Contest outcomes are usually determined by differences in resource holding potential, the social histories of the combatants, and perceptions of resource value. One understudied aspect of gaining an advantage is the residency effect. Prior occupancy of a particular place can affect the knowledge and motivation of the resident. There could be a tactical advantage in knowing the terrain or an increased willingness to fight to maintain control of a familiar area. In this study we evaluated the importance of shelter residency effects relative to size differences between rusty crayfish (Orconectes rusticus) as potential competitors for access to shelter. The intensity of any residency effects was manipulated by altering the number of shelters in the arena. Our results suggest that any residency effect is very weak in this system, and if present may often be masked by the strong and pervasive influence on contest outcome of the relative body sizes of the contestants. We also found that both shelter number and crayfish size asymmetries had strong, independent effects on levels of aggression. Dominance, but not residency status, was a factor in shelter use.

This study shows the importance of the basic spectro-temporal song parameters in territorial contests between male songbirds. Chiffchaff and willow warbler were selected as model species for their phylogenetic and ecological similarity, strong territorial behaviour, but very different singing styles. We found that bigger males of both species sing lower pitched songs and that males of both species adjust their behaviour according to song pitch of their rival simulated by playback. Thus, it seems that both species use song pitch to acoustically assess the body size of their rivals. We also show that song length and syllable rate are important features of the song, probably signalling short-term motivation to escalate territorial conflict.

Competition over mating opportunities is a conspicuous characteristic of animal behaviour. In many butterfly species the males establish territories in places advantageous for encountering females. This thesis addresses questions about how territoriality has evolved and is maintained in butterflies. The studies have been conducted using the speckled wood butterfly, Pararge aegeria, as a model species. Males of P. aegeria are found in sunspots on the forest floor (paper I-V), on the lookout for females visiting the sunspots. However, males are only found in sunspots above a certain size (paper III). This behavior is maintained by a mating success advantage, where using large sunspots instead of small sunspots as perching areas generates a higher reproductive output (paper I). The mating success asymmetry is not explained by female choice or by a female preference for large sunspots per se (paper I, V), but rather the large sunspot facilitates visual performance of perching males and improves flight pursuit and interception of females (paper III). Winners of territorial contests gain sole ownership of large sunspot territories, while losers search for a new suitable sunspot territory (paper I, II & IV) or use smaller, suboptimal sunspots as perching sites (paper II). Territorial contests between P. aegeria males are not settled due to an obvious morphological/physiological asymmetry (paper I). Rather, variation in resource value and motivational asymmetries are important for settling contests (paper IV). A majority of male-female interactions (paper V) and matings (paper I) are initiated by a perching male detecting and intercepting a flying female. Furthermore, females can affect their chances of being detected by a perching male by behaving more conspicuously (paper V). This thesis highlights the role of female behaviour, variation in resource value and motivation asymmetries to understand the evolution of territoriality in butterflies.

Social dominance relationships are formed within numerous animal species and reduce costly fights over resources. Dominant individuals often enjoy greater access to important resources such as food and mating partners, and are generally more aggressive, bold, active and explorative compared to subdominant individuals. These behavioural traits can differ among individuals, but they can also be consistent within the individual, thereby describing the individual’s personality type. However, the causal direction of the observed correlation between dominance and personality is not well studied. One possibility is that some personality types have higher chances of obtaining a dominant social position. This would suggest that personality has consequences for fitness. Another possible explanation is that possessing different social positions gives rise to consistent behavioural differences among individuals on various timescales. If social status has a lasting effect on behaviour, social status would constitute a ‘stable state’ that explains some of the diversity of personality types that has been observed in a multitude of animal species. Dominance and personality may also share underlying proximate factors. In this thesis, I investigate the relationship between social dominance and personality using male domestic fowl, Gallus gallus domesticus. The species is group-living with pronounced dominance hierarchies, and dominance increases male access to mating partners. I show that some aspects of personality, exploration, vigilance and in particular aggressiveness, increased a male’s chances of obtaining dominance (paper III, IV, V), and that aggressiveness can be even more important than body weight and ornament size (comb size, paper V) or recent experience of winning or losing (paper IV). Winning a social interaction resulted in an increase in aggressiveness, while a decrease was seen in males that experienced a loss (paper IV). By observing behaviour before and after changes in male dominance relationships, I further show that a recent (2 days earlier) change in social status induced behavioural adjustments to the new social status in activity, exploration and vigilance (paper I). By extending the time of the new social relationship to 3 weeks, I show that such behavioural changes did not continue (paper II). Finally, I show that the social environment during juvenile development had little impact on adult male competitiveness (paper V). Social interactions appear to have several short-term effects on behaviour, but did not contribute significantly to variation and long-term consistency of personality in male fowl. Instead, a male's personality, and in particular his aggressiveness, affected the outcome of dominance interactions. Overall, my studies reveal important consequences of individual differences in behaviour, and contribute to the highly sought-after empirical testing of hypotheses explaining variation in animal personality. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 5: Manuscript.</p>

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.