore, males who were able to fend off their larger opponent in their first clash were significantly more likely to win their overall encounter. This hyper-aggressive response may therefore represent an adaptive mechanism to extreme odds and is worthy of further study. Female contact is known to be a significant promoter of male aggression and fighting enthusiasm, and mate guarding aggression is well documented in G. bimaculatus. A recreation of two contradictory studies, including one which concluded that mating makes males lose fights, highlighted that female contact after spermatophore transfer can overcome the loser effect and cause a male to re-engage with a previously dominant opponent. Fighting behaviour in this species is therefore highly flexible and factors affecting the outcome of contests are complex. There is much scope for further studies on this topic.