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.

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This thesis represents a comprehensive examination of the phonotactic behaviour (i.e. attraction to sound) of the female Gryllus bimaculatus under laboratory conditions. Chapter 2 is the first study to analyze the effect of substrate texture on walking performance in crickets. Substrate texture is found to play an essential role in the phonotactic responses of G. bimaculatus. Smooth substrate texture has a detrimental effect due to slipping, whereas a rough texture results in optimal walking performance due to the friction with the walking legs. Chapter 3 represents the first detailed lifetime study analysing phonotaxis in crickets. My results demonstrate that the optimal age to test phonotaxis in G. bimaculatus females is from day 7 to 24 after the final moult. I also found that selectiveness was persistent with age. These findings contradict the female choosiness hypothesis. This study is also the first to describe the effect of senescence on phonotaxis in insects, as responsiveness decreases with age. Chapter 4 compares the phonotactic behaviour of female crickets from different laboratory-bred colonies. From six tested cricket lab colonies, I found three groups statistically different from each other. Females raised under laboratory conditions at the University of Cambridge and Anglia Ruskin University were most reponsive at a frequency of 4.5 kHz, whereas females bred in Tokushima University in Japan were tuned towards a higher frequency of 5 kHz. These results suggest a degree of artificial allopatric speciation. Comparisons with crickets bred under low-quality conditions in a local pet shop demonstrate a loss of responsiveness, indicating that breeding conditions have a direct effect on phonotactic responsivity. Chapter 5 is the first study to report the presence of phonotaxis in males of G. bimaculatus. Previously it was unknown if G. bimaculatus males were able to perform phonotaxis, given that they were only recognised as endurance signal producers. In the present study, only 20% of the studied males (N=70) performed a weak phonotactic response. This finding has potential ecological implications in terms of male cricket territory establishment, and male-male interactions in the wild, which are discussed. Chapter 6 explores the song pattern recognition of the female G. bimaculatus by changing the duration of either the first, second or third pulse of the chirps. A long first pulse decreased the phonotactic response whereas phonotaxis remained strong when the third pulse was long. Chirps with three pulses of increasing duration of 5, 20 and 50 ms elicited phonotaxis, but the chirps were not attractive when played in reverse order. The data are in agreement with a mechanism in which processing of a sound pulse has an effect on the processing of the subsequent pulse, as outlined in the flow of activity in a delay-line and coincidence-detector circuit.