Darwinian sexual selection predicts that males with the most extravagant secondary sexual traits suffer elevated mortality. Although correlative evidence has generally not borne this idea out, recent research, including a field cricket study, showed that investment in sexually selected traits is costly to survival. I investigated male survival, ornamentation (song) and mating success in a North American grylline, Gryllus pennsylvanicus, to test the generality of previous work and highlight the importance of ecology differences to resource allocation. As the calling songs of older male G. pennsylvanicus are highly attractive to females, in Chapter 2 I tested whether male age correlated with calling song and found a weak but statistically significant correlation, thus leaving open the possibility that choosy females use an age-based indicator mechanism. In Chapter 3, I tested the condition dependence of male survival and calling effort. In contrast to previous work, I found that high condition males both called more and lived longer than low condition males, although there was no trade-off between survival and calling effort. The substantial condition dependence of calling effort suggests that calling effort is under strong directional selection. In Chapter 4 I tested whether female mating preferences resulted in strong selection on male calling effort. I also tested for the condition dependence of female mating preferences. I found that female choosiness was condition-dependent, but the rank of preferred male songs (preference function) was not. Both low and high condition females preferred high calling effort over low calling effort song. In Chapter 5 I tested for evidence of nonlinear selection on male survival that might explain the nonlinear pattern of male investment in survival seen in Chapter 3 (i.e. male survival leveled-off with increasing condition). I found that socially experienced females, but not virgin and naive females, exerted linear selection on male age. I discuss these and the other results of my thesis in the context of previous work on field crickets and condition-dependent ornamentation. Finally, Appendix A reports results that confirm ancient Chinese cultural knowledge that large headed male crickets are more successful in male-male combat.

Acoustic communication in orthopterans and anurans provides a suitable model system for studying the evolutionary mechanisms of sexual selection mainly because males use acoustic signals to attract females over long distances for pair formation. Females use these signals not only to localize conspecific males but also to discriminate between potential mates. Investigations on the effect of sexual selection on acoustic signals requires an understanding of how female preferences for different features of the acoustic signal affect male mating success under ecological constraints in wild populations. The effect of female preferences on male mating success depends on the mate sampling strategy that females employ to search for potential mates. Despite its relevance, female mate sampling strategies based on male acoustic signals have rarely investigated in orthopterans and anurans, especially in the field. Considering the elaborate knowledge of the role of sensory physiology in female phonotaxis behaviour and characterization of the male acoustic signal, I used the field cricket species Plebeiogryllus guttiventris as a model system in this study. In this thesis, I first investigated the ecology of callers in wild populations. I then investigated female mate sampling strategies by incorporating relevant information on the ecology of signalers and the sensory physiology of receivers. Amount of calling activity is a strong determinant of male mating success in acoustically communicating species such as orthopterans and anurans. While many studies in crickets have investigated the determinants of calling effort, patterns of variability in male calling effort in natural choruses remain largely unexplored. I therefore investigated the spatio-temporal dynamics of acoustic chorusing behaviour in a wild population. I first studied the consistency of calling activity by quantifying variation in male calling effort across multiple nights of calling using repeatability analysis. Callers were inconsistent in their calling effort across nights and did not optimize nightly calling effort to increase their total number of nights spent calling. Next, I investigated calling site fidelity of males across multiple nights by quantifying movement of callers. Callers frequently changed their calling site across calling nights with substantial displacement but without any significant directionality. Finally, I investigated trade-offs between within-night calling effort and energetically expensive calling song features such as call intensity and chirp rate. Calling effort was not correlated with any of the calling song features, suggesting that energetically expensive song features do not constrain male calling effort. The two key features of signaling behaviour, calling effort and call intensity, which determine the duration and spatial coverage of the sexual signal, are uncorrelated and function independently Acoustic signal variation and female preference for different signal components constitute the prerequisite framework to study the mechanisms of sexual selection that shape acoustic communication. Despite several studies of acoustic communication in crickets, information on both male calling song variation in the field and female preference in the same system is lacking for most species. First, I quantified variation in the spectral, temporal and amplitudinal characteristics of the male calling song in a wild population, at two temporal scales, within and across nights, using repeatability analysis. Carrier frequency (CF) was the most repeatable call trait across nights, whereas chirp period (CP) had low repeatability. I further investigated female preferences based on song features with high and low repeatability (CF and CP respectively). Females showed no consistent preferences for CF but were more attracted towards calls with higher rates (shorter CP). I also examined the effect of signal intensity, which is known to play a critical role in female phonotaxis behaviour, on female preferences for faster calls. Females preferred louder calls over faster ones, implying a dominant role for signal intensity in female evaluation of potential mates based on acoustic signals. Call intensity was also the only signal feature that was positively correlated with male size. In the final chapter, I investigated female mate sampling strategies based on acoustic signals using both theoretical and empirical approaches. Analytical models of mate sampling have demonstrated significant differences in individual fitness returns for different sampling strategies. However these models have rarely incorporated relevant information on the ecology of signalers and the sensory physiology of receivers. I used simulation models to compare the costs and benefits of different mate sampling strategies by incorporating information on relative spacing of callers in natural choruses and the effect of signal intensity on female phonotaxis behaviour. The strategy of mating with males that were louder at the female position emerged as the optimal sampling rule in the simulations. When tested empirically in the field using callers in natural choruses, females seemed to follow the optimal strategy of mating with males that were perceived as louder at their position.

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.